Annals

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Volume XIV

1927

With Forty-one Plates and One Hundred and

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TABLE OF CONTENTS

PAGE

Revision of the Genus Townsendia . . . Esther L. Larsen 1-46

Studies on South American Labiatae, III
Carl C. Epling 47-86

A Monograph of the Genus Pholiota. .L. 0. Overholts 87-210

A Monograph of the Section Oreocarya

of Cryptantha Edwin B. Payson 211-358

Cotton Wilt : A Pathological and Physiolog-
ical Investigation David C. Neal 359-424

Species of Cercospora on Smilax in the

United States L. 0. Overholts 425-432

General Index to Volume XIV 433-436

Annals

of the

Missouri Botanical Garden

Vol. 14 FEBRUARY, 1927 No. 1

A REVISION OF THE GENUS TOWNSENDIA 1

ESTHER LOUISE LARSEN

Jessie R. Barr Research Fellow in the Henry Shaw School of Botany

of Washington University

History of the Genus

The genus Townsendia was first described by William Jackson
Hooker in his 'Flora Boreali- Americana ' in 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 Townsendia, recognized only one
species, Townsendia sericea, which he described in detail and
carefully illustrated.

Thomas Nuttall, 3 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 "on the Black Hills (or eastern chain
of the Rocky Mountains) near the banks of the Platte." During
the next four decades T. Parryi Eaton, 4 T. condensata Parry, 6

1 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 Sci

Wash

8 Hook. Fl. Bor. Am. 2: 16. 1834.

»Nutt. Trans. Am. Phil. Soc. N. S. 7: 304. 1841.

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

•Parry, Am. Nat. 8: 213. 1874.

Issued June 8, 1927

Ann. Mo. Bot. Gard., Vol. 14, 1927

(1)

2 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

T. scapigera Eaton, 6 T. Rothrockii Gray, 7 and T. Wilcoxiana
Wood 8 were described by individual workers, thus making a total
of ten known species. In 1880 Dr. Asa Gray 9 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 10 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 a 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 Townsendia exscapa
(Richards.) Porter, 11 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. A 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., 12 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.

'Gray, Wheeler Kept. 6: 148. 1878.

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

•Gray, Proc. Am. Acad. 16: 82. 1880.

"Gray, Syn. PL N. Am., ed. 2, 1*: 166. 1886 and 1888.

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

"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. The 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 Townsendia develop rather
slender tap-roots. The roots of some of the more caespitose
species, however, become very coarse and woody and more or less
branched. The 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, erect, scapiform stems, while T.
grandifiora, T. texensis, and T. strigosa have numerous ascending
branched stems. The 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. The broader leaves are attenuated
at the base into a petiole, while the narrower ones are only ob-
scurely petiolate or sessile. The range in the leaf size is from
0.5 to 6 cm. in length and from 0.2 to 1.0 cm. in width. Entire
leaf margins prevail throughout the genus. The surface is usually
pubescent, but sometimes it is glabrate, as in T. glabella. In T.
spathulata, on the other hand, the leaves are villose-lanate. How-
ever, a strigose pubescence of closely appressed hairs as in T.
eximia and T. strigosa is most prevalent in the genus.

[Vol. 14

4 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Inflorescence. — The heads resemble those of the closely related
genus Aster. They 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 involucre 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. A lacerate-ciliate margin prevails throughout the
genus. Furthermore, in most species the involucral bracts also
have a membranaceous margin. The 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. The pappus of the
ray-flower is somewhat shorter, or often reduced to a crown of
short squamellae. The pappus in T. formosa in both ray- and
disk-flowers is scarcely more than a vestige of the squamellate
crown. Townsendia eximia develops a coroniform pappus of
coalescent rigid squamellae sometimes bearing two or more
prolonged awns. The condition in T. 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. The variability in the length of the pappus in certain
species, particularly in T. sericea, has been emphasized by Gray,
Meehan, 1 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

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

1927]

LARSEN — REVISION OF THE GENUS TOWNSENDIA 5

but

both rav- 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 T. strigosa and T. grandiflora
as " infertile or neuter." A 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

within the snecies and is important in the classificat

of the

Geographical Distribution

The genus Townsendia 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. T. mexicana is endemic to Mexico. The
only other representative of the group occurring in that country
is T. 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, T. leptotes occurs only in the
Middle and South Park Region, T. glabella in southwestern Colo-
rado, while T. Rothrockii extends from the north-central part of
the state to the Uncompahgre Mountains. Two other species

6

[Vol. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Plate 1. Geographical distribution of the genus Tovmsendia.

• • •

X

X

III— III

• •*

X

ill IM

7 owtcsendia exsrapa.

Townsendia Rothrockii.
T ointserulid eximia.

Tontiserulia jnontana.

Tovmsendia sericea.

Tovmsendia stngosa.

Toumsetulia mexicana.

Tmvfi8€ndia scapigera.

t

Tovmsendia sjxithulaia.

o

+

+ + +

Tovmsendia glabella.

nvnxendia florifer
Tovmsendia grandifior
Tovmsendia incana*
Tovmsendia Parryi,
^Tovmsendia WcUsonL
Tovmsendia arizonica.
Tovmsendia texensis.
Tovmsendia fornwsa.
Tovmsendia leptotes.

1927 J

LARSEN — REVISION OF THE GENUS TOWNSENDIA 7

which seem to be restricted to rather local areas are T. formosa of
southwestern New Mexico and adjacent Arizona and T. texensis of
northwestern Texas. The difference between the mountain
species T. sericea and the plains species T. exscapa is very well
brought out by material collected in Colorado. The 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. T. 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 T. 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;

8 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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

Taxonomy

Townsendia Hook. Fl. Bor. Am. 2: 16. 1834; DC. Prodr. 7:
273. 1838; Nutt. Trans. Am. Phil. Soc. N. S. 7: 304. 1841;
loir. & Gray, Fl. N. Am. 2: 185. 1842; Benth. & Hook. Gen. PI.
2: 208. 1873; Gray, Proc. Am. Acad. 16: 82. 1880; Syn. Fl.
N. Am. P: 166. 1884, and ed. 2, 166. 1886 and 1888; Coulter,
Man. Rocky Mt. Region, 156. 1S85 ; Engl. &Prantl, Nat. Pflanzen-
fam. 4 5 : 161. 1890; Nelson in Coulter & Nelson, Man. Cent.
Rocky Mis. 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. Hay-flowers nu-
merous, in a single series, pistillate, fertile, rays linear, 2S-
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, barbcllate-scabrous bristles as long as the corolla; that of
the ray-llowers 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: T. sericea Hook. Fl. Bor. Am. 2: 16, pi. 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

A. Bracts of the involucre attenuate-acuminate.
B. Stems erect, simple.

C. Pappus plurisetose 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 hut with two stout awns S. 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.

F. Involucral bracts scarious-margined.

G. Pappus of ray-flower a crown of short squamellae. . . .5. T. texensis

GG. Pappus of ray-flower plurisetose 6. T. glabella

FF. Involucral bracts not scarious-margined 7. T. Rothrockii

EE. Leaves persistently pubescent.

H. Achenes glabrous or glabrate 8. T. montana

HH. Achenes persistently pubescent.
J. Hairs on achenes bidentate.

K. Plants cinereous with short hirsute pubescence.
L. Involucral bracts linear-lanceolate.

M. Leaves all obovate-spathulate 9. T. Watsoni

MM. Leaves of stem mostly linear-spathulate 10. T. florifer

LL. Involurral bracts broadly lanceolate 11. T. scapigera

KK. Plants densely subsericeous, villous 12. T. spathulata

JJ. Hairs on achenes glochidiate-tipped.

N. Plants with branched ascending stems.

O. Involucral bracts in two series, equal 13. T. mexicana

00. Involucral bracts in several series, unequal 14. T. strigosa

NN. Plants depressed, caespitose.
P. Densely canescent, pubescent.

Q. Leaves 1.5-3 cm. long, narrowly spathulate

to oblanceolate 15. T. incana

QQ. Leaves 1-1.5 cm. long, obovate-spathulate. .. .16. T. arizorrica

PP. Sparsely hirsute pubescent 17. T. leptotes

NNN. Plants strictly acaulescent.

R. Leaves narrowly oblanceolate, somewhat glabrate,

distinctly flattened 18. T. exscapa

RR. Leaves linear-lanceolate, densely pubescent, subterete./£. T. sericea

1. T. Parryi Eaton, Am. Nat. 8: 212. 1874; Gray, Proc. Am.
Acad. 16: 82. 1880; Syn. Fl. N. Am. I 2 : 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. FL
Rocky Mts. 874. 1917, and ed. 2, 874. 1922.

10 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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 cm.
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 (F); canyon, Helena, July, 1892, Alton (F,
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 801 (F); Gallatin Co.,
May, 1888, Tweedy 228 (F); mountain meadows, alt. 1537 m.,
Bozeman, June 1, Livingston, June 8, 1906, Blankinship 801a
(F, M, R); on a clayey gravelly slope in the foothills ten miles
east of Monida, Madison Co., June 18, 1899, A. & E. 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 (M);
1888, Kelsey (M, No. 783933) ; July, 1894, Mrs. Moore (M).

Yellowstone National Park: Electric Peak, alt. 2923 m.,
July 26, 1902, Smith 16 (F); Mt. Washburn, July 20, 1902,
Smith (F, No. 121846); Mammoth Hot Springs, June 15, 1902,
Mearns (F, No. 121848); subalpine, alt. 2923 m., Aug., 1884,
Tweedy (F, 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,

Eikenberry 59 (F).

1927]

LARSEN — REVISION OF THE GENUS TOWNSENDIA 11

Wyoming: Wind River Mts., alt. 2764 m., 1873, Parry 144
(F, M, co-type) ; Gros Ventres Fork, alt. 2000 m., June 10, 1860,
Hay den (M); slopes at timber-line, Wyoming Range, 15 miles
west of Merna, Sublette Co., July 18, 1922, E. & L. Pay son
(M, R); sage-brush slopes 20 miles west of Big Piney, Sublette
Co., July 10, 1922, E. & L. Payson 2632 (M, R) ; Gros Ventres
Fork, alt. 2400 m., June 5, 1860, Hay den (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,
Lincoln Co., alt. 3169 m., Aug. 2, 1923, E. & L. Payson

(M).
Idaho: 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).

Canada: "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).

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

2. T. formosa Greene, Lean. 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 ace. 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 a stout
upright monocephalous stem about 25 cm. 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 cm. 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; involucre 2-3-seriate; bracts with
broad membranaceous margins, minutely lacerate-ciliate, those

12

[Vol. 14

BOTANICAL

of the outer series broadly ovate?, those of the inner series linear-
lanceolate, distinctly attenuate-acuminate; achenes glabrous.

Plate 2; pi. 5, fig. 19-24.

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

Specimens examined:

New Mbxico: 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
1484 (M, co-type); Sacramento Mts., July 28, 1899, Wooton (R).

Arizona: White Mountains, Aug. 0-15, 1903, Griffiths 5340
(M); Bonita Creek, White Mts., July 23, 1912, Goodding 1235
(R); Thorn] sons Ranch, Black River, White Mts., July 13, 1910,

Goodding 501 (R).

3. T. eximia Gray, .Mem. Am. Acad. (PL Fendl.) N.S. 4: 70.
1849; Walp. Ann. Bot, Syst. 2: 822. 1851-1852; Syn. Fl. N.
Am. I 2 : 107. 1884, and ed. 2, 169. 1886 and 18S8; Proc. Am.
Acad. 16: 83. 1880; Coulter, Man. Rocky Mt. Region, 156.
1885; Coulter k Nelson, Man. Cent. Rocky Mts. 508. 1909;
Wooton 6l 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 cm. high; leaves spathulate or the upper

solitary

ceolate, 2-5 cm. lt>ng, nearly glabrate; heads terminal,
involucre 3-4-seriate, 1-1.5 cm. broad, 2-3 cm. in di-
ameter; bracts ovate-lanceolate and somewhat cuspidate-acumin-
ate with a narrow membranaceous lacerate-ciliate margin; ray-
flowers about 35-40, fertile, blue or purple, with a 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, type of T. Vreelandii) .

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. & 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 U12 (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. (PI. Fendl.) N.S. 4: 70. 1849; Proc. Am. Acad. 16: 83.
1880; Syn. Fl. N. Am. I 2 : 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. S. Nat. Herb. 19: 692. 1915;
Rydb. Fl. Rocky Mts. 873. 1917, and ed. 2, 873. 1922.

Caulescent, divaricately branched from the base, 5-20 cm.
high; leaves linear, sublanceolate, acute, nearly glabrous, 2-5
cm. long, 0.3-1 cm. 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.

14 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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

Specimens examined:

South Dakota: 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 2188 (F).

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

Wyoming: 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 253 (M).

Colorado: Douglas Co., 1892, Walker (F, 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 3188 (F) ; 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 (F, No. 290193) ; Colorado Territory, 39^0° 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 (F, No. 208980);
near Golden City, 1870, Greene (F, No. 15363); Fremont Co.,
near Canyon City, 1873, Greene (F, No. 15364) ; 39-40° 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 OF THE GENUS TOWNSENDIA 15

flora, 39-41° lat., 1862, Hall & Harbour 289 (F, M); eastern
Colorado, Carleton (F, No. 353119) ; southern Colorado, Brandegee
(F, No. 204740) ; Manitou, Aug. 11, 1884, Letterman 83 (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 157 (M).

5. T. texensis Larsen, n. sp. 1

Herbaceous perennial; the caudex giving rise to ascending
branched stems 6-30 cm. high, leafy throughout; leaves oblance-
olate, 1-5 cm. long, 0.2-0.8 cm. broad, apiculate, narrowed at the
base into a petiole, sparsely pubescent with closely appressed
hairs, occasionally glabrate; heads usually solitary, terminal,
sessile or short-pedunculate, 1.5-3 cm. in diameter including the
ray; involucre 4-5-seriate, 1-1.5 cm. 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 (M, Nos. 121021, type, 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

1 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, utrinque dense strigoso-pilosis, rarius glabratis; capitulis
plerumque solitariis, terminalibus, sessilibus vel brevi-pedunculatis, radio incluso
1.5-3 cm. in diametro; involucris campanulatis, 4-5-seriatis, 1-1.5 cm. in diametro;
bracteis involucri oblanceolatis, acutis ad apices roseis, marginibus membranaceis
lacerato-ciliatisque ; floribus femineis 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 "rocky bluffs of the Red River," Texas, Aug. 13,
1900, H. Eggert (Mo. Bot. Gard. Herb., No. 121021, type).

(Vol. 14

16 ANNALS OF THE MISSOURI BOTANICAL GARDEN

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

6. T. glabella Gray, Proc. Am. Acad. 16: 86. 1880; Syn. FL
N. Am. I 2 : 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. Bakeri Greene, Pittonia 4: 157. 1900.

Subacauiescent ; caudex thick, woody, bearing tufted leaves;
leaves thick, pilose when young, soon becoming glabrous, linear-
spathulate, 2-4 cm. long, 0.2-0.5 cm. broad, gradually narrowed
into a slender petiole ; heads solitary, on naked peduncles 2-5 cm.
long; heads 1.5-2.5 cm. 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 (M,
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 (F, M, R); Los
Pinos (Bayfield), May 16, 1899, Baner 727 (F, M, R, type of T.
Bakeri); mature yellow pine forest, Piedra, June 31, 1924, Hazel
M. Schmoll 1348 (R) ; open places between pines, Piedra, June 21,

1914, Hazel M. Schmoll 1212 (R).

7. T. Rothrockii Gray, ace. to Rothrock in Wheeler Rept. 6:
148, L 7. 1878; Gray, Proc. Am. Acad. 16: 85. 1880; Syn. Fl.
N. Am. I 2 : 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
875 (F, type) ; South Park, Aug., 1873, Rothrock (F, 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. S.
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 cm. long, 0.3-0.5 cm. 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 cm. 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).

[Vol. 14

18 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Wyoming: Piney Mt., 25 miles west of Big Piney, Sublette
Co., Summit, July 12, 1922, E. & L. Payson 2694 (M, R); 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, R); Sheep Mt.
(Ferry Peak), Snake River Range, near Alpine, Lincoln Co., July
11, 1923, Payson & Armstrong 3474 (M, R); in vicinity of Green
River Lakes, Sublette Co., alt. 3169 m., Aug. 5, 1925, E. & L.

Payson 45 4% (M).

Idaho : 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).

Utah: canyon above Tropic, alt. 2154 m., May 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 T. mon-
tana); loose stony soil, Uintah Mts., Dyer Mine, July 3, 1902,
Goodding 1238 (R, M, co-type of T. 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. Fl.
N. Am. I 2 : 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 N. Am. I 2 : 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 cm. in diameter including the rays;
involucre 2-seriate; bracts oblong-lanceolate, margins membran-
aceous, lacerate-ciliate ; ray-flowers with a reduced pappus of
unequal capillary bristles shorter than the diameter of the achene ;

1927)

LABSEN — 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:

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

Nevada: 1891, A. J, Jones (M).

Oregon: sandy soil near Vale, May, 1896, Leiberg 2067 (M);
common on hills, in the region of Malheur River, June 19, 1898,
Cusick 1935 (F, M).

10. T. florifer (Hook.) Gray, Proc. Am. Acad. 16: 84. 1880; Syn
Fl. N. Am. I 2 : 167. 1884, and ed. 2, 167. 1886 and 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. Bor. Am. 2: 20. 1834.
Aplopappus florifer Hook. & Arn. Bot. Beechey Voy. 351.

1841, excl. var. p.
Stenotus florifer Torr. & Gray, Fl. N. Am. 2: 238. 1842, excl.

var. p.

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.

20 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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, II ay den (M).

Idaho: ditch banks along fields, Challis, Custer Co., alt. 1662

m., July 15, 1916, Macbride & Payson 3225 (M, R); 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 8095 (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).

Utah: Joseph City, Sevier Co., alt. 1692 m., May 13, 1899,
Jones 6379 (M); Marysvale, alt. 1846 m., May 31, 1894, Jones

(M, F); rim of Great Salt Lake Desert, May 6, 1889, H.
Engelmann (M) ; sage-brush slopes, Milford, June 5, 1902, Good-
ding 1 046 (R) .

Washington: Ritzville, Adams Co., alt. 480 m., June 6, 1893,
Sandberg & Leiberg 169 (F, M); Craigs Ferry, Kittitas Co., July
15, 1903, Cotton 1361 (M); Columbia River opposite Umatilla,
Apr. 20, 1882, Howell (F, 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 (M).

Oregon: near Lexington, Morrow Co., alt. 420 m., May 7,
1894, Leiberg 84 (F, M); near Umatilla, May 1, 1882, Howell (F,
No. 396898); on open plains, Cline Falls, Crook Co., May
1905, Nelson 815 (M, R) ; dry banks of Deschutes River five miles
below Bend, July 30, 1920, Peck 9708 (M) ; stony hills west of
Silver Creek (and common westward), June 28, 1901, Cusick 2616
(F, M, R).

11. T. scapigera Eaton, Bot. King's Exp. 145. L 17. 1871;
Gray, Proc. Am. Acad. 16: 84. 1880; Syn. Fl. N. Am. I 2 : 168.
1884, and ed. 2, 168. 1886 and 1888; Rydb. Fl. Rocky Mts. 874.

19271

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. Torr. & Gray, Fl. N. 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. I 2 : 168.
1886, and ed. 2, 168. 1886 and 1888.

T. scapigera var. ambigua Gray, Proc. Am. Acad. 16: 84. 1880;
Syn. Fl. N. Am. I 2 : 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 cm. 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:

Wyoming: 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.).

Utah: Rabbit Valley, alt. 2092 m., Aug. 6, 1875, Ward 523
(U. 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, type of T. scapigera var. caulescens) ; eastern Nevada, 1883,
Meehan (Phil.).

California: Buffalo Ravine near Surprise Valley, Apr. 1879,

Lemmon 29 (M).

22 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

12. T. spathulata Nutt. Trans. Am. Phil. Soc. N.S. 7: 305.
1841; Torr. & Gray, Fl. N. Am. 2: 186. 1842; Eaton, Am. Nat.
8: 213. 1874; Gray, Proc. Am. Acad. 16: 86. 1880; Syn. Fl. N.
Am. I 2 : 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. N. Am. I 2 : 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. N. Am. I 2 : 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 cm. 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, E. C. Smith (F).

Wyoming : high alpine ridge between the valleys of the Stinking
Water and the Yellowstone, 1873, Parry 1J$ (F, M) ; northwestern
Wyoming, 1873, Parry 145 (F, G, M, type of T. Parryi var.
alpina); high alpine peak, Owl Creek Range, July, 1874, /. D.
Putnam (G, type of T. condensata); "Black Hills of Platte,"
Nuttall (Phil.); shale flats, Bush Ranch, June 10, 1910, Nelson
7054 (R).

1927 J

LARSEN — REVISION OF 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 cm. 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. N. Am. I 2 : 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 cm. broad; heads usually solitary on terminal pe-
duncles, 1-2 cm. 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:

Mexico:

Coahuila: Saltillo, March 22, 1877, Gregg 327 (M, type);
Saltillo, alt. 1650 m., June 5, 1909, Arsene 8387 (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).

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

[Vol. 14

24 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Aug. 6, 1896, Pringle 6578 (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. N. Am. 2: 186. 1842; Gray, Mem. Am. Acad.
N.S. (PI. Fendl.) 4: 70. 1849, in part; Syn. Fl. N. Am. I 2 : 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. S. 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. N.
Am. I 2 : 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. S. 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 cm. long, 0.3-0.6 cm. 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 8205 (M, R) ; Church Buttes, Fort
Bridger, July, 1873, Porter (M, Phil.); without definite locality,
Parry (Phil); Green River, May 30, 1897, Nelson 3081 (M, R).

1927J

LARSEN — REVISION OF THE GENUS TOWNSENDIA 25

Colorado: "R. Mts. Platte," Nuttall (Phil., type); Salida,
June 19, 1898, Baker, EarU & Tracy 1016 (M); Texas Cr., Fre-
mont Co., 1874, Brandegee 951 (M) ; Arkansas River near Puncha
Pass, Sept. 24, 1878, Jones 767 (M) ; Huerfano, Aug., 1867, Parry
98 (M, R); dry slopes, alt. 1662 m., Paradox, Montrose Co.,
June 17, 1912, Walker 98 (M) ; adobe plains of San Juan Valley,
July, 1875, Brandegee (M) ; southern Colorado, Brandegee (Phil.) ;
sands of Huerfano Cr., Sept., 1875, Brandegee (M) ; McElmo Cr.,
June 3, 1892, Eastwood (F, No. 82217); alkaline hillsides, Natur-
ita, 1662 m., May 19, 1924, Payson 821 (F, M, R); Arkansas
Valley, Sept., 1873, Wolf 517 (F).

New Mexico: near Espanola, Santa Fe Co., alt. 1723 m.,
May 17, 1897, Heller & Heller 8547 (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., ait.
1184 m., Apr. 19, 1907, Wooton & Standby 8287 (M); sand hills
near Mesilla ; May 4, 1906, Standley (M) ; gravelly hillsides, Santa
Fe, May-July, 1847, Fendler 850 (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 851 (Phil., M); Aztec, May 4, 1899, Baker 728 (F, M, R);
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 528 (M); sandy soil, Beaver Cr.,
Sept., 1903, Purpus 8800 (M); Smart 92 (F); Voth 7 (F).

15. T. incana Nutt. Trans. Am. Phil. Soc. N.S. 7: 305. 1841;
Torr. & Gray, Fl. N. Am. 2: 155. 1842; Walp. Rep. 2: 575.
1843; Gray, Syn. Fl. N. Am. I 2 : 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.

[Vol. 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 cm. high; leaves spathulate, sometimes apiculate,
petiolate, 2-4 cm. long and 0.2-0.5 cm. broad, the uppermost
clustered at the base of the heads and seldom surpassing them in
length; heads 1-2.5 cm. in diameter including the rays, usually
sessile; involucre 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 Mexico, west into eastern Arizona and Utah.

Specimens examined:

Wyoming: " Black Hills of the Platte," Nuttall (Phil., type);
on a stony flat, Granger, Uinta Co., June 14, 1899, A. & E. Nelson
5403 (M, R); Granger, June 10, 1898, A. Nelson 4622 (M, R);
deep hot sands, Alcova, 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 (F, 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, R).

Utah: stony slopes, Thompson Springs, alt. 1630 m., May-
Oct., 1899, Purpus 6765 (M); Thompson Springs, May 7, 1891,
Jones (Pomona, No. 39664, type of T. 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.

19271

LARSEN — REVISION OF THE GENUS TOWNSENDIA 27

Chepeta Well, alt. 1540 m., May 23, 1908, Jones (Pomona, No

, type of T. incana var. prolixa).
Arizona: near Oraibi, 1900, Voth 19 (F); Hackbeny, May 26

1884, Jones A5 16 (F,R).

16. T. arizonica Gray, Proc. Am. Acad. 16: 85. 1880; Syn.
Fl. N. Am. 1«: 169. 1884, and ed. 2, 169. 1886 and 1888 ; Wooton
& Standley, Contr. U. S. Nat. Herb. 19: 692. 1915; Rydb. Fi.

Rocky Mts. 875. 1917, and ed. 2, 875. 1922; Tidestrom
Contr. U. S. Nat. Herb. 25: 554. 1925.

T

Jones. Zoe 2: 248. 1891.

Depressed subacaulescent and multicipital, branching from a
perennial root, forming a loose pnlvinate tuft 3-4 cm. high, mi-
nutely sericeous-canescent ; leaves short, obovate-spathulate, 2-3
cm. long, 0.2-0.5 cm. 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:

Colorado: Naturita, June 1, 1917, Payson 989 (M, R); rocky
hillside, Naturita, alt. 1670 m., Apr. 21, 1914, Payson 21*2 (F,
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, type of T. 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.

T. sericea var. leptotes Gray, Proc. Am. Acad. 16: 85. 1880;

28

|Vol. 14

GARDEN

Coulter, Man. Rocky Mt. Region, 157. 1885; Gray, Syn. Fl.
N. Am. I 2 : 169. 1884, and ed. 2, 1886 and 1888

T. kpotes 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 cm. long, 0.1-0.2 cm.

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:

Colorado: Middle Park, coll. of 1864, Parry (G, type; M,
co-type, No. 121020); "Estes Park," coll. of 1864, Parry (F,
No. 209717); Kremmling, Grand Co., June 22, 1907, Osterhout
8487 (R); Kremmling, Grand Co., May 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, in part,
excluding synonym T. 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. S.

, Nat. Herb
mensana.

excluding T. sericea and

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: 273. 1838, in part; Walp. Rep. 2: 575 and 957. 1843, in part,
as to Aster exscapus; Torr. & Gray, Fl. N. Am. 1: 185, 1842, in
part, as to Aster exscapus and plant of Richardson: Grav. Proc.

1927]

LARSEN — REVISION OF THE GENUS TOWNSENDIA 29

Am. Acad. 16: 85, 1880, in part, as to Aster exscapus; Syn. Fl. N.
Am. I 2 : 168. 1884, and ed. 2, 168. 1886 and 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. N. Am., ed. 2, V:
168. 1886 and 1888; Coulter, Man. Rocky Mt. Region, 157.
1885; Rydb. Fl. Rocky Mts. 875. 1917, and ed. 2, 875. 1922.

T. sericea p papposa Gray, Mem. Am. Acad. N. S. (PI. Fendl.)
4: 70. 1849; Proc. Am. Acad. 16: 84. 1880; Syn. Fl. N. Am. I 2 :
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 cm. long and 0.2-0.5 cm. broad, somewhat
apiculate, sparsely pubescent with fine appressed hairs, somewhat
glabrate; heads large, 2-3 cm. high and 3-5 cm. 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;
aehenes 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:

South Dakota: clay-hills, near Hot Springs, May 10, 1924,
Mcintosh 21 (R); hills near Hot Springs, June 2, 1924, Mcintosh

166 (R).

Kansas: Trego Co., Rich 718 (R, M,iso-type 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, Apr. 5, 1925,
Wilcox (F, M); hillside, Alva, Apr. 17, 1913, Stevens 216 (M).

30 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

Texas: infrequent, rocky slopes, alpine, March 22, 1919,
Hanson (M) ; rare on high mountains, Limpia, March, 1914, Allen
86 (M); calcareous bluffs, Falls Creek, Hood Co., Apr, 1884,
Reverchon 1583 (F, M).

Colorado: near foothills, Fort Collins, Apr. 19, 1898, Crandall
8132 (F, R) ; plains, Colorado Springs, May 4, 1878, Jones 25 (F) ;
Cheyenne Canyon, May 4, 1891, Smith (M, No. 121177); Canyon
City, Apr. 1875, Brandegee (M); Rocky Mountain FL, 39-41
lat., 1862, Hall & Harbour 290 (F, No. 456674; M, No. 121176);
Los Pinos, May, 1899, Baker 730 (F, M, R); 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 (M, co-type of T. sericea $ pap-
posa); near Silver City, March 29, 1889, Greene (F).

Arizona: Flagstaff, May-Oct., 1902, Purpus 4 (M); 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 <Sc
Palmer 365 (M) ; Fort Whipple, Apr. 20, 1865, Coues & Palmer
815 (M).

Canada: at Carlton House, Richardson (Kew, type); sand
hills, Aweme, Manitoba, May 28, 1900, Griddle (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, pi 119. 1834, in
part, excluding synonym; DC. Prodr. 7: 273. 1838, in part;
Nutt. Trans. Am. Phil. Soc. N. S. 7: 304. 1841; Walp. Rep. 2:
575 and 957. 1843, in part, excluding Aster exscapus; Torr. &
Gray, Fl. N. Am. 2: 185. 1842, excluding Aster exscapus and
the plant of Richardson; Gray, Proc. Am. Acad. 16: 85. 1880,
in part; Syn. Fl. N. Am. I 2 : 168. 1884, and ed. 2, 168. 1886 and
1888, in part, as to plant from " Rocky Mountains in lat. 54°."

T. mensana Jones, Contr. Western Bot. 13. d. 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 T. sericea; Nelson in 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.

d 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; pi. 7, fig. 32-34.

Distribution: Rocky Mountains, Alberta to Colorado and

Utah; also in the Black Hills of South Dakota.

Specimens examined:

South Dakota: dry hillside near Pringle, alt. 1508 m., Apr. 19,

1909, Murdoch 3510 (F).

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

121170).

Wyoming: Laramie, May 8, 1897, A. Nelson 2862 (F) ; Laramie
Hills, Apr. 28, 1896, A. Nelson 1883 (F); Laramie plains, Apr. 12,
1894, A. Nelson 7 (F, M); Laramie, Albany Co., May, 1899,
A. Nelson 7055 (M, R); March 8, 1860, Hayden (M, 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 (R); Rocky
Mts., 39-41° lat., 1862, Hall & Harbour 290 (F, Nos. 314634 and
17721); Larimer Co., March 30, 1896, Baker 1250 (F, M); West

32

[Vol. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Cliff, Custer Co., Apr. 1888, Cockerell (F, No. 352961); steep
slope of "The 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, Vestal 368 (M) ; infrequent,
upper mesas near Boulder, alt. 1477 m., March 19, 1921,
Hanson cl60 (M); South Park, Aug., 1873, Rothrock (F, No.
304922 in part) .

Utah: Theodore, Benches of the Uintas, alt. 2308 m., May 14,
1908, Jones (Pomona, No. 40603, type of T. mensana).

Canada: Rocky Mountains, Drummond (Kew, type; 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 Wrightii Gray, Bot. Mex. Bound. Surv. p. 78.

1859

Aster Wrightii Gray, PI. Wright. Part II, p. 75

1853

List of Exsiccatae

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

Aiton, G. B. (1).

Alcott, W. P. (3); (14).

Allen, Miss E. A. 36 (18).

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

Arsene, Bro. G. 3387 (13).

Baker, C. F. (4); 727 (6); 728 (14); 105, Cockerell, T. D. A. (19).

Brumback, Miss F. M., and Davis, Miss
C. A. 178 (4).

Burglehaus, F. H. (1).
Carleton, M. H. (4).

Clokey, I. W. 2810 (4); 4338 (19).

729 (15); 730 (18); 1250 (19).

Cotton, J. S. 1361 (10).

Baker, C. F., Earle, F. S., and Tracy, Coues, E., and Palmer, E. 523 (14); 315,

S. M. 548 (6); 1016 (14).
Bethel, E. (19).

365 (18).
Cowdry (19).

Bethel, E., Willey, F. S., and Clokey, I. Crandall, C. S. (4); 3132 (18).

W. 4340 (6).

Criddle, M. 900 (18).

Blankinship, J. W. 301, 301a (1); 147 Curtis, C. (18).

(19).
Brandegee, T. S. (3); 487 (4); (10); 951
(14); (18).

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

(9); 2616 (10).
Dawson, G. 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).

Ellis, Miss C. C. 56 (3).

Engelmann, G. (3).

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

Fendler, A. S5S (3); 157, 583 (4); 351

(11); 350, 351 (14); 349 (18).
Flodman, J. H. 828 (1).

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

Mulford, Miss I. (10).

Murdoch, J. 3510, 4503 (19).

Nelson, A. 385, 7373, 8312 (4); 815 (10);

4622 (15); 7, 1883, 2862, 7055 (19);

7054 (12); 3031 (14).

Nelson, A., and E. 5425 (1); 5403 (15).
Nelson, A., and Macbride, J. F. 1129

(10).
Nelson, E. 815 (10).
Newton, G. W. (19).

Goodding, L. N. 1288 (8); 166 (15); 1235, Norris, H. W. (18).

561 (2); 1046 (10).
Gordon, A. L. 42 (18).
Greene, E. L. (4); (18).
Gregg, J. 827 (13).
Griffiths, D. 4361 (18); 5840 (2).
Hall, E. (4).

Nuttall, T. (12); (14); (15).
Osterhout, G. E. 6116 (16); 5221, 8487
(17); (18).

Palmer, E. J. 14170, 14586 (5).
Palmer, Ed. 252, 499, (13); 204 (16);
(18).

Hall, E., and Harbour, J. P. 289 (4); Parry, C. C. 144 (1); (4); 14%, 145 (12);

290 (18); 290 (19)

95, 94, (14); (17); 35 (19).

Hanson, H. C. cl59 (4); (18); cl60 (19). Patterson, H. N. 49 (4).

Hapeman, H. (19).

Hayden, F. V. (1); (4); (19); (10).

Payson, E. B. 1160 (6); 821 (14); 989,
242 (16); 326 (18).

Heller, A. A., & E.G. 37«ff (3); 3547 (14). Payson, E. B., and L. B. 1979, 2026,

Howell, T. J. (10).

Johnston, E. L. 253, 41 4b, 492 (4); 253a

(18).
Jones, A. J. (9).
Jones, M. E. 4157 (3); 284 (4); (8); (9); Peck, M. E. 9708 (10).

5323, 6379 (10); (11); 767 (14); 4516 Porter, T. C. (4); (14).

2682, 2762, 4642, 8693 (1); 4542, 1986,
2078, 2694, 4542 (8).
Payson, E. B., and Armstrong, G. M.
8693 (1); 8474, 8706 (8); 3205 (14).

(15); 1794 (16); 25 (18); (19).

Kelsey, F. D. (1).
Kern, R. H. (11); (14).
Kirkwood, J. E. 110 (13).
Leiberg, J. B. 2067 (9); 84 (10).
Lemmon, J. G. 29 (11).
Letterman, G. W. 83 (4).
Lloyd, C. G. 110 (13).
Macbride, J. F. 90, 1017 (10).

Pringle, C. G. 6573, 7580, 9967 (13).
Purpus, C. A. 582 (7); 1345 (13); 8300

(14); 6765 (15); 4 (18).
Putnam, J. D. (12).
Ramaley, F. 654 (19).
Redfield, J. H. 478 (4).
Reverchon, J. 3320 (5); 1533 (18).
Rich, J. A. 718 (18).
Richardson, John (18).

Macbride, J. F., and Payson, E. B. 8095, Rothrock, J. T. 875 (7); (19).

8225 (10).
MacDougal, D. T. 31 (18).
Macoun, J. (1); (18); (19).
Mcintosh, A. C. 21, 166 (18).
Marshall, W. F. 8188 (4).
Mearns, E. A. (1); (8).
Meehan, T. (11).
MeU and Knopf (18).

Robbins, W. W. 8189 (3).

Rusby, H. H. 660 (16).

Rydberg, P. A., and Bessey, E. A. 5132

(1); 5181 (8).
Sandberg, J. H, and Leiberg, J. B. 169

(10).
Schmoll, Hazel M. 1348, 1212 (6).
Shear, C. L. 2 (18).

Metcalfe, O. B. 418, 1434 (2); 15, 1510 Smart, D. 92 (14).

(14).

Smith. B. H. (6).

[Vol. 14, 1927]

34

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Smith, E. C. 16 (1); (12); (18).
Spread borough, W. (18).

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

Spreadborough,W.,andMacoun, J. (19). Walker, Mrs. S. B. (4).

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); 804 (4).
Vestal, A. A. 868 (19).
Visher, S. S. 2188 (4).
Von Schrenk, H. (11).
Voth, H. R. 7 (14); 19 (15).
Vreeland, F. K. 689 (3).

Ward, L. F. 92 (9); 523 (11); 176 (15).

Watson, S. 519 (11).

Webber, H. J. (4).

Wilcox, T. E. (18).

Williams, T. A. (4).

Wolf, J. 517 (14); 416 (19).

Wooton, E. O. (2).

Wooton, E. O. and Standley, 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.

Aplopappus „ . . . 19

florifer 19

florifer var. 21

Aster Wrightii 32

Aster

28

ex sea-pus 28

Erigeron 19

florifer 19

Stenotus

19

florifer 19

florifer var. @ 21

Townsendia 8

alpigena 17

alpina 22

ambigua 21

arizonica 27

arizonica X incana 27

Bakeri 16

condensata 22

dejecta 17

eximia 12

exscapa 28

exscapa 31

exscapa Wilcoxiana 29

Fendleri 24

florifer 19

florifer var. communis 19

formosa 11

Frernontii 26

16

glabella

grandiflora 13

incana 25

incana var. ambigua 26

incana var. prolixa 26

intermedia 29

lepotes 28

leptotes 27

mensana 30

mexicana 23

montana 17

Parryi 9

Parryi var. alpina 22

pinetorum. 11

Rothrockii 16

scapigera 20

scapigera var. ambigua 21

scapigera var. caulescens 21

sericea 30

sericea 28

sericea var. leptotes 27

sericea & papposa 29

spathulata 22

strigosa 24

strigosa Eaton 18

strigosa Gray 19

texensis 15

Vreelandii 12

Watsoni 18

Wilcoxiana 29

Wrightii 32

(Vol. 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. Bot. Gaud., Vol. 14, 1927

Plate 2

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LA RSEN— REVISION OF THE GENUS TOWNSENDIA

(Vol. 14, 19271

38 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 3

Toumsendia texensis Larsen

Northwestern Texas

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

Ann. Mo. Bot. Gard., Vol. 14, 1927

Plate 3

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LARSEN— REVISION OF THE GENUS TOWNSENDIA

40

[Vol. 14, 1927]

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 4

Townsendia

Fig. 1. Style-branches of the disk-flower, gre*

Fig. 2. Style-branches of the disk-flower, X 5.

Fig. 3. Corolla of the disk-flowei

Fig. 4. A seta of the disk-flower.

snlarged

showing

Fig. 5. Disk-flower, X 5.
Fig. 6. Glochidiate- tippe«

niarged

enlarged

Fig. 7. Style-branches of the ray-flower, X 5.
Fig. 8. Style-branches of the ray-flower, great
Fig. 9. A seta of the ray-flower, greatly enlarged.
Fig. 10. Achene of the ray-flower with ]
Fig. 11. Corolla of the ray-flower, X 5.
Fig. 12. Photos

enlarged

attached

Philadelphi

Academy

Ann. Mo. Bot. Gaud., Vol. 14. 1927

Plate 4

v.

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[Vol. 14, 1927]

42 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 5

T. eximia Gray.
Fig. 13. Ray-flower, X 2]/ 2

showing

larged

Fig. 16. Pappus of the disk-flower, greatly enlarged.

Fig. 17. Achene of the disk-flower with the corolla and pa

Fig. 18. Glochidiate-tipped hair.

T. formosa Greene.
Fig. 19. Ray-flower, X 2]/ 2 .

Fig. 20. Style-branches of the ray-flower, greatly enlarged.

Fig. 21. Disk-flower, X 2Y 2 .

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

Fig. 23. Stylo-branches of the disk-flower, greatly enlarged.

Fig. 24. A seta of the pappus, greatly enlarged.

T. florifer Gray.

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

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

Ann. Mo. Bot. Gard., Vol. 14, 1927

Plate 5

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LARSEN— REVISION OF THE GENUS TOWNSENDIA

IVol. 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. Hot. Gaud., Vol. 14, 1927

Plate 6

Ex Herbario Mn i Britannia

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LARSEX— REVISION OF THE GENUS TOWXSENDIA

[Vol. 14, 1927|

46 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 7

Toumsendia exscapa (Richards.) Porter

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

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

Ann. Mo. Hot. Gaud., Vol. 14, 1927

Plate 7

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33

LARSEN— REVISION OF THE GENUS TOWNSENDIA

STUDIES ON SOUTH AMERICAN LABIATAE. Ill 1

Synopsis of the Genus Satureia

CARL CLAWSON EPLING

Instructor in Botany, University of California, at Los Angeles

Lackland

)f Washingt

Satureia (Tourn.) L.

Gardoquia Ruiz et Pavon, Prodr. 86, pi. 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 pi. 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 longiore, trifido, lobo medio majore et
subpatente; staminibus 4, supra tubi medium saepius sitis,
didymis, posticis paulo brevioribus, filamentis glabris, connectivo
crasso thecas saepius separante, his 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 a
certain diversity of habit and flower character, the species of

1 Issued June 8, 1927

Ann. Mo. Bot. Gard., Vol. 13, 1927

(47)

(Vol. 14

48 ANNALS OF THE MISSOURI BOTANICAL GARDEN

which, however, form a continuous and interlocking series, so
that subgeneric lines are difficult to distinguish. As concluded by
Briquet, 1 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 a
Micromeria non nisi longitudine corollarum, limbique lobis sub-
erectis, distinctum," 2 while Micromeria (including Xenopoma) 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 multifiora R. & P.

Conspectus Specierum

A. Flores in axillis solitarii, pedicellis calyces subaequantibus vel longioribus

elatis, bracteolis absentibus /. S. Brownei

AA. Flores 3-6 et ultra in axillis vel solitarii, pedicellis quam calycibus

brevioribus, bracteolis praesentibus.

B. Herbae fruticulique humiles, altitudine 5-20 cm., floribus in axillis

solitariis.

C. Bracteola ad pedicellorum medium sita; folia obovata, margine

integra.

D. Calyces 13-venis, dentibus duobus anticis quam posticis sub-

duplo longioribus, erectis 33. S. pusilla

DD. Calyces 15-venis, dentibus subaequalibus, subconniventibus.

32. S. Darwinii

CC. Bracteola ad pedicellorum basim sita; folia ovata vel rotundato-

ovata, margine leniter crenata 31. S. nubigena

BB. Suffrutices vel frutices, altitudine 30-200 cm., floribus 1-6 et ultra in

axillis.

C. Calyces 1.5-3.5 mm. loneri. corollae tnho inr.lnon wl nouln ovcortn

rarius quam calyce duplo longiore.

5-2

EE. Folia puberal
DD. Calyces 3-3.5 mm. longi, corollae tubo breviter exserto.

axillaris

E. Folia ovata vel rotundato-ovata, leniter crenata 34. S. vana

EE. Folia lanceolata, oblanceolata vel ovata, subintegra.

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

1927|

EPLING — SOUTH AMERICAN LABIATAE. Ill 49

F. Folia lanceolata, acuta; flores saepius tres in axiilis.

36. S. simulans

FF. Folia oblanceolata, oblonga vel ovata, obtusa; flores saepius

solitarii.

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

vel elliptica, 3-5 mm. longa 38. S. brevicalyx

GG. Calyces tubulosi; corollae 9 mm. longae; folia oblance-
olata vel oblonga, .5-1.5 cm. longa 35. S. boliviana

CC. Calyces 4.5-10 mm. (3.5-4 mm. in S. rigidvla et breviflora), saepius

6-8 mm. longi, corollae tubo saepius longe exserto quam calycibus
duplo triplove longiore.
D. Folia 2-5 cm. longa, sat tenuia, ovata vel elliptica, crenato-

serrata; flores 3-3.5 cm. longi.

E. Flores pedicellis gracilibus calyces subaequantibus elevati; caly-
ces glabri 2. S. mvltifiora

EE. Flores pedicellis quam calycibus brevioribus; calyces extus

villosi 3. S. elliptica

DD. Folia raro 3 cm. longa, saepius .5-1.5 cm. vel breviora, forma

diversiore saepius integra; flores saepius 2-2.5 cm. longi.
E. Folia obovata vel oblanceolata, omnino obtusa; calycis dentes

acuminati (acuti in S. guamanie nsis) etiam subaristati.
F. Folia rhomboideo-obovata, vel subrotunda.

G. Folia rhomboideo-obovata, subtus puberula; flores fere

2 cm. longi 6. S. Matthewsii

GG. Folia rotunda, subtus parce villosa; flores vix 6 mm. longi.

8. S. breviflora

FF. Folia oblanceolata.

G. Flores in axiilis solitarii 7. S. guamaniensis

GG. Flores 3-7 in axiilis 5. S. Gilliesii

EE. Folia elliptico-lanceolata, ovata, vel linearia, raro obtusa;

calycis dentes saepius acuti, raro subaristati (S. Lindeniana,
rigidvla, glabrata).
F. Folia 1-2.5 cm. longa, acuta (frequenter obtusa in S. tomentosa)

pagina superiore subglabra, inferiore incana (sericea in
S. acutifolia).
G. Calyces vix 5 mm. longi, tubo extus villosissimo . .12. S. discolor

GG. Calyces 6-8 mm. longi, tubo extus appresso-villoso vel

puberulo.
H. Folia ovata, crenata, subtus incano- tomentosa. .9. S. tomentosa
HH. Folia elliptico-lanceolata, subintegra.

J. Pagina superiore sericea, inferiore tomentosa; margo

revoluta 13. S. acutifolia

J J. Pagina superiore glabra, inferiore tomentella; margo

plana 15. S. Pavoniana

FF. Folia 2-15 mm. longa utrinque glabra (tomentella in S. An-

drei) saepius subnitida, venis prominulis, ascendentibus,
rectis, parallelis; margo leniter vel nihil revoluta.
G. Folia 5-15 mm. longa.
H. Folia glabra.
J. Calycis dentes subaristati, 2 mm. longi 17. S. glabrata

50 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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

18. S. taxifolia

KK. Folia utrinque obtusa, internodiis subaequilonga.

16. S. pallida

HH. Folia praecipue subtus puberula 19. S. Andrei

GG. Folia 2-5 mm. longa.

H. Folia in basi angustata, haud cordata.

J. Margo integra; calycis dentes erecti 20. S. striata

J J. Margo serratula; calycis dentes paten tes 21. S. plicatvla

HH. Folia in basi cordata.

J. Corolla 18-20 mm. longa; calycis dentes tres postices

omnino connati 23. S. connata

J J. Corolla 20-25 mm. longa; calycis dentes tres postices

fere ad medium liberi 22. S. Jamesoni

FFF. Folia 2-6 mm. longa, pubescentia, saepe sericea (pagina

superiore glabra in S. Lindeniana), venis obscuris; margo
saepius valde revoluta.
G. Calycis dentes duo antici subulati, fere aristati.
H. Dentes antici 3 mm. longi, posticos superantes.

29. S. Lindeniana

HH. Dentes antici 1.5 mm. longi, posticos aequantes.

26. S. rigidula

GG. Calycis dentes duo antici anguste lanceolati, nullomodo

aristati, posticos vix aequantes vel breviores.
H. Folia linearia, utrinque sericeo-villosa; calyces 6 mm.

longi 27. S. sericea

HH. Folia ovata, pagina superiore glabrata vel puberula;

calyces 5 mm. longi.
J. Foliorum margines valde revolutae; pagina inferiore

tomentosa 28. S. revoluta

JJ. Foliorum margines revolutae; pagina inferiore pu-
berula, venis prominulis 2J+. S. argentea

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

Micromeria sect. Hesperothymus Benth. Lab. Gen. et Sp. 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. Ill 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. 3a, 300. 1897.

Herba humilis, caule prostrato, repente, ramis filiformibus,
ascendentibus, glabris vel tenuiter pilosis, quadratis, angulis
acutis, submarginatis ; f oliis 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-
crenata 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
calyce 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 ma j ore, 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 corolla subexserto; nuculis .8-1 mm. longis,

oblongis, atris.

Subsp. eubrownei nom. nov.

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

Ramulis foliisque fere glabris; petiolis laminas aequantibus vel
parte dimidia brevioribus; f oliis nee sessilibus; florum pedicellis
saepius calycibus aequilongis vel paulo brevioribus.

Specimens examined:

Venezuela: Galipan, 1250 m., June 7, 1885, Jahn 201 (known
as "Poleo") (US) 1 ; Paramo de la Sal, 3000 m., Menda, 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 Botanical Garden; NY, New York Botanical Garden; UC, University
of California; US, United States National Herbarium.

52

[Vov. 14

BOTANICAL

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., Merida, Nov. 14, 1921, Jahn 786,738 (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 Jab6n, 3000-3200 m., Oct. 2, 1910, Jahn
41 (US); Paramo de Pinango, 2600 m., March, 17, 1915, Jahn
410 (US).

Colombia: Sabana de Bogota, Bro. Ariste-J oseph A 903 (US);
Cuesta de Tocota, 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 Caldas, 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. 11 41 (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., "a 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,
Purac6, 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) .

19271

EPLING SOUTH AMERICAN LABIATAE. Ill 53

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

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

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

Frutices f ruticulive ramulis saepius ascendentibus, brevioribus ;
foliis saepe fasciculatis, forma diversioribus, maximam partem sub-
integris, glabris vel pubescentibus ; floribus 1-6 in f oliorum 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. brevifiora and 8. tomentosa or S. boliviano and
S. Gilliesii, and in floral characters, as in S. rigidula, brevifiora,
argentea, boliviano and vana. The closest alliance between the
two sections appears to lie between 8. boliviano and members of
the subsection Obovatae, or between S. brevifiora, tomentosa and

vana.

A

Foliis 2-5 cm. longis, sat tenuibus, ovalibus vel ellipticis, glabris
vel subglabris, crenatis; floribus 3-15 in cymulis in f oliorum
axillis, corollis 3-3.5 cm. loneris.

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 cm. longis, 1-2 cm. latis,
lanceolatis vel ovatis, apice obtusis, in basi rotundatis, margine

[Vol. 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 secondares .3-1 cm. longis; calycibus
membranaceis, 8 mm. longis, in basi paulo angustatis, extus
glabris, f auce 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
circa 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:

Chili: unknown collector 245 (NY); 1828-34, Gay 148 (NY);
Gay (GH) ; Chanco, Reed (GH) ; Valdivia, Ufergebusch 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)

Panguipulli, July 1924, Bro. Claude- J oseph

(US)

3. Satureia elliptica Briq. in Engler u. Prantl, Nat. Pflanzen-
fam., ed. 1, IV. Abt. 3a, 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. Misc. 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. Ill 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, circa 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:

Peru: Obrajillo, Wilkes Exp. (US); rock ledges at cascades of
Rio Chillon, above Obrajillo, Lima, 3100-3300 m., June 13-23,
1925, Pennell 14401 (GH); abundant on open rocky slopes,
Huaros, Lima, 3200-3600 m., June 23, 1925, Pennell 14784 (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.

(Vol. 14

56 ANNALS OF THE MISSOURI BOTANICAL GARDEN

"Peru: Prov. Huamachuco, Depart. Libertad, liber Huama-
chuco, Grassteppe mit eingestreuten immer oder regengriinen
Straiichern, 3400-3500 m. ii. M., 1 m. hoher Strauch, Bliiten
violett (Fl. 3. VII. 1914.— Weberbauer n. 7008!).

"Die Art gehort in die Nahe von S. elliptica (Ruiz et Pav.)
Briq. und S. Matthewsii Briq., die erstere Art weicht schon durch
eine viel langere Korolla ab, die letztere besitzt nach der Be-
schreibung eine langere, innen kahle Krone und kurzere Kelch-
z aline."

Mihi ignota; ad S. ellipticam affinis videtur. Inquire praeterea
in S. rugosani Briq. in Engler u. Prantl. Nat. Pflanzenfam., ed. 1,
IV. Abt. 3a, 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. 3a, 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. Geneve

2: 191. 1898.

Fruticulis suaveolens altitudine ad 1-1.5 m., caulibus in basi
lignosis, procumbentibus, ramis teretibus, cortice discedente,
ramulis pubescentibus, quadratic, 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. Ill 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 :

Peru: Dombey 291 (GH).

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

Cauquen

17, 1882, Ball (GH; NY

Maule, Reed (GH) ; Valparaiso, Arnott (NY)
Statin (NY); no data, Wilkes Exp. (ASP; US; NY); Santiago j
montana, Dec. 5, 1920, Bro. Claude- J oseph 1821 (US) ; Valpar-
aiso, Eights (US) ; Valparaiso, in Gebtischen, 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; UC); 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- J oseph 956 (US) ; near Valparaiso,'
June 1885, Rusby 1061 (ASP; US); no data, Styles (ASP); in
fruticetis apricis collium St. Jago, Quillota, Feb., 1829, Sept-

, 1829, Bertero 291, 1015 (MBG; GH; NY); Prov. Curic6,
hacienda Monte Grande, 1000 m., Dec, 1924, Werderman 547
(UC); Cerro Echaurrina (San Fernando), Prov. Colchagua, Oct.
1925, Montero IS (GH).

Oct

6. Satureia Matthewsii Briq. in Ann. Conserv. et Jard. Bot.
Geneve. 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 cm. distantibus; foliis 1-1.5
cm. longis, elliptico- vel rhomboideo-obovatis, frequenter subro-
tundatis, apice saepius obtusis, in basi rotundato-extenuatis.

[Vol. 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 approximates, 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 cm. 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 :

Peru: Near San Felipe, Cajamarca, 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. 3a, 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, antic is e corolla
5-6 mm. exsertis, posticis e tubo exsertis; thecis late divaricatis,
.6 mm. loneis: stvlo e corolla 8 mm. exserto; nuculis non visis.

1927]

EPLING — SOUTH AMERICAN LABIATAE. Ill 59

S. guamaniensis , as suggested by its author, 1 is scarcely separ-
able from Gardoquia obovata R. & P.

Specimens examined:

Peru: between Cuancabamba and Oyavaca, 3200 m., May,
1912, Weberbauer 6823 (FM, type collection of S. guamaniensis
Mansf ) .

mm

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 ; f oliis .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
longis, anguste lanceolatis, acutissimis, duobus anticis quam
posticis paulo longioribus, pedicellis gracilibus circa 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 :

Ecuador: Quitensian Andes, Jameson (US).

C. Discolor es

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

am nachsten stehende Art S. obovata (Ruiz et Pav.) Briq. unterscheidet sich

: (nicht mehr ausreichenden) Beschreibung nur durch langere Blatter,

behaarte zweige an der Basis behaarte kelch und innen Kahle Krone."— Mans-

nach

feld, I.e.

(Vol. 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, IV. Abt. 3a, 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
cm. longis, ramulis gracilibus, ascendentibus, quadratis, angulis
obtusis, superne cano-puberulis vel tomentosis, internodiis
maximam partem quam foliis brevioribus; foliis 1-1.5 cm. 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, inforiore 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 cm. longis, extus
pubescentibus, tubo superne gradatim ampliato, intus tenuiter
piloso, labro subrecto 5 mm. longo, emarginato, labiolo subaequi-
longo, lobo medio lateralibus nunc 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. Kunihii {grandi flora) , 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. Ill 61

Ecuador: Banos, Prov. Tunguragua, 2000 m., Tate 607 (US);
ad sepes, in planitie Rumibamba, Quito, Hartweg 1340 (NY);
ad pontem Guapalo prope Quito Hartweg 1839 (NY) ; Punin,
Quebrada Chalan, 2779 m., Oct. 28-Nov. 4, 1923, Anthony &
Tate I$3 (US) ; ad vicum Guapalo prope Quito, 2076 m., Jameson
659 (ASP); no data, Jameson (US); Quitensian Andes, 1855,
Couthouy (GH) ; 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., May 31-June 3, 1922, Macbride & Featherstone 2084
(forma villosa) (MBG; FM).

10. Satureia thymoides Briq. in Engler u. Prantl, Nat. Pflan-
zenfam., ed. 1, IV. Abt. 3a, 300. 1897.

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

"Frutex ramosissimus ; ramis tetragonis, pubescentibus. Folia
opposita, brevissime petiolata, 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 ; inf eriore
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. 3a, 300. 1897.

Gardoquia foliolosa Benth. in DC. Prodr. 12: 238. 1848.

[Vol. 14

62 ANNALS OF THE MISSOURI BOTANICAL GARDEN

"foliis 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
nunc fere glabris, nunc densissime pilis longis ornatis imprimis
differt.

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

Gardoquia 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 nee 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 nun. 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. prof undo, labiolo aequilongo, lobis sub-
aequalibus; staminibus didymis supra tubi medium sitis, anticis
longioribus, omnibus inclusis, thecis divergentibus, .6 mm. longis;
stylo e corolla vix exserto; nuculis non visis.

Specimens examined :

Venezuela: prope coloniam Tovar, 1854-5, Fendler 2058

1927]

EPLING — SOUTH AMERICAN LABIATAE. Ill 63

(MBG; GH); Silla de Caracas, 2600 m., Apr. 27, 1884, John 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
-3 ft. tall; fl. violet) (US; NY).

13. Satureia acutifolia Briq. in Engler u. Prantl, Die Nat.
Pflanzenfam., ed. 1, IV. Abt. 3a, 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) f asciculatis ; foliis 1.5-3 cm. 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) cm. 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 visis.

Specimens examined:
Peru : f 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 circ. 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 circ. usoue ad medium con-

[Vol. 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 circ. 9 mm. longis, posticis 4-5 mm. longis,
antheris divergentibus, anticis et stylo exsertis. Stylus 32 mm.

longus.

"Peru: Depart, und Prov. Huancavelica, sudliche Talwand des
Mantaro iiber Iscuchaca, Grassteppe mit eingestreuten Strauch-
ern, 3600 m.u.M., bis 2 m. hoher Strauch mit roten Bluten (Fl.
15. VI. 1910.— Weberbauer n. 5677!).

' ' Die Art steht nach der Lange der Krone z wischen S. Pavoni-
ana Briq. (Gardoquia incana Ruiz et Pav.) und S. discolor (Kunth)
Briq. in der Mitte ; S. discolor hat f erner eingeschlossene Stamina
und inne kahle Kronrohre, S. Pavoniana oberseits bleiche Blatter
(nach der nicht mehr ausreichenden Beschreibung) . S. thymoides
(Kunth) Briq. weicht durch den fast herzformigen Blattgrund
und zuriickgerollten deutlich gezahnten Blattrand ab, steht aber
sonst besonders im Bliitenbau der S. mantaroensis wohl am

nachsten."

Ex descriptione ab S. Pavoniana vix distincta videtur.

15. Satureia Pavoniana Briq. in Ann. Conserv. et Jard. Bot.

Geneve 2: 189. 1898.

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 cm.-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. Ill 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 cm. 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 circa 5
mm. exserto; nuculis circa 1.5 mm. longis, oblongis, 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 1J±11 (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 f oliorum axiliis ;
corollis 1.5-3 cm. longis.

16. Satureia pallida sp. nov.

Frutex ramosus, altitudine ut videtur circa 1 m., ramis as-
cendentibus, teretibus, lignosis, cortice discedente, ramulis di-
varicatis, glabris, quadratis, angulis acutis, internodiis foliis sub-
aequilongis vel brevioribus; foliis 1-1.5 cm. longis, maximam
partem ovalibus vel elliptico-oblongis, obtusis, in basi saepius
rotundatis, utrinque glabris et pallidioribus, margine leniter
revoluta, integra, petiolis 1-2 mm. longis elatis; floribus 1-3 in
foliorum supremorum axiliis 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

[Vol. 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 $♦ Pavoniana and at first thought to be a
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. The 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 S. pallida. The species
is intermediate with the groups Discolores and Striatae.

Specimens examined:

Bolivia: Bang, no data (NY); Turedon, Bolivian plateau,
1891, Bang 1127 (US; NY, type; GH).

17. Satureia glabrata Briq. in Engler u. Prantl, Die Nat.
Pflanzenfam., ed. 1, IV. Abt. 3a, 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 elliptieo-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
confornialibus, 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
brevioribus, 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. Ill 67

Specimens examined:

Ecuador: vie. of Tabl6n de Oiia, Sept. 27, 1918, Rose

(US).

18. Satureia taxifolia Briq. in Engler u. Prantl, Nat. Pflanzen-
fam., ed. 1, IV. 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 1 1 : 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:

Ecuador: Loja, between San Lucas and Oiia, 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 S. glabratae, foliis ramulisque
tamen utrinque tomentellis, eis 10-12 mm. longis, oblanceolatis,

[Vol. 14

68 ANNALS OF THE MISSOURI BOTANICAL GARDEN

obtusis, in basi angustatis, venis prominulis, margine subserrata,

subrevoluta, petiolis subnullis; fioribus 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:

Colombia: Cisne, Oct. 30, 1876, Andre 4331 (NY, type, 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 Pa von, 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 circa 4-6, pedicellis subnullis, pubes-
centibus; floribus in axillis solitariis, bracteolis foliis conformali-
bus; calycibus 6-7 mm. longis, profunde sulcatis, 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, anticis labrum subaequantibus, omnibus supra tubi medium
sitis; thecis divergentibus, .5-6 mm. longis; stylo e corolla 3 mm.
exserto; nuculis non visis.

Specimens examined :

Peru: Mito, July 23- Aug. 14, 1922, Macbride & Featherstone
1734 (FM; MBG); 12 mi. south of Panao, 3,076 m., July 4-10,
1922, Macbride & Featherstone 2213 ("a common hillside shrub")

(FM;MBG).

1927J

EPLING — SOUTH AMERICAN LABIATAE. Ill 69

21. Satureia plicatula sp. no v.

Suffrutex procumbens, caule duro, ramisque teretibus, ramulis
subquadratis, puberulis, nodis approximatis, saepe in ramulis
lateralibus brevibus conf ertissimis ; 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 nee 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., Llata, Aug. 21,
1922, Macbride & Featherstone 2238 A (FM, type).

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

Gardoquia Jamesoni Benth. Lab. Gen. et. Sp. 404. 1834, et in
DC. Prodr. 12: 239. 1848.

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, subglabris, fauce intus nudis, den-

[Vol. 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 patenter exserto; nuculis non visis.

Specimens examined:

Colombia: Ayapel, Andre K 1564 (NY) (doubtful station).

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

23. Satureia connata nom. nov.

S. microphylla Briq. in Engler u. Prantl, Nat. Pflanzenfam., ed.
1, IV. Abt. 3a, 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.

f Satureia ericoides Willd. ex Benth. in Linnaea 1 1 : 328. 1837.

Frutex ut videtur altitudine circa 1 m., ramis ascenden tibus,
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 f oliosis, pedicellos breviter superantibus ;
calycibus 5 mm. longis, extus puberulis fauce intus glabris,
dentibus anticis ovatis, acutis, 1 mm. longis, erectis, tribus posticis
connatis, subnullis, labia itaque subintegra, pedicellis 1-1.5 mm.
longis; corollis coccineis, 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-
nccmilonffis. naulo suora tubi medium sitis, 1.5 mm. longis, in

19271

EPLING — SOUTH AMERICAN LABIATAE. Ill 71

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

Specimens examined:

Ecuador: ranch between Ibarra and Tulcan, 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 S. rigidula);
corollis 9-25 mm. longis.

This group represents the extremes of adaptation to an arid
habitat to be found in the genus. The 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. 3a, 300. 1897.

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

Suffrutex procumbens, ramosus, ramis teretibus, cortice dis-
cedente, ramulis pubescentibus, quadra tis, 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 nudis, 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 :

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

[Vol. 14

72 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. ianceolato-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-flori; 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 Angasmarca, lockeres, von Grassteppe durchsetztes
Gestraueh, 3000-3100 m., l /2 m. hoher Strauch mit purpurnen
Bliiten (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. no v.

S . fasciculata Briq. in Engler u. Prantl, Nat. Pflanzenfam., ed.
1, IV. Abt. 3a, 300. 1897, nee Rafin. Prec. Decouv. 39. 1814;
nee Tausch in Syll. Ratisb. 2: 248. 1828.

Gardoquia fasciculata Benth. PI. 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.

1927]

EPLING — SOUTH AMERICAN LABIATAE. Ill 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-
cussatim instructis, in spicam densam 1-4 cm. longam ad ramu-
lorum apices confertis, bracteis subulatis, calycibus brevioribus;
calycibus 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, circa 10 mm. longis, extus pubescentibus, tubo
superne ampliato, decurvo, intus infra stamines pubescentibus,
labro bifido, sinu circa .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,

fuse is.

Specimens examined:

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

Ecuador: plains of Pamasqui and San Antonio, 2615 m.,
Jameson (NY); in planitie Rumibamba necnon juxta pontem
Guapalo prope Quito, Hartweg 1338 (NY) ; no data, Lehmann 6347
(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); Alausi, Chim-
borazo, 2500 m., July 19, 27, 1923, Hitchcock 20705 (US).

27. Satureia sericea Briq. in Engler u. 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 f asciculatis ; floribus
tribus in axillis, rarius solitariis, brevissime pedunculatis, brac-
teolis primis omnino foliosis, secondariis conformalibus sed

74 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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 cocci neis 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:

Peru: sunny slopes, "Chunmis," Chasqui, 3230 m., Sept. 27,
1922, Macbride & Feather sUme 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; FM); 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 Pa von, 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 subf astigiatis ; 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; calycibus 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 3HL
mm. longis; corollis coccineis 20-22 mm. longis, tubo extra caly-
cem multo dilato, extus molliter villoso intus infra labiolum pi-

1927]

EPLING — SOUTH AMERICAN LABIATAE. Ill 75

losis, labro erecto 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 non 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 segregation from S. revoluta as understood by him.

Specimens examined:

Peru: Wilkes Exp. (US; GH); between Cuancabamba and
Oyavaca, 3200 m., May 1912, Weberbauer 6333, (FM, type collec-
tion of S. insignis 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.
Geneve 2: 191. 1898.

SufFrutex ut videtur, altitudine circa 15 cm., ramis e caudice
lignoso erectis, virgatis, subteretibus, pubescentibus, nodis inter
se 2-5 mm. distantibus, ramulis brevibus, erecto-ascendentibus;
foliis circa 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 a 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 paulo exserto; nuculis non visis.

Specimens examined:

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

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

(Vol. 14

76 ANNALS OF 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. In Peruvia (Ruiz et Pavon!). Frutex ra-
mosissimus, ramis duris tetragonis junioribus rufo-villosis. Folia

1-1 J/2 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. 3a, 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. 3a, 300. 1897.

Thymus nubt 'genus Kunth in Humboldt et Bonpland, Nov. Gen.

etSp. PL 2:310. 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. Ill 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., Merida, Sept. 14, 1922, Jahn 1151 (US); Laguna Verde,
Paramo Mucuchies, Merida, 3384 m., 1922, de Bellard 19 (US) ;
summit of Paramo, Quirora, 3200 m., Merida, Feb. 24, 1922,
Jahn 882 (US) ; Sierra de Nevada de Merida, Laguna del Gallo,
4070 m., Dec. 1910, Jahn 74 (US).

Colombia: in monte ignivomo Azufral, May 18, 1876, Andre
8249 (NY); Hacienda de Antisana, Hartweg 1337 (NY); Ruiz,
3000 m., 1918, Dawe 750 (NY); Canaan, Mt. Purace, 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; MBG; GH; NY);
Westabhange des Param6 de Ruiz, El Cauca, 3000-3500 m., Sept.
11, 1883, LehmannSlll (US).

Ecuador: Andes near Quito, 4000 m., Couthouy (NY); Andes
of Quito, 4000 m., Jameson 217, "crescit in graminosis alpinis
(vernacule "paramo de los Andes"), 4000 m., Jan. 21, 1856 (US;
GH); Paramo de Tuza, 3400 m., Jan. 31, 1881, Lehmann 3092 a
(US) ; between Urbina and Mt. Chimborazo, 3600-4500 m., Prov.

[Vol. 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
Cuenca, 2700-3300 m., Azuay, Sept. 9-10, 1923, Hitchcock 21635

(US).

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 nee 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).

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

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

32. Satureia Darwinii Briq. inEngleru. Prantl, Nat. Pflanzenfam.,

ed. 1, IV. Abt. 3a, 300. 1897.

Micromeria Darwinii Benth. in 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 circa 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-
curvis, 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).

1927]

EPLING SOUTH AMERICAN LABIATAE. Ill 79

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

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

Herba perennis humilis, altitudine 5-6 cm., 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 circa .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, subaequalibus sed duobus anticis
tamen longioribus, intus tenuiter hirsutis, pedicellis subnullis;
corollis 7-8 mm. longis, extus villosulis, tubo superne gradatim
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 characteristics of sections Gardoquia and
Xenopoma.

Specimens examined:

[Vol. 14

80 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Peru: rainy-green formation, 3100 m., Carumas, Moquegu
Feb. 21-March 6, 1925, Weberbauer 7259 (FM, type).

35. Satureia boliviana Briq. in Engler u. Prantl, Nat. Pflan-
zenfam., ed. 1, IV. Abt. 3a, 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 u. 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, graeilibus, 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
cm. longis, eis in ramulis florentibus densis saepius oblongis,
obtusis, in basi angustatis, margine integra et revoluta, plerum-
que .5-1 cm. 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 3 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 divergent ibus; stylo corolla aequilongo;
nuculis non visis.

Satureia Kuntzeana was based by Briquet on a collection by
Kuntze at Tunari, 4000 m., Apr. 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'etre ecarlates, 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 S.

1927]

EPLING — SOUTH AMERICAN LABIATAE. Ill 81

boliviano,. It corresponds in every way to Briquet's description
of the type.

Specimens examined :

Bolivia: 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 (GH;
NY); an sonnigen Abhangen, 3600 m., La Paz, Sept. 3, 1906,
Buchtien 438 (US) ; Tiahuanaco, 3900 m., Nov. 1913, Buchtien 438
(GH; NY); La Paz, sonnige Abhange, 3700 m., Nov. 3, 1906,
Buchtien (GH); vie. of La Paz, Aug. 15, 1914, Rose 18894 (US;
NY) ; no data, Bang (NY) ; viciniis Achacache, monticula Arichaca
in petrosis, 4000 m. alpine, Jan. 1859, Mandon 518 (apparently a
dwarf from 15 to 20 cm. 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; GH; NY).

Var. tarijense comb. nov.

Xenopoma bolivianum Griseb. var. tarijense Wedd. in 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 1037 (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 circa 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,

[Vol. 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 S. boliviano, speciminibus substerilibus valde
similis, floribus tamen differt.

Specimens examined :

Bolivia: 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 axillare Rusby, Mem. Torr. Bot. Club 6: 108.

1896.

B. unifiorus Busby in Briq. 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 examined :

Bolivia: 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. Ill 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
circa 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 :

Peru: 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.

Micromeria Gilliesii 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.
Wystropogon minutus Briq. in Bull. THerb. Boiss. 4: 803.

1896.
Satureia Gilliesii Briq. in Engler u. Prantl, Nat. Pflanzenfam.,

ed. 1, IV. Abt. 3a, 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

84 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Voi.. 14

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:

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

Bolivia: Oruro, Tapacari, 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 254 (NY); Sierra de Tucuman, Jan. 10-17, 1878, Lorentz &
Hieronymus 722 (US) ; Capillitas, Catamarca, 1875, collector un-
known (ASP); El Candado, Andalgala, Oct. 2, 1916, Jdrgensen
1189 (MBG;UC).

Species Mihi Omnino Ignotae

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

Gardoquia salviaefolia Colla Mem. Accad. 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.

"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? An eadem ac Hedeoma multiflora?" (Benth. in DC.
Prodr. 12: 223. 1848.)

1927]

EPLING — SOUTH AMERICAN LABIATAE. Ill

85

Index Specierum

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

PAGE

Bystropogon

axiilare 82

minutus 83

uniflorus 82

Gardoquia 47

acutifolia 63

argentea

71

breviflora 59

chilensis 56

discolor 62

elliptica 54

fasdculata 72

foliolosa 61

Gilliesii 56

glabrata

incana

66

64

Jamesoni 69

microphylla 70

multiflora 53

obovata .
revoluta

rugosa

58

74

76

salviaefolia 84

sericea 73

striata 68

taxifolia 67

thymoides 61

tomentosa

60

Micromeria 47

boliiriana 80

bonariensis 84

Danvinii 78

eugenioides 83

Gilliesii 83

nubigena 76

pasilla 79

pulchella

84

Micromeria sect. Hesperothymus .... 50

Rizoa 47

ovatifolia 53

Satureia 47

— Discolores 59

Ellipticae 53

PAGE

Obo vatae 56

Revolutae 71

Striatae 65

acutifolia 63

Andrei 67

argentea 71

axillaris 82

boliviana 80

boliviana var. tarijense 81

bonariensis 84

brevicalyx 82

breviflora 59

Brownei 51

Brownei subsp. eubrownei 51

chilensis 56

connata 70

Darwinii 78

discolor 62

elliptica 54

ericoides 70

eugenioides 83

fasdculata 72

foliolosa 61

Gilliesii 56

Gilliesii 83

glabrata 66

guamaniensis 58

insignis 74

Jamesoni 69

Kuntzeana 80

Lindeniana 75

Loesneriana 55

mantaroensis 63

Matthewsii 57

microphylla 70

multiflora 53

nubigena 76

nubigena var. glabrescens 78

obovata

58

oligantha 83

pallida 65

Pavoniana 64

plicatula 69

86

[Vol. 14, 1927]

ANNALS OF THE MISSOURI BOTANICAL GARDEN

PAGE

pusilla 79

revoluta 74

rigidula 72

rugosa 75

sericea 73

simulans 81

striata 68

taxifolia 67

thymoides 61

tomentosa 60

vana 79

Weberbaueri 72

PAGE

Stachys

8peciosa 54

Thymus 47

Brownei 51

humifusus 76

nubigenus 76

taxifolius 67

Xenopoma 47

bolivianum 80

bolivianum var. tarijense 81

eugenioides 83

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,
Mycologist 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. No 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 in 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. Mo. Bot. Gard., Vol. 14, 1927

(87)

[Vol. 14

88 ANNALS OF THE MISSOURI BOTANICAL GARDEN

running each description, as it were, into a mold or through a
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.

The paper is the outgrowth of a critical study of the species of
the genus Pholiota of the Agaricaceae for the 'North American
Flora.' The drawings illustrating the microscopic details are all
made to the same scale in the different species so that direct
comparisons can be made. All were outlined under the camera
lucida from free-hand transverse sections of the gills of dried
plants after softening with alcohol and water, mounting in KOH,
and staining with dilute eosin. The larger drawings of the spores
were made in the same way except under the oil-immersion lens,
yielding a magnification of about 1800 diameters. The 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 1
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 2 on the species occurring in the region of the Great
Lakes and to which additions were made in two subsequent
papers, 8 The illustrations published by him constitute the finest

1 Bull. N. Y. State Mus. 122: 141-158. 1908.

* Trans. Wis. Acad. Sci. 17: 470-502. 1912.

»I. 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. The 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 1 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 a new angle of at-
tack for taxonomic 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 a sufficient extent in a single genus of that group.
The nearest approach to this has been in the work of Lange on
the Agarics of Denmark. 2 However, the illustrations there are

— _ __ _____ ___ i

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

* Lange, J. E. Studies in the Agarics of Denmark. I. The genus Mycena.
(Dansk Bot. Arkiv 1*: 1-40. 1914); II. Amanita. Lepiota. Coprinus. (I.e.
2': 1-53. 1915); III. Pluteus. Collybia. Inocybe. (I. c. 2 7 : 1-50. 1917); IV.
Pholiota. Marasmius. Rhodophyllus. (I.e. 2 n : 1-46. 1921.)

[Vol. 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.

III. 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
on 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, etc.,
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 a 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 a 10 X or a 12 X 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, etc.). 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

92 ANNALS OF THE MISSOURI BOTANICAL GARDEN

(Vol. 14

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 a 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. The recog-
nition of these groups is a relatively easy matter, since in most
cases they do not intergrade. On 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 phylogenetic 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 a mark of relationship
between P. dura, P. praecox, P. vermiflua, P. temnophylla, P.
Howeana, and P. Acericola. It so 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 a small group centering around P. erebia
that shows close agreement in spore characters and in the char-
acteristic cystidia with long projecting necks. Here 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 cases representing a non-plastic 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.
P. praecox, P. vermiflua, P. Howeana, P. temnophylla, P. Aceri-

cola (also P. dura of Europe).

While P. A cericola frequently grows on rotten wood and perhaps

1 Items are 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.

94 ANNALS OP THE MISSOURI BOTANICAL GARDEN

{Vol. 14

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 a 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 n long,
not truncate. Cystidia abundant, flask-shaped with long necks.

P. erebia, P. ombrophila, P. subnigra, P. platyphylla.

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

P. caperata, P. Mcmurphyi.

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

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

P. terrestris, P. angusiipes.

This group may not be a natural one but does show some
transition over towards the Truncigeni. While not known to
grow on wood, the latter species grows around stumps and is likely
attached to buried wood. P. 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. P. trachyspora would be included were
it not for its rough spores.

Unattached Species of the Humigeni. — (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 Togularis group as referred by Harper.
(3) P. aurea is difficult to interpret. It may grow attached to
buried wood and belong in the Truncigeni 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. oreqonense, P. rubecula.

9. The Spectabilis Group. — Gills bright-colored (yellow or och

and remaining so on drying). Pileus dry

Sp

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.

96

[Vol. 14

MISSOURI

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

I have 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. P. limonella, I would prefer to class in the Tuberculosa
group, rather than here as in Harper's synopsis. P. flammans
departs from the other species of the group in the small stature
and the small spores. P. 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 Group. — Small plants with erect-pointed
dark scales on pileus and stem. No cystidia. Spores smooth.

P. muricata, P. erinaceella.

IS. The Fulvo-squamosa Group. — Large plants with fibrous cu-
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 Truncigeni. — (1) P. Aegerita. Close to
the Mulabilis 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 \i 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. xMUSCIGENI

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

1927]

OVERHOLTS

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 |i
long, and the latter has spores not truncate, rough-walled, and
6-8.5 \x 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
a second annulus-like structure comparable to an imperfect volva,
at its base. For that reason the species is sometimes separated
off in the genus Rozites. The spores are 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
in the genus. The 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 con-
sidered as genera is unthinkable. The 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 a 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.

[Vol. 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. As a
result, some of my conclusions as to the identity of our species
are at variance with those of my predecessors. The 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. My 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 y
due to the early disappearance of the partial veil or the annulus,
and in Flammula the veil fragments may at times persist as an

mcom

Key to the Species 1

1. Plants at maturity not more than 4 cm. broad 2

Plants at maturity 4-15 cm. or more broad 29

2. Plants growing on the ground or among mosses 3

Plants growing on stumps, trunks, or rotting logs 14

3. Pileus distinctly fibrillose or squamulose, especially at the center 4

Pileus entirely glabrous at the center, the margin occasionally with white

fibers from the veil 5

4. Stem distinctly scaly below the sheathing veil remnants; gills bright

cinnamon in dried plants; known only from Oregon; spores smooth,

5.5-6.5 X 3.5-4.5 M SO. P. terrestris

Stem not scaly, somewhat fibrillose; gills deep brown in dried plants;
known only from Colorado; spores rough-walled, 7-9 X 5-6 \i.
19. P. trachyspora

5. Plants growing among Polytrichum mosses 18. P. minima

Plants growing in wet places among Sphagnum mosses 9. P. mycenoides

Plants growing on the ground or on leaf mold 6

6. Pileus slimy-viscid; stem heavily white-floccose below the annulus; known

only from Washington and Oregon 22. P. albivelata

Pileus and stem not as above 7

1 For a supplementary key to the species see p. 169. It 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.

[Vol. 14

100 ANNALS OP THE MISSOURI BOTANICAL GARDEN

7. Cystidia present and projecting conspicuously from the sides of the gills

(figs. 40, 48) 8

Cystidia absent 11

8. Cystidia 15-20 /i in diameter at the broadest part; spores truncate at one

end, 8-10.5 m long 9

Cystidia 7-12 n in diameter at the broadest part; spores not at all trun-
cate, 10-15 m long 10

9. Spores 4.5-6 ju broad 2. P. praecox

Spores 6-7 fi broad 5. P. temnophylla

10. Plants small, less than 2 cm. in diameter, uniformly fuscous in color;

known only from Washington IS. P. subnigra

Plants somewhat larger; lighter in color than the above; spores 10-12 /x

long 11. P. platyphylla

Plants as in the last but spores up to 15 m long 12. P. erebia

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

changing on drying 8. P. rugosa

Stem 2-10 cm. long; pileus w T atery-brown, drying to ochraceous 12

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

10. P. filaris

Stem not filiform, 2 mm. or more thick; pileus at maturity more than 2

cm. broad 13

13. Spores 6.5-9 ^ long 6. P. blattaria

Spores 9-10.5 m long 7. P. togularis

14. Pileus entirely glabrous and hygrophanous, 1 the margin usually striatulate

when moist (pi. 15) 15

Pileus scaly 2 or densely fibrillose from the first or becoming so in mature

plants; hygrophanous only in P. confragosa, but often viscid 20

15. Stem distinctly scaly; gills not forked SO. P. mutabilis

Stem not at all scaly; gills forked 28. P. furcata

Stem not scaly; gills not forked 16

16. Pileus viscid when fresh 27. P. discolor

Pileus not viscid 17

17. Stem hollow, with irregular transverse partitions; known only from Cali-

fornia; on small twigs 21. P. anomala

Stem stuffed or hollow, not internally partitioned; on logs or on sawdust 18

18. Cystidia absent; spores entirely smooth, slightly truncate at one end (fig.

95) 29. P. marginella

Cystidia present, flask-shaped, projecting (fig. 83); spores not truncate,

smooth at first, usually somewhat roughened when mature 19

19. Annulus funnel-shaped, persistent, conspicuous; pileus never more than

2 cm. broad 26. P. unicolor

Annulus not funnel-shaped, sometimes persistent as a ring but often

evanescent; pileus often 3-4 cm. or more broad 25. P. marginata

20. Pileus dull cinnamon, dull tawny, or brown 21

Pileus bright-colored, i.e., pinkish-red, yellow, ochraceous-orange, or

bright tawny in fresh plants 22

1 i. e. watery-brown when moist, ochraceous as the pileus begins to lose moisture.

2 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. Pileus densely floccose-fibrillose or floccose-squamulose as seen under a

lens 47. P. confragosa

Pileus densely covered with small erect spine-like scales #?. P. erinaceella

22. Spores 3-5.5 m long; dried plants with a yellow pulverulent appearance.

38. P. flammans

Spores 6 m or more long; pileus not pulverulent 23

23. Pileus viscid when fresh 24

Pileus dry when fresh 26

24. Pileus lemon-yellow or bright sulphur-yellow 25

Pileus with greenish or purplish tinges and not uniformly nor brightly

yellow Flammula polychroa 1

25. Stem concentrically ringed or peronate with white fibrils 60. P. ludfera

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 plants 27

Gills not bright-colored, rather cinnamon or brown 28

27. Young plants bright-colored and only silky, becoming scaly and tawny

when mature; taste mild; spores smooth 53. P. curvipes

Young plants dark red and plush-like, becoming somewhat areolate and
pinkish-red when mature; taste bitter; spores minutely roughened (fig.
107) 35. P. luteofolia

28. Pileus drying brown or cinnamon-brown, densely covered with tawny

fibrils or tufted and sub-erect scales (pi. 14, fig. 3); stem not bulbous
49. P. muricata

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

a distinct bulb just at base (pi. 14, fig. 4) 52. P. tuberculosa

29. Plants growing on the ground and not around old stumps and logs or from

buried wood 30

Plants growing on wood, on sawdust piles, or around old stumps or rotting

logs, sometimes from buried wood 42

30. Pileus entirely glabrous or at most only with marginal fibers from the veil. ... 31
Pileus fibrillose, floccose, squamulose, or scaly, at least over the center 37

31. Spores more than 10 m long 32

Spores averaging less than 10 n long 34

32. Cystidia pyriform to subglobose (figs. 1, 3, 5), not projecting conspicu-

ously, about 20 m in diameter; spores ovoid with a truncate apex (fig. 2)
1 . P. vermiflua

Cystidia elongated, projecting prominently (20-40 v) (fig. 43), 5-7 m in

diameter; spores elongate-elliptic, not truncate 33

33. Pileus 2-5 cm. broad; gills 2-4 mm. broad 12. P. erebia

Pileus 4-10 cm. broad; gills 4-6 mm. broad H. P. ombrophila

34. Plants western; pileus slimy-viscid; stem conspicuously white- tomentose

just below the annulus 22. P. albivelata

Plants eastern; pileus dry or very slightly viscid in very wet weather;
stem not white-tomentose 35

35. Spores 4-6 m long; annulus membranous, evanescent 23. P. duroides

1 This species is so frequently met and at times presents a fairly well-formed annulus,
so I have thought it best to include it in the key at this point. No description is
given in the text.

102 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

Spores 5-8.5 m long; annulus not membranous but forming a cottony roll

on the stem 24- P. Johnsoniana

Spores 8-10.5 m long 36

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

open grassy woods, or among straw or other waste vegetable matter

carried into the woods; pileus not rugose 2. P. praecox

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

times 3. P. Acericola

37. Spores 12-17 n long, decidedly rough-walled (fig. 58) 38

Spores 9-12 m long, smooth or nearly so 15. P. aurea

Spores 6-9 ju long, decidedly rough-walled (fig. 64) 19. P. trachyspora

Spores 6-9 m long, entirely smooth 39

Spores 4-6 m long, smooth 23. P. duroides

38. Pileus dry or moist, in youth showing fine whitish fibrils under a lens;

plants eastern 16. P. caperata

Pileus slimy-viscid, glabrous; plants western 17. P. Mcmurphyi

39. Pileus very slightly squamulose only at the center 24. P. Johnsoniana

Pileus uniformly squamulose or nearly so 40

40. Cystidia present, rather abundant, hyaline, fusoid, projecting rather

prominently (fig. 149a) 46. P. Schraderi

Cystidia absent or represented by small inconspicuous brown bodies in

the hymenium 41

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

45. P. rigidipea

Scales of the pileus minute and dot-like (pi. 23), abundant; pileus brown

or pinkish-cinnamon 5 4- P. angustipes

42. Pileus glabrous or essentially so at all stages 43

Pileus more or less distinctly fibrillose or scaly 48

43. 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
or tawny, concolorous or darker in dried plants; not hygrophanous;
cystidia none 46

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

ameter in the broadest part, not with an elongate tip (figs. 9, 11)

3. P. A cericola

Spores not truncate, smooth; cystidia 6-10 /i in diameter in the broadest

part, not with an elongate tip (fig. 98) 31. P. Aegerita

Spores not truncate, rough-walled at least at maturity; cystidia 10-15 m

in diameter in the broadest part, with an elongate tip (figs. 83, 89) 45

45. Pileus viscid 27. P. discolor

Pileus dry or moist 25. P. marginata

46. Stem 3-6 cm. long 34. P. rubecida

Stem 6-15 cm. long 47

47. Spores rough-walled (fig. 103); gills not interveined; plants known only

from the eastern states S3. P. cerasina

Spores smooth; gills strongly interveined; plants known only from Oregon
32. P. oregonense

1927]

OVERHOLTS THE GENUS PHOLIOTA IN THE UNITED STATES 103

48. Spores averaging 4-6 /* long 49

Spores averaging 6-11 y. long

50

Spores averaging 11-14 fx long 89. P. albocrenulata

49. Pileus lemon-yellow, zinc-orange, or tawny, with concolorous floccose-

fibrillose, sub-appressed scales; stem 2-5 mm. thick 88. P. flammans

Pileus pale cinnamon or cinnamon-buff, with concolorous erect-pointed

scales (pi. 22); stem 5-12 mm. thick 43. P. squarrosoides

50. Gills bright-colored, i.e., yellow, bright ferruginous, or ochraceous-orange,

and remaining so in dried plants 51

Gills cinnamon to dark ferruginous, tawny, or brown 53

51. Pileus with greenish tints and strongly areolate from an early stage (pi. 18);

spores slightly roughened (fig. 113) 87. P. aeruginosa

Pileus pale pinkish-red, frequently slightly areolate at the center; spores

slightly rough (fig. 105) 35. P. luteofolia

Pileus apricot-yellow to zinc-orange or tawny, not at all areolate 52

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

86. P. spectabilis

Spores smooth; pileus 2-5 cm. broad; stem 2-5 mm. thick 53. P. curvipes

53. Plants with one of the following characteristics:

(a) Spores decidedly rough-walled (fig. 109) 86 P. spectabilis

(b) Pileus uniformly covered with minute dot-like scales (pi. 23)

scarcely visible except under a lens 54.. P. angustipes

(c) Pileus with large white floccose patches or soft white scales

56. P. destruens

Plants with none of the above characters 54

54. Pileus dry 55

Pileus viscid or slimy-viscid 58

55. Cystidia entirely lacking 56

Cystidia at least in part as brown organs among the basidia 57

Cystidia altogether hyaline, projecting prominently (fig. 149a) . .46. P. Schraderi

56. Pileus 2-6 cm. broad, with small scales; stem 1.5-3 cm. long, 1-5 mm.

thick 62. P. tuberculosa

Pileus 6-12 cm. broad, with a conspicuous fibrous-scaly covering (pi. 13);
stem 5-8 cm. long, 8-10 mm. thick 55. P. fulvo-squamosa

57. Pileus with many large recurved, often floccose scales (pi. 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

58. Cystidia none; plants less than 5 cm. broad at maturity 59

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

broad Flammula polychroa

i

Cystidia as brown organs between the basidia (fig. 127a, 135a); mature

pileus 5-15 cm. or more in diameter 60

59. Stem marked with rings of white fibrils; spores 7-9 m long 50. P. lucifera

Stem not so marked; spores 6.5-7.5 m long 51. P. limonella

60. Spores 9-1 1 m long; known only from Colorado, New Mexico and Wy-

oming 40. P- aurivelloides

Spores mostly 7-9.5 m long, occasionally slightly longer 61

1 See footnote, bottom p. 101.

[Vol. 14

104

ANNALS OF THE MISSOURI BOTANICAL GARDEN

61. Fileus with large appresscd, spot-like scales; stem increasingly scaly-
downward; pileus tawny or rusty in color 4L P. aurivella

Pileus with medium-sized scales, often recurved; stem increasingly scaly

upward; pileus yellow to zinc-orange J+2. P. adiposa

I. Spores with a truncate apex (fig. 2) (see also II, p. 117).

A. Cystidia present in the hymenium: largest plants more than 4 cm. broad
(see also B, p. 113).
1. Spores 10-14 m long.

1. Pholiota vermiflua Peck, Rept. N. Y. State Mus. 31: 34.

1879.

PL 8.

Pileus 2-12 cm. broad, hemispheric to campanulate or plane,
white, cream-color or massicot-yellow, retaining these colors in

drying, dry or moist or

in wet weather almost
viscid, glabrous or the
margin fibrillose from
the veil, the center of-
ten becoming 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,

forming a superior, often

Figs. 1-5. P. vermiflua: 1, section of hymenium Qn ocnonf on-milnc rw

the tvne sneeimens showinc the inflated cvstidial evanescent annUlUS, Or

of the type specimens showing the inflated cystidial
cells (a) and the spores (b), X550; 2, spores, X1200
(note the truncate apex); 3, three isolated cystidia
from the type specimens, X 550; 4 and 5, more glo-
bose cystidia not projecting, X 550.

adhering to the margin
of the pileus; stem cen-
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 n; cystidia present,
subglobose to pyriform with a tapering base, 35-45 X 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
Scarsdale, N. Y.; State College, Pa.; Chapel Hill, N. C; West
Elkton, Ohio; Greencastle, Ind.; Lexington, Ky.; St. Louis, Mo.;
Rooks Co., Kan.; Denver, Colo.; Berkeley and San Francisco,

Cal.

Illustration: Harper, Trans. Wis. Acad. Sci. 17: pi. 28 C-F, 29;

18: pi. 11-13 (as P. dura).

Closely related to P. praecox with which it has often been con-
fused. The larger spores will invariably separate it from that
species. The thin, slender-stemmed, Panaeolus-like form with a
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 jjl long, and indicates that in the true P. dura
they are the same, although all earlier records report them as ap-
proximately 8-10 [i 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 is distinct in the larger spores. In
P. praecox they rarely reach a length of more than 10 p.

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. praecox.

2. Spores 8-10.5 ju long.

106

(Vol. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

2. Pholiota praecox (Pers.) Fries, Syst. Myc. 1: 282. 1821.

Agaricus praecox Pers. Comm. Fung. Bavar. 89. 1800.

PI. 9.

Pileus 2-14 cm. 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

Figs. 6-8. P. W aeco7: 6, section of the hy- decurrent tooth, medium-
^^r&^^^yffft^/tf ?ftf th f yf°l close, whitish, becoming

brown or rusty-brown,

honey-yellow to clay-color
or snuff-brown in herbarium specimens, 3-12 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 cm. long, 3-20 mm. thick; spores ovoid, usually with a trun-
cate apex, smooth, deep brown, 8.5-10 X 4.5-6 [&; cystidia pres-
ent, flask-shaped or vcntricose, sometimes rare, projecting some-
what, 14-18 (x 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.;

. ... 'P'

(b), X 550; 7, three types of cystidia isolated
from the hymenium, X 550; 8, spores, X 1200.
All from Overholts Herb. 3318.

1927]

OVERHOLTS

107

West Elkton, Ohio; Tolland, Lake Eldora, and South Boulder
Canon, 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, pi 42; ed. 2, pi. 46; Berke-
ley, Outl. Brit. Fung., pi 8,f. 1; Bresadola, Fung. Mang. pi 49;
Cooke, 111. Brit. Fung. pi. {360) 381; Gillet, Champ. Fr. pi {292)
524; Hard, Mushrooms, /. 209; Harper, Trans. Wis. Acad. Sci.
17: pi 27, 28 A, B; Kauffman, Agar. Mich, pi 59; Murrill,
Mycologia 3: pi 49, f. 1; Patouillard, Tab. Anal. Fung. /. 112;
Peck, Rept. N. Y. State Bot. 49: pi 46; Peck, N. Y. State Mus.
Mem. 3: pi 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). The spores are
distinctly smaller than in that species, however. Kauffman's
record (9-13 X 6-7 y.) 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. The writer
collected this plant in abundance in the Missouri Botanical

Garden

No 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-

[Vol. 14

108

ANNALS OF THE MISSOURI BOTANICAL GARDEN

of these

It seems auite likely that Hebeloma hor

from P

ifi

P. dura), and

ir imperfect system of Agaric classification necessitates its ref-
ence to the former genus.

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

3.

1873

Peck, Bui. Buffalo Soc. Nat. Sci. 1: 50.

PL 10.

Pileus 2.5-7.5 cm. broad, broadly convex or nearly plane, rarely

somewhat umbo-

nate, cream-buff to
clay-color or buck-
thorn-brown when
young and moist,
buff-yellow or Na-
ples-yellow when
old, the center fre-
quently somewhat
darker than the
margin, not strong-

hygrophanous
but changing color
from young to old
condition, glab-
rous, clay-color to
tawny in herbar-
ium specimens,
often rugosely re-
ticulated or cor-
rugated; margin
even, sometimes

Figs. 9-14. P. Acericola: 9, section of the hymenium,

owimr the neculiar thrpe- or four-nointed evstidia fa and

d

flesh

thin,
and

white, taste
dor farina-

b), and the spores (c), X 550; 10, spores, X 1200 (Over-
holts Herb. 3986); 11, section of the hymenium showing
flask -shaped cystidia (a) (Overholts Herb. 984); 12, iso-
lated cyst idia, X 550 (Overholts Herb. 3953); 13, spores, .
X 1200 (Overholts Herb. 3765); 14, spores, X 1200 (Over- ceous; gills Sinuate-

holts Herb. 3911). adnate or with a

small decurrent tooth, medium-close, 2.6-7 mm. broad, grayish,

1927]

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 or
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 X 5-6 [*; 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 (x 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. The 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 a 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. pi. 645) shows a rosy or
purplish tint to the gills of that species very comparable to the
color of the gills in our specimens. A collection from Colorado

[Vol. 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 a
rotten log and can as well be referred to P. Acericola, although the
spores are a 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 prolific 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 A. placomyces. Specimens growing on the ground are likely
to be mistaken for P. praecox which is not usually a woodland
plant. The 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, Bui. Buffalo Soc. Nat. Sci. 4: 53.

1873.

Pileus 2.5-8 cm. 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-

19271

OVERHOLTS

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
times with white mycelial
cords at the base; spores

cm

4-7 mm. thick, some

oid or ovoid

trun

cate at the apex, smooth,
dilute brown, 8-10x4.5-5.5ea;
cystidia present but quite rare,
broadly flask-shaped, project-

=" *\ ' . Figs. 15-16. P. Howeana: 15, section of

Habitat: on the ground m hymenium, showing a cystidium (a), and

woods and bushy places, often J^™ *«£•»■ 8 »° re8 ' X 1200 ' AU
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. sphaler omorpha , and
Patouillard shows that species with a rosy or purplish tint to the
erills as would be indicated for Peck's plant by its original in-

the genus Stroph

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.

•pha, as I have remarked in another place, is

P

mo

extremely similar to the terrestrial form of P. Acericola. My 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
renorts the soecies as occurring in open grassy fields, while Peck

woods and

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

112

[Vol. 14

GARDEN

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

5. Pholiota temnophylla (Peck) Sacc. Syll. Fung. 5: 740. 1887.
Agaricus temnophyllus Peck, Rept. N. Y. State Cab. 23: 90.

1873.

Pileus 2.5-5 cm. broad, hemisph

becoming

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,

Figs. 17-19. P. temnophylla: 17, sections taken at i i i 11 r_in

different places along hvmenium, showing various gl<*Dr0US, UOllOW, O— 1U

types of cystidia (a-d) X .550; 18, spores, X 550; cm . long, 4-8 mm.
19, spores, X 1200. All from type specimens. ,1 • , ... .

thick ; spores elliptic
to ovoid, one end usually slightly truncate, smooth, 9-10.5 X

ent but rather rare, flask-shaped or fusoid,
diameter, projecting prominently.

7

t*

)

dia

hyal

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

Harper, Trans. Wis. Acad. Sci. 17: pi. 83, A

dently

I have been unable to detect this species in my own co
grounds or in the herbarium material examined. It is ev
too close to P. praecox to be recognized from dried plants
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 late]
examination showed their presence, but they are not abundant

Harper's specimens were identified by

The obliquely and

character

proximal edge of the gills is stressed as an im

1927

OVERHOLTS — THE GENUS PHOLIOTA IN THE UNITED STATES 113

B. Cystidia absent from the hymenium; largest plants not more than 4 cm- broad

long

a. On wood.

and P. marginella).

b. On the ground.

Fries, Syst. Myc

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

side; stem

annulus, often Striate on the Figs. 20-26. P. blattaria: 20, section of

hymenium showing basidia, X 550; 21,
spores, X 550; 22, spores, X 1200 (from

or tapering upward, covered specimens at New York, collected on

Long Island by Dodge and Seaver); 23,
section of hymenium of No. 967, X 550;
24, spores of same, X 550; 25-26, various
types of spores from same collection, X550.

small white fibrils either
entirely or only at the base,
whitish or slightly brownish,
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 [>.;

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 p 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 p, long
as in P. toaularis Bull, (non Fries) . The only specimens of this

1 1 A lV ° L - 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. S i ores 9-11 n long.

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

1872.

Agaricus togularis Bull. Herb. Fr. pi 595, f. 2. 1793.
Pileus 1-4 cm. broad, campanulate or convex, becoming

PI. 15.

r^

brown when fresh, hygrophanous, becoming och-
— -v r\ r\ raceous on drying, warm-buff

£ — -\^~) ( N-4» 8 or ochrace °us-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,
Fi K s. 27 30. P. togularis: 27, section yellow becoming pale ferrugin-

of hymenium showing basidia, X 550; ous f tpn pV,rflpprm« huff ™
28, spores, X 550; 29, spores, X 1200! ' OIien OCUraceoUS-DUtt in

All from collection in New York State dried specimens; veil formine

Museum, from North Elba, New York. • . . ,

30, spores, x 1200, from collection at a conspicuous, persistent,

by 3 Murrili. fr ° m FallS ChUrd1 ' Virgink ' 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 cm. long, 2-5 mm. thick;
spores narrow-ovoid or ovoid, truncate at the apex, smooth,
9-10.5 X 4-6 [x; 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. pi. 595, f. 2; Boudier, Ic. Myc.
pi. 101; Cooke, 111. Brit. Fung. pi. 350 (379); Gillet, Champ. Fr.

taria).

Harper, Trans. Wis. Acad. Sci

seems to be well separated from P. rugosa by
throughout and the ochraceous or buff colon

drying. As 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.

type specimens.

Pileus 8-25 mm. broad, broadly conic or campanulate to c
or plane, sometimes umbonate, yellowish-red or dark ferrug
cinnamon or tawny in herbarium specimens, hygroph
glabrous, slightly rugose in the
center; margin striate, often up-
turned in age; flesh very thin, fl^Jf\
concolorous, odor and taste not \j \^L ( J
characteristic; gills at first ad- (^ )*&
nate, becoming adnexed or free
with age, medium-close, usually
minutely denticulate on the edge, ,

J . Figs. 31-33. P. rugosa: 31, section of

1-3 mm. broad, yellowish White, hymenium showing basidia; 32, spores,

becoming ferruginous or brownish- ^JSrSST 1 X im M fr ° m
ferruginous, amber-brown in dried
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 [i; 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 ferruginous and remains so
in 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. Myc. 1: 246. 1821.

[Vol. 14

116 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Pileus 0.5-2 cm. 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,
pAjO odor none, taste mild or sub-farinaceous;

i5 g ^ s a(mate 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-
spicuous, nearly median, membranous an-
nulus, often striate on the upper side ; stem
central, equal or tapering upward, slender,
pallid to brown, glabrous or nearly so, hol-
All from Oyerholts Herb- low, 4-10 cm. long, 1-3 mm. thick; spores

anum No. 2380. . „ . , ' *

ovoid or broadly ovoid, the apex truncate,
smooth, 9-11.5 x 6-7.5 y.; 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. Myc. pi. 102; Cooke, 111. Brit. Fung.
pi 508 (405) B.

Figs. 34-30. P. my-
cenoides: 34, section of

Api

The very slender plants, with

long stem, narrow pileus, median annulus, and the truncate spores
point to a relationship with the P. togularis group. The 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 elliptic without sign of truncate

apex.

144. 1908.

(Fries) Peck, Bui. N. Y. State Mus. 122

Pholiota togularis fdaris Fries, Ic. Hym. 2: 2. 1877.

1927]

OVERHOLTS — THE

117

Pileus 0.5-2 cm. 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 a distinct,
persistent, median or su-

perior though distant
nulus, striate on the

Figs. 37-39. P. filaris: 37, section of hymen-

•j ^+«^ ™,,+™l 3!), spores, X 1200. All from collection at

per Side; Stem central, N ' York from WasWton. bv Murrill.

labrous 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 X
4.5-5 [l; cystidia none.

Habitat : on the ground in woods or pastures.

Distribution: specimens have been examined from Catskill
Mts., N. Y.; Washington (s';ate).

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 \l long. However, it
may intergrade too much into P. togularis, though the habit
seems distinct enough. I have followed Peck in recognizing the

species.

//. Spores not with truncate apex.

A. Growing on the ground or among moss (see also B, p. 182).

1. Plants with large spores 10-15 fx 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) cm. 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-

118

(Vol. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

distant, thin, at the very first pallid, soon 'clay color' to 'buck

brown' (Ridg

Stem 3-5 (6) cm. lone, 2^4.5 mm. thick

slightly tapering upwards, subflexuous in age, stuffed,
•ally 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

Figs. 40-42. P. platyphylla: 40, section of hymenium, with °Ut Under the veil

a cystidium (a) and spores (l>), X 550; 41, a single remnants, fuscescent.
cystidium, X 550; 42, spores, X 1200. One spore c ' . „. . ,

showing the faint minute roughness. All from type spores SUbellipsoia,

specimen8 " but narrower toward

one end, inequilateral in one view, smooth, 10-12 x 5-6.5 n,

9

de rusty brown, cystidia scattered on sides of gills, 60-70

-11 [i, 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.

Syst. Myc

1821.

PI. 11.

Pileus 1-5 cm. 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 olants : veil

1927]

OVERHOLTS — THE GENUS PHOLIOTA IN THE UNITED STATES 119

spores, X 1200 (both from type specimens of P.
aggericolo) ; 45, isolated cystidia from collection at New-
York fro ii Bronx Park by Murrill, X 550.

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-
tic or elongate-ovoid,
smooth, 11-15 X 5.5-
7.5 [i; cystidia pres-
ent, usually abun-
dant, hyaline, 7-12 [i
broad at the enlarged
base, projecting 30-
40 n.

Habitat: on the Yigs. 43-45. P. erebia: 43, section of hymenium
ground by roadsides snowm g a cystidium (a), and spores (b), X 550; 44,

and in woods.

Distribution: spec-
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, and 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

[Vol. 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 on the margin where a 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-
ior membranous annulus;

Figs, 46-47. P. siibnigra: 46 section of gtem cent ral, equal, pallid,
hymenium showing cystidia (a), and spores (b), . ,

X 550; 47, spores, X 1200. All from type speci- rough with short Soft

mens ' whitish conic scales point-

ing upward, solid, 2 cm. long, 2.5 mm. thick; spores somewhat
elongate-ellipsoid, smooth, 10-13 X 5-6 p; cystidia abundant,
projecting, conspicuous, flask-shaped.

Habitat: attached to a small buried root.

Distribution : known only from the type locality, Seattle, Wash,

b. Plants 4-15 an. broad at maturity,
aa. Spores smooth (see also p. 123).

14. Pholiota ombrophila Fries, Hym. Eur. 216. 1874.

Pileus 3-10 cm. 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 a few white
floccose fibers or scales on the margin, soon glabrous, hygrophan-
ous or sub vise id 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 IN THE UNITED STATES 121

enlarged below, whitish or brownish, often dark below, stuffed
or hollow, 4-8 cm. long, 4-15 mm. thick; spores cylindric-elliptic
or fusoid-elliptic,
smooth, 11-15 X 5-
7 pi ; cystidia abun-
dant, conspicuous,
projecting 20-40 p,
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.

Illustrations : Fries,
Ic. Hym. pi 103, f. 2;
Harper, Trans. Wis.
Acad. Sci. 17: pi. 81.

QJ\

v

Figs. 48-53. P. ombrophila: 48 ; section of hymenium
This species repre- showing cystidia, X 550; 49, spores, X 550; 50, spores,

X 1200 All from specimens at New York from Bresa-

51, section of hymenium showing cystidia, X

All from

sents the height of dola.

development in a p); 52 s P ores x 550; 53 spores, x 1200.

„ ^ , type specimens of P. washingtonensis.

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 otiers 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 (x long) are correct, his plants cannot belong to this species.

15. Pholiota aurea (Mattuschka) Fries, Syst. Myc. 1: 241.

1821.

PL 12.

[Vol. 14

122 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Agaricm aureus Mattuschka, En. Stirp. Sil. Herbor. 351. 1779.
Pileus fleshy, 10-15 cm. broad, convex to plane, often upturned
In 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,
compact, yellowish; gills rounded at
the base, rather close, 0.7-2 cm.
broad, ochraceous-buff to ochrace-
ous-tawny; veil forming a firm,
large, pendulous annulus, lemon-
color on the upper side, dark buff
Figs. 54-56. P. aurea: 54, section beneath, eventually disappearing;

MO; 3 «fSSl. d 7«8& i ■#£;„£ stem central, tapering upward from

x 1200. All from Overholts Herb. a n 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 cm. long, 2-3 cm. thick, inserted 2-5 cm.
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 X 4-6 [a; 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. At 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. Odeli of the Dominion Experimental Farms, Ottawa,
Canada.

bb. Spores rough.

16. Pholiota caperata (Per 3.) Fries, Syst. Myc. 1:241. 1821.

PL 13.

Agaricus caperatus Persoon, Syn. Fung. 273. 1801.

Pileus 5-15 cm. broad, hemispheric or ovate, then campanulate
or convex to nearly plane, o 3hraceous-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
a 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 corrugate; margin even, at first
incurved; flesh white, taste mild;

gills sinuate-adnate or adnate to
slightly decurrent, medium-close,
5-15 mm. broad, at first whitish,
then clay-color, finally ferrugin-
ous, becoming cinnamon in dried
plants, usually variegated with

close, narrow, light and dark

Figs. 57-58. P. caperata: 57, section
of hymenium showing basidia and

cross-bands and appearing trans- spores, x 550; 58, spores, x 1200. All

versely wrinkled, the edge floe- ^om Overholts Herb. No. 5241.

cose-crenate; veil forming a persistent, superior or almost
median, white, deflexed, membranous 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 cm. long,
7-25 mm. thick; spores b::oadly and inequilaterally elliptic,
echinulate, 11-19 X 7-11 \x; cystidia none.

Habitat : on the ground in woods.

Distribution : specimens ha ve been examined from Bar Harbor,
Me.; Ontario, Canada; Stow, Mass.; Chappaqua, Adirondack
Mts., West Ft. Ann, and Gansevoort, N. Y. ; Lamar, Stone Valley,

[Vol. 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. pi. 48; Fl. Dan. pi. 1675;
Gillet, Champ. Fr. pi. 520 (518) ; Harper, Trans. Wis. Acad. Sci.
17: pi. 24; Hard, Mushrooms,/. 212; Peck, Rept. N. Y. State Bot.
54: pi. 73, f. 1-5; Ricken, Blatterp. pi. 55, f. 2.

Among the true Pholiotas this species is a 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, elliptic 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
in which the enclosing fibers are too scanty to leave such a 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 4r-8 cm. 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) [x; 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 annuus is of a more delicate texture and
more evanescent.

2. Plants with small spores up to 9 n long,
a. Spores rough-walled (see also p. 126).

18. Pholiota minima Peck, Rept. N. Y. State Mus. 41: 65

1888.

Pileus 3-8 mm. broad, henispheric or campanulate, umbonate

brown when moist, nale buff

09c

o

lowish- white when dry, brown in /*3K5» f\

herbarium specimens, hygrop
glabrous, margin striatulate when
moist; gills adnexed, subdistant, fer-
ruginous, cinnamon in dried plants,

1-2 mm. broad; veil forming an FigSi 59 _ 61 . P . minima : 59,

median or superior an- *? cti °ii of hymenium showing

r dia, X 550; 60, spores, X 550; 61,

central, equal, concolor- spores, x 1200. All from type

stem

ous with the pileus, shining, glab- *"**"»•

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 [&; cystidia none.
Habitat : among hair-cap mosses.
Distribution : specimens hs/ve been examined from the Catskill

Mts., N. Y.; Auburn, Ala.

The 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 cm. broad, convex or plane, deep golden- or coffee-

[Vol. 14

126 ANNALS OF THE MISSOURI BOTANICAL GARDEN

lmens.

brown to dark melleous, entirely covered with matted floccose

fibrils of the same color or these
collected into squamules particu-
larly at the center of the pileus,
dry; gills adnate to adnexed, deep
brown, medium-close, 3-6 mm. broad;
veil forming a persistent or evanes-
Figs. 62-04. P. irachyspom: 62, cent, superior annulus ; stem central,

section of hymenium showing basi- 1 £i •« 1 t A

dia, x 550; 63, spores, x 550; 64, e <l ual > fibrillose, umber-brown, 4-8

?E!Y3 x 12(X)- AU f rom type spec " cm - lon g> 4-10 mm - thick; spores

broadly ovoid or subglobose, dark

under the microscope, decidedly rough- walled, 7-9 X 5-6 nj

cystidia none.

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 (I. c.) 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 photograj >h 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 cm. in diameter; from the Pacific Coast only (see also

p. 128).

20. Pholiota terrestris Overholts, N. Am. Fl. 10: 268. 1924.

Pileus 1-3 cm. 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

127

the stem, 3-6 cm. long;, 2-5 mm. thick; spores oblong-ellipsoid

smooth

•6.5 X 3.5-4

cystidia of several inconspicuous

types, none projecting strongly, some

brown

ith a 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 Figg ^^ p terrestris . 65>

These lat- section of hymenium showing

basidia and several types of
inconspicuous cystidia, X 550;

with a slightly projecting tip and a dis- Mjjpojj ^M^

tinct ball-like content that h as a strong

ystidia more

fusoid

affinity for the eosin stain, so that

Herbarium.

left in it for half an hour show these sterile structures

tremely

P

of Europe is similar, but accordin

to specimens from Bresadola has spores 8-9 u, long and entir
lacks cystidia. Harper's plants referred to P. terrigena seem
belong here and he gives the smaller spore record of this speci

21. Pholiota anomala Peck, Bui. Torr. Bot. Club 22: 202.

1895.

5-2.5 cm. broad, at first hemisph

sub

then convex, broccoli-brown when moist, pale yellow or cream-
when dry, warm-buff in dried
5, hygrophanous, glabrous ; gills
or decurrent, medium-close
tlv distant, 3-4 mm. broad,

adnate

becoming brownish

ochraceous
dried plan

to cinnamon in

formin

slight,

fibrillose or glabrous

Figs. 68-70. P. anomala: 68, finally evanescent, annulus; stem

section of hymenium with a single __„ trQ i pmin l

narrow cystidium-like body, X 550; Central, equal,

69, spores, x 550; 70, spores, x whitish or brownish, hollow with

I 200 -^ A1 L ^.^^i^Ti!! i«**ni1*r transverse nartitions or the

York.

filled with a cottony tomen

turn, 3-6 cm. long, 2-6 mm. thick; spores oblong-ellipsoid, smooth

128

[Vol 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

very dilutely colored, 7.5-10 X 4-5 \l; 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 (x beyond
the basidia, 40-50 X 5-7 (a.

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 nearlv or auite crone.

annulus is nearly or quite gone. Sections of
the stem show the peculiar-chambered internal structure in good
shape. The 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. The species has not
been recognized since its discovery and its affinities are in doubt.

bb. Mature plants more than S cm. in diameter.

22.

albivelata Murrill, Mycologia

1912.

Pileus 2-6 cm. broad, convex to plane, sometimes somewhat
umbonate, isabelline tinted with rose, resembling the color of some

species 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
tv ».««.,,.,, - . pendant, superior or median,

Figs. 71-73. P. albivelata: 71, section of . . f ,

hymenium showing different types of cystidia persistent annulus, pure

(a-e), and spores (f), X 550; 72, spores, X white
1200; 73, isolated cystidia showing the

rounded body near the apex, X 550. All Upper side brown from the
from Overholts Herb. No. 6027.

the lower side

and

from the

the

heavily white-floccose just be-

floccose above, becoming

subglabrous and rarely yellowish toward the base

1927]

OVERHOLTS — THE GENUS PHOLIOTA IN THE UNITED STATES 129

5-8 cm. lone, 4^10 mm. thick; spores ovoid to narrow

soid, smooth, slightly

end. 7.5-9 X 4.5-5.5

ystidia abundant, not projecting conspicuously, small, hyal

with a rounded conspicuous dark

body in the apex,

sometimes pointed, but more often obtuse, 30-35 X 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. The cystidia with their colored
contents are different from t hose 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, Bui. N. Y. State Mus. 122: 148.

1908.
Pileus 2.5-9 cm. 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-squamos'3

or

with

ot-like scales or d(
the center, margin <

flesh white
nexed or

mild

ad

sinuate-adnate, some-
times with a decurrent tooth,

close, narrow, 2-5 mm. broad,
whitish becoming brown or i usty-
brown, with a white-crerulate

dge

cinnamon-buff or imuff

brown when mature; veil forming
a superior, white, membranous,

74-76. P. duroides: 74, section

pendant or rolled

, • , , ui mmsiiiuui ^,,^15 the inconspicu-

bpersistent OU8 cystidia (a), and spores (b), X 550;

«rmiilii<3 nftpn (tfriftt.p on thp iin- 75 > spores, x 1200. All from type

annulus. oiten striate on tne up 8pecimen8 76( ^^^ of hymenium as

or in 75, X 550. From Overholts Herb.
No. 7479.

r side; stem central, eqtal
larged below, glabrous, or at

times fibrillose-scaly below, whitish, stuffed or hollow, 3

cm

[Vol. 14

130 ANNALS OF THE MISSOURI BOTANICAL GARDEN

long, 4-15 mm. thick; spores ellipsoid or ovoid, smooth, 4-6 X
3-4.5 [i ; eystidia present, obtuse, not projecting strongly yet

rather conspicuous in point of size, often mucronate-tipped,
8-11 (j. in 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 Botanical 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 R. 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. The 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

justified at present.

Harper describes (Trans. Wis. Acad. Sci. 17: 476. pi 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

1872.

Pileus 3-10 cm. 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

£>

OQo

OOO

-°<p

4£0&

brow

ff

brown to bister in
dried specimens; veil

formi

a

th

Figs. 77-82. P. Johnsoniana: 77, section of hymen-

ium showing basidia, X 550; 78, section of hymenium

white, persistent an- showing inconspicuous cystidium (a), X 550; 79,

spores, X 550; 80, spores, X 1200. > All from type

specimens. 81, section of hymenium with cystidia (a),

labroUS, light- and spores (b), X 550; 82, spores, X 1200. All from

Overholts Herb. No. 3946.

nulus; stem central,

colored, solid

the top, 7

cm

5 cm. thick ; spores ovoid

or ellipsoid, smooth, 5-8.5 X 3-4 (i; 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 Brmx Park, N. Y.; State College, Pa.;
also reported from North Carolina by Atkinson and from Michi-
gan by Kauffman.

Illustrations: Atkinson, Mushrooms, pi. 49 {H\f- H9 {H^);
Murrill, Mycologia 7: pi. 138, f. 10; Peck, Rept. N. Y. State

Cab

f.4

Atkinson and Kauffman stress the turbinate shape of the
as a character for recognizing the species. In the one coll
of fresh plants I have seen this character was not partk
pronounced, and in the type specimens is not evident ir
present condition although s hown in two of the three illusti

132 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

published by Peck. The species has some of the aspects of P.
praecox, 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 y. long; those of P. Johnsoniana vary
between 5 and 8.5 p. and are slightly more irregular in shape. The
cystidia of P. duroides, types, are not conspicuous but are present
as slightly projecting mucronate-tipped organs much larger than
the basidia. 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. IHleus not distinctly scaly (see also 2, p. 142).

a. Spores rough-walled, prominent flask-shaped cystidia present in the

hymenium (fig. 83) (see also b, p. 137).

25. Pholiota marginata (Batsch) Fries, Epicr. Syst. Myc. 169.
1836-38. PL 14.

Agaricus marginatus Batsch, Elench. Fung. 65. 1783.

Pileus 1-7 (-8) cm. broad, convex to plane or slightly depressed,
sometimes somewhat umbilicate, more rarely slightly umbonate,

1927]

OVERHOLTS

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, hygiophanous, glabrous; margin usually
striatulate when moist, often extending somewhat beyond the
gills; context fleshy, thin, concolorous with the pileus, taste and
odor farinaceous
uate-adnate to

to adnate or
slightly decurrent, me-
dium-close to slightly dis-
tant. 2-7 mm. broad, light

ish-brown

buck

brown; veil forming
tcious or subpersist-

stem

somewhat
the apex,
>se at the

fibrillose, pruinose
top and with a white
mentum at the base, <
colorous or lighte

than

Figs. 83-86. P. marginata: 83, section of hy-
the pileUS, hollow, 2-8 cm. menium showing cystidia (a), and spores (b),

** « / +*\ j.1.- i X 550 (from Overholts Herb. No. 1562); 84,

long, 2-0 (-10) mm. thick; „p res, X 1200; 85, section of hymenium with

cystidium, X 550; 86, spores, X 1200. All from
;ype specimens of P. autumnalis.

spores elliptic or ovoid,

often smooth when young,

with a rough wall when ma :ure, 7-10 X 4.5-6 \

but usually not abundant, flask-shaped with

ystidia

that

hyaline, 50-80

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, 111.; Creve Coeur Lake, Wicks,
Meramec Highlands, and St. Louis, Mo.; Minnesota; Vancouver
Island; also reported from Michigan by Kauffman.

Illustrations: Atkinson, Mushrooms, /. 143 (147); Cooke, 111.
Brit. Fungi, pi 372 (403), Hard, Mushrooms, /. 215; Harper,
Trans. Wis. Acad. Sci. 17: pi 54, 55; Ricken, Blatterp. pi. 56, f. 7.

134

[Vol. 14

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 enoutih constant to

enable one to distinguish the

>y

from the other. On the Pacific

Also

there is an apparently common plant which dries a bright

has a persistent, well-developed, superior membranous an-

and the spores are less roughened or altogether smooth.

scarcely more than 1-2 mm. thick.
micolor. The same type is also found

stem is slender

This plant I

East where these characters seem to be less constant but

more 1

distinguishable

P. maroinata the stem

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
a synonym.

26. Pholiota unicolor (Vahl) Fries, Epicr
Syst. Myc. 170. 1836-38.

Agaricus unicolor Vahl, Fl. Dan. 6 18 : 7
1792.

Pileus 0.5-2 cm. broad

cam

to

campanulate or somewhat con-
vex, often sharply umbonate, cinnamon-
buff to ochraceous-buff or ochraceous-
orange, herbarium specimens cinnamon
to ochraceous-tawny, glabrous, hygro-

dn striate or fluted: flesh

mar

Figs. 87-88. P. unicolor:
87, section of hymenium ph
showing a cystidium (a), and ±u;
spores (b), X 550; 88, spores,
X 1200. All from specimens
at New York, from Washing- ,

ton, collected by Murriii. somewhat free with the expansion of

the

gills squarely adnate, often becoming

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 lloccose-mealy above the annulus,
fibrillose below or the base white-tomentose, hollow, 2.5-5 cm.
long, 2-4 mm. thick; spores ovoid or elliptic, smooth or slightly
rough when mature, 7.5-10 X 5-6 [x; 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.; Bronx 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. pi. 530, f. 2 (as Ag. xylophilus) ;
Cooke, III. Brit. Fungi, pi. 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, Bui. Buffalo Soc. Nat. Sci. 1: 50.

1873.

PI. 15.

Pileus 1-3 (-5) cm. broad, convex then expanded or slightly

depressed, cinnamon-rufous,
bright ochraceous-yellow when
dry, cinnamon-buff, ochraceou 3-
orange, or cinnamon in dried
plants, hygrophanous, smooth,
viscid, margin striatulate when
moist, even when dry; gills ad-
nate or with a decurrent tooth,
medium-close or somewhat di 3-
tant, 1-3 mm. broad, pallid then

pale ferruginous, mikado-brown JSBStLS.'StSfirS: X
or russet in dried plants; veil

forming a subpersistent, dis-
tinct annulus; stem central, terete, equal, pallid, fibrillose-striate,
hollow, 2.5-8 cm. long, 1.5-b mm. thick; spores elliptic, rough,

550; 90, spores, X 550; 91, spores X 1200.
All from type specimens.

[Vol. 14

136

ANNALS OF THE MISSOURI BOTANICAL GARDEN

dilute brown under the microscope, 7-9 X 4-6 (x ; cystidia present
but sometimes rare, projecting 20-40 y. or more, flask-shaped,
40-70 X 10-12 j*.

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: pi. 61 B.
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 a de-
cidedly darker color to the edge of the basidial layer as seen under
the microscope. I 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.

Figs. 92-93. P. furcata:
92, section of hymenium
showing a cystidium (a), and
spores (l>), X 550; 93, spores,
X 1200. All from type speci-
mens.

28. Pholiota furcata Overholts, N. Am.
Fl. 10: 272. 1924.

Pileus 1-3 cm. broad, convex to plane,
somewhat gibbous at times, reddish-
brown when moist, ochraceous when dry,
cinnamon-buff in dried plants, hygro-
phanous, glabrous; margin even, at first
incurved; flesh concolorous, odor none;
gills slightly decurrent, close, ochraceous
to dull cinnamon, 2-3 mm. broad, con-

1927J

OVERHOLTS — THE GENUS PHOLIOTA IN THE UNITED STATES 137

spicuously forked and connected by veins so as to appear some-
what poroid; veil forming a superior, distinct but somewhat evan-
escent annulus; stem central, equal, dark watery-brown, floccose-
pruinose above the annulus, v rhite-fibrillose below, solid or spongy,
1.5-2.5 cm. long, 2-3 mm. thick; spores ovoid or ellipsoid, some-
what rough at maturity, 7.5-9.5 X 4.5-6 &; cystidia present and
fairly abundant, hyaline, flask-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 alio c, p. 141)-

29. Pholiota marginella Peck, Rept. N. Y. State Mus. 51: 289.
1898.

Pileus 1^1 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-devc sloped fuga- 94 > 8ection of , v h y™ en ^™

, , , , , /m m s howm g spores (a), X 550;

cious annulus; stem central, squal, fibril- 95, spores, x 1200. All from
lose below, pruinose above the annulus, type 8 P ecimens -
stuffed or hollow, whitish or pallid, sometimes with a white
tomentum at the base, 3-10 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 ^; cystidia none.

Habitat: decaying wood 01 on sawdust piles.

[Vol. 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, I 'a.; Oxford, Ohio; Tolland, Colo; Seattle, Wash.; Yaku-
tak Bay, Alaska.

Illustrations: Peck, Rept. N. Y. State Mus. 51: pZ. B,f. 12-20.

Though apt to be confused on external appearances with P.
margin (da 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. A favorite haunt
for it is an old sawdust pile, and most of the collections examined
have been from such a habitat. On 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. I found it to be
the most abundant species in the vicinity of Tolland, Colorado,
in 1913 and 1914, and at least a dozen collections are preserved
in my herbarium from that locality.

30. Pholiota mutabilis (Schaeff.) Fries, Epicr. Syst. Myc. 169.

1 836-38.

CxQ

Figs. 96-97. P. mutabilis:
96, section of hymenium with
spores (a), X 550; 97, spores,
X 1200. All from speci-
mens at New York from
Bresadola.

Agaricus mutabilis Schaeff. Fung. Bavar.
4: Ind. 6- 1774.

Pileus 1.5-3 cm. broad, convex to plane,
cinnamon when moist,paler when dry,
ochraceous-buff in dried plants, hygro-
phanous, glabrous; gills adnate or slightly
decurrent, medium-close, 2-4 mm. broad,
pallid then cinnamon; veil forming a
white or dark, superior, evanescent or
persistent annulus; stem central, equal,
concolorous with the pileus, decidedly
scaly below the ring, pruinose above,
stuffed then hollow, 3-7 cm. long, 3-5
mm. thick; spores ovoid or elliptic, slightly
truncate at one end, smooth, 6-7.5 X4-5 \l;
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. pi. 38, f. 208; Bresadola,
Fung. Mang. pi 51; Cooke, 111. Brit. Fungi, pi 355 (402); Fries,
Sverig. Atl. Svamp. pi. 47; Lanzi, Fung. Roma, pi 76, f. 3;
Schaeffer, Ic. Fung. pi. 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

em is scalv with scattered

the annulus, in which character it is distinct from P. m

31. Pholiota Aegerita (Brig.) Fries, Epicr. Syst. Myc. 164.
1836-38.

Agaricus Aegerita Brig. Funghi Litogr. Napol. pi. 1. 1824.

Pileus 3-13 cm. broad, si b-hemispheric 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-clos e, 4-8
mm. broad, dark brown; veil form-
ing a median-superior, persistent,
conspicuous annulus; stem central,
nearly equal, brownish, more or less
white-fibrillose, solid, 4-15 cm. long,
4-10 mm. thick; spores ovoid or
more often elongate-ovoid or elon-
gate-elliptic, smooth, 9-11 X 4.5-6 p,; cystidia present but not
conspicuous, projecting somewhat, hyaline, 6-10 (x in diameter,
some pointed at the apex.

Habitat: on recently felled trunks or from wounds in deciduous
trees.

Figs. 98-99

P.

Aegerita: 98,
section of hymenium showing the

inconspicuous cystidia (a-c), and
spores (d), X 550; 99, spores, X
1200. AH from specimens at New
York from Portugal by Bresadola
and Torrend.

[Vol. 14

140

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Distribution: Michigan.

Illustrations: Bresadola, Fungi Mang. pi. 50; Briganti, Hist.
Fung. Nap. pi. 32; Cooke, 111. Brit. Fung. pi. 365, 453.

I have seen no American specimens of this species. It is re-
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. The colors are similar to P.
praecox but a deeper tan and the pileus may become areolate,
particularly in 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

marked

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. The cystidia seem to present a well
characteristic, 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
agreeable nutty or amygdaline taste in
dried plants ; gills adnate, medium-distant
,„ %M D to distant, yellow or yellowish-brown,

Figs. 100-101. P. Oregon- . \ , . . . , ,

ense: loo, section of hymen- becoming darker, strongly mterveined,
(b), x 550; 101, spores, x perior or nearly apical, irregular, yellow-

1200. All from type speci- . , , ., , , , ,

lsn-white annulus ; stem central or excen-

su-

mens.

1927]

OVERHOLTS — THE GENUS PHOLIOTA IN THE UNITED STATES 141

trie, terete or compressed, equal or enlarged upward or down-
ward, yellowish above, fulvous below, with small scattered uni-
colorous subfloccose evanescent scales pointing upward, solid,
6-10 cm. long, 8-20 mm. th.ck; spores ovoid or elliptic, smooth,
7.5-10 X 3-5 (x; 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. The gills 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 as to the status o: the species.

c. Spores rough-walled; no cystidia in the hymenium.

33. Pholiota cerasina Peck, Bui. Buffalo Soc. Nat. Sci. 1: 50.

1873.

Plants cespitose, 5-12 cm. broad, convex to plane, cinnamon-
color to tawny, perhaps light er at times, dry or somewhat hygro-
phanous, glabrous or nearly e o ; margin even ; context fleshy , rather
thin, bitter to the taste, wi;h an amygdaline odor that is best
noticed in young plants; gills
medium-close or slightly distant,
sinuate to adnate or slightly de-
current, yellow, becoming cinna-
mon or ferruginous, finally prui-
nose from the spores, 5-12 mm.
broad; veil present, forming an Figg lQ2 _ m p ^^ ^

early evanescent, spore-Stained section of hymenium with spores (a),

annulus; stem central or more $"%%!£?* * 12 °°' A " fr ° m
often excentric, 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

[Vol. 14

142 ANNALS OF THE MISSOURI BOTANICAL GARDEN

to ovoid, slightly roughened, brown, 6-9 X 4.5-5.5 [x; 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.

The 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. If there is a 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 cm. 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 cm. long, 4-7 mm. thick; spores ovoid or elliptic,
8-9.5 X 4-6 [i, 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 a water-color sketch by Miss
Banning. The sketch shows a pileus orange-cinnamon, ochre-
red, or tawny, and gills and stem similar but lighter. The 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, among which its affinities are in doubt.

2. Pileus distinctly scaly at maturity.

a. Spores rough-walled (see also 6, p. 1^8).

1927]

OVERHOLTS — THE GENUS PHOLIOTA IN THE UNITED STATES 143

35. Pholiota luteofolia Peck, Rept. N. Y. State Mus. 27: 94.

1875.

cm. broad, convex, very young specimens dark red

or reddish-brown, becoming pinkish-

red or

red whe n mature

buff or ochrac(

m

dried

dry, appressed fibril

squamulose and sometimes

late in the center, fibrillcse on the

mar

fleshy, thii

lavender in fresh

I

gills

adnate or uncinate, sometimes becom

ing

adnate, mediu

or

somewhat distant, 3-8 mm. broad

bri

yellow becoming

mostlv ochraceous-buff or ochraceous

grrugmous

ing a

i dried specimens; ve
ght, fugacious, spore
stem central or somewhat

form

Figs. 104-107. P. luteofolia:

104, section of hymenium with

Stained spores (a), X 550; 105, spores,

X 1200. All from Overholts
Herb. No. 3741.

centric, equal

ed downward

with the nileus, fibrillose

ran

106, section of
hymenium, showing cystidium
(a), and spores (b), X 550;
107, spores, X 1200. All from
type specimens.

;htly

gh

5-4.5

solid, 3-9 cm. long, 3-10
spores ellipsoid to elliptic,
cystidia none or not noteworthy.

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: pi. 1+8.

This species is most closel y 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

identification of dried

P. luteofolia seems

to lack the green hue of P. aeruginosa and does not become
rimose-areolate and the scales appear to be more innate. It i:
smaller plant than P. speck bilis , and the gills are much bright

144 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

colored in dried plants. From P. cerasina it differs in the dis-
tinctly squamulose pileus. The color of mature plants is typic-
ally a 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.

Figs
bilis: 108, section of hy

36. Pholiota spectabilis Fries, Elench. Fung. 28. 1828.

PL 16, 17.
Pileus 4-15 cm. broad, convex becoming nearly plane, buff-
llow 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
young plants practically glabrous at
times, to distinctly fibrillose or rivulose
or in mature plants squamulose; margin
even; context yellow, taste bitter or
amygdaline; gills adnexed to adnate or
with decurrent teeth or lines, medium-
close, 3-8 mm. broad, yellow becoming
ferruginous, yellow-ochre to ochraceous-
orange or tawny in dried specimens; veil
menium with spores (a), x forming a distinct, superior or apical,

550; 109, spores, X 1200. , . , .' , r , . . J

All from type specimens of spore-stained, persistent or subpersistent
?o™ ule % 1I0 ' 8p . ores ' * annulus, sometimes striate on the upper

1200. From specimen at ^^

New York from Bresadola side; stem central or nearly so, nearly
from Holland. equal tQ decidedly ven tricose or bulbous-

enlarged at the base, yellow or tawny, yellow and floccose above
the annulus, fibrillose or furfuraceous below, 3-15 cm. long, 0.5-3
cm. thick, solid; spores elliptic, rough, 7-9 X 4.5-6 [>.; 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, N. J.;
Frederick, Md.; Washington, D. C; Montgomery and Bir-
mingham, Ala.; Clary ville, 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, Or 3.; Seattle, Wash.; also reported by
Kauffman from Michigan.

Illustrations: Bernard, Champ. Roch. pi. 55, f. 1; Cooke, 111.
Brit. Fung, pi. 352 (394); Fr.es, Ic. Hym. pi. 102; Gillet, Champ.
Fr. pi 527 (529); Harper, Trans. Wis. Acad. Sci. 17: pi. U, 50;
Kauffman, Agar. Mich. pi. 01; Murrill, Mycologia 1: pi. 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 minute fibrils or fibrillose squamules
covering the entire pileus at maturity, the subpersistent, nearly
apical annulus, and the rou^;h spores 7.5-9 \i 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 distinctly. In other collections the squam-
ules are large enough to be n adily 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 bitter (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.

of still other

Most

curious of all, in one collectiDn 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 bur i ed 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

[Vol. 14

146 ANNALS OF THE MISSOURI BOTANICAL GARDEN

are available. As 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. I be-
lieve, however, that there is no other European illustration that
shows so squamulose a pileus as does Fries'. Cooke's illustration
(pi. 352) approaches it most closely but here the pileus is shown
with appressed rivulose squamules formed by the separation of
the 1< > ng fibers of the cuticle. This type is represented in America,
at least on the Pacific Coast. One 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. It 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. pi. 529), although the other col-
lection agrr<>s with Fries' figure and Cooke's illustration in being
obtuse or enlarged at the base. All 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

fibrillose to squamulose.

be asked, "Does the tvoical P

Eur

be in the affirm

ative, on the basis of an illustration of P. spectabilis published by
Bernard in Champ. Rochelle, pi. 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

147

New York from Alabama by Earle, another from Bronx Park by
Murrill in 1911, and two oi three others, there would be more
basis for keeping it distinct. For this typ

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 fibril: 5 on the pileus and in stature compare

There is a specimen at Albany, from New

birch, that

matches Bresadola

well with our p
Brunswick, on

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.

PL 18.

>

Pileus 2-10 cm. broad, convex, greenish becoming tinged
with yellow or brown, dral
to cinnamon or ochraceous-
buff in dried specimens, dry,
at first glabrous, usually

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 in dried
plants; gills adnate or sinu-
ate-adnate, easily separating ,
3-7 mm. broad, pale-ochra-
ceous when young, becoming
ochraceous-orange or apri-

113, spores, X 1200. All from Overholts
Herb. No. 8081. 114, spores, X 1200 (from
type specimens); 115, spores, X 1200 (from
Overholts Herb. No. 8773, spores probably

less mature).

buff on drying; veil leaving only a slight fibrous, lacerated

stem

[Vol. 14

148 ANNALS OF THE MISSOURI BOTANICAL GARDEN

or nearly so, glabrous or slightly fibrillose, sometimes sulcate-
striate, colored like the pileus, solid, 3-8 cm. long, 4r-10 mm.
thick; spores ellipsoid or elliptic, rarely entirely smooth, typically
slightly echinulate when mature, 6-8 X 3.5-1.5 [i; 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.,
New York. Another Pennsylvania collection is at New York
City, by Mcllvaine. Dr. Murrill has already referred his Flam-
mula viridans from the Pacific Coast to this species. The 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 pulehrifolia, 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 3.5-5.6 n long (see also 2, p. 149).

38. Pholiota flammans (Batsch) Fries, Syst. Myc. 1: 244

1821.

PI

Agaricus flammans Batsch, Elench. Fung. 30. 1783.

1927]

OVERHOLTS — THE GENUS PHOLIOTA IN THE UNITED

149

Pileus 2-5

cm. broad, convex to plane, sometimes um

bonate, lemon-yellow or ta^ roy-yellow, zinc-orange or tawn
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

fug

stem central, equal, with yellow, re-
curved, floccose scales, or scarcely more
than densely yellow-floccose up to the
annulus. stuffed or hollow, yellow, 2-7

cm

Figs. 116-119. P. flammans:
116, section of hymenium show-

long, 2-5 mm. thick; spores ob-
long, smooth, 3-5.5 X 2-3 x; cystidia
abundant, flask-shaped or clavate- m . .,.,,.,

„ . , t if j.' mg cystidia (a), and spores (b),

fusoid, brown or hyaline, projecting x 550; 117, spores, x 1200;1 18,
slightly, 30-40 X 6-12 ji.

Habitat: on dead wood of both de-
ciduous and coniferous trees .

Distribution: specimens have been
examined from Lake Placid, Osceola,
Greig, and Fourth Lake (Herkimer Co

section of hymenium showing
cystidia, and two cystidia iso-
lated, X 550; 119, isolated
cystidia, the one on the right
hyaline, the other two brown, X
550. AH from Overholts Herb.
No. 3753.

N. Y. : Glen Brook

Ore. ; also reported from Mi chigan by Kauffman and by Harp

Illustrations: Fries, Ic. Hym

104, /• 1; Cooke, 111

Fungi, pi. 396 (368): Harper, Trans. Wis. Acad. Sci. 17: pi 41C.

Ricken, Blatterpilz

/

In one collection at New York, from a birch stump, the col-

jtor's notes say "viscid."

P. flammans is distinct from all related species in the very

small, oblong spores, measurin
marked species with bright cc

> (x long. It is indeed a well-
and soft floccose scales, and

while generally considered its occurring only on coniferous wood,
yet a collection at New York is very evidently on the wood of

birch

and I have seen no American

mens more than 4 cm. broad

2. Spores between 9.5 and 15 n long (see also S, p. 152).

150

[Vol. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

39. Pholiota albocrenulata Peck, Bui. Buffalo Soc. Nat. Sci. 1:

49. 1873.

Pileus 2.5-12 cm. broad, conic-campanulate to broadly

PL 19.

or nearly plane, sometimes
umbonate, uniformly yellow-
ish-brown to sayal-brown or
tawny, the darker of these

dried

viscid, with

th

large

scales that are appressed and
sub-gelatinous when wet, but
become suberect, floccose,
and lighter in color on dry-
ing, easily separable and
sometimes disappearing in
old plants; gills sinuate-ad-
nate or slightly decurrent,

medium

or in

m

large
rather distant,
mm. broad, grayish be-
coming ferruginous, the edge

3

d

white

and remaining so at least in
part in dried plants; veil
forming a torn fugacious an-
nulus or partly appendicu-

late to the mar

of the

Figs. 120-126. P. albocrenulata: 120,
Bection of hymenium showing central medulla

(a), X 550; 121, spores, X 550; 122, spores,
X 1200. All from type specimens. 123,
section of hymenium showing the occasional
pointed cystidia, X 550; 124, spores, X 550;
125, spores, X 1200; 126, section of edge of
gill showing the clavate cells that give to the
edge the crenulate appearance, X 550. All
from Overholts Herb. No. 9368.

fusoid

pileus; stem central, equal
or somewhat enlarged below,
sparingly or abundantly
squamose with fibrillose
scales up to the annulus,
pallid or brown below, white
and furfuraceous at the apex,
5-11 cm. long, 0.5-2.5 cm.
thick; spores broadly-elliptic
smooth, brown, 11-14 x 6-8 [i; cystidia ap-

parent in fresh specimens as occasional clavate proj ecting hyal

1927]

OVERHOLTS — THE GENUS PHOLIOTA 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: pi J$, 48.

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. The plants usually occur singly. The 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
cm. broad. Professor Har-
per illustrates, and I have
also found, a large form
with pileus 10-12 cm.
broad, but otherwise with

the characteristics of
Peck's types.

40. Pholiota aurivelloides

Overholts, sp. nov.

Pileus 5-8 cm. broad,
hemispheric or broadly
campanulate to convex,
ferruginous to tawny,
lighter-colored on the
margin, sometimes carob-
brown in dried plants,

!Figs. 127-130. P. aurivelloides: 127, section
of hymenium showing an imbedded cystidium
(a', spores (b), and central medulla (c), X 550;
123, spores, X 1200. All from type specimens.
12"), section of hymenium showing imbedded
cyistidia (a), and central medulla (b), X 550; 130,
spores, X 1200. All from specimens in New
Ycrk State Museum, from New Mexico, by
Co ckerell.

[Vol. 14

152 ANNALS OF THE MISSOURI BOTANICAL GARDEN

probably viscid, with a few scattered, spot-like or appressed
scales; flesh yellow, rather thick; gills sinuate-adnate or with a
decurrent tooth, medium-close or slightly distant, 7-12 mm.
broad, whitish then ochraceous-tawny or russet; veil forming a
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 cm. long, 5-10 mm. thick; spores oblong-
ellipsoid, smooth, deep-brown, 9-11 X 6 \x; cystidia numerous or
rare, brown, not projecting, 25-35 X 6-8 |i.

Habitat: on dead trees or from wounds in living Alnus, Salix, 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. The 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.

S. Spores 6-9 n or rarely longer.

aa. Cystidia present and rather conspicuous as broum 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./. 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

1827]

OVERHOLTS — THE GENUS PHOLIOTA IN THE UNITED STATES 153

smooth, 7-9.5 X 4-5 ^;

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, floccose above
the annulus, fibrillose
below and increasingly
scaly or shreddy down-
wards with fibrillose
scales that may become
recurved, solid, 5-8 cm.
long, 5-15 mm. thick;

spores exactly and con- Fig8 m _ m p auriveRa: m gection of

stantly oblong-ellipsoid, hymenium showing the imbedded cystidia (a),

spores (b), and central medulla (c), X 550; 132,
section of hymenium from another gill showing

Cystidia present, often projecting cystidia ,(a), and I spores (b) X 550. AH

from specimens at New York from Colorado, by
rather rare, brown, Betiel. 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 [i 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, 111.

Illustrations: Batsch, Elench. Fung. pi. 22, f. 114, 115; Cooke,
111. Brit. Fungi, pi. 390 {351)) Harper, Trans. Wis. Acad. Sci. 17:
pi. 38, 39.

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 collection have I seen these present — that
collected by Gilbert in Oregon, on willow, in 1915. The col-

[Vol. 14

154

ANNALS OF THE MISSOURI BOTANICAL GARDEN

lection is preserved at New York. Here 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. The 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. PL 20, 21.

Pileus 3-16 cm. broad, hemispheric
to convex or plane, antimony-yellow
to zinc-orange or finally the center
somewhat tawny, decorated with
rather medium-sized squamules of a
darker color than the rest of the
pileus, and drying down to small
dark spots on a 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 or 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,
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

Figs. 135-138. P. adiposa: 135,
section of hymenium showing the
imbedded cystidia (a), spores (b),
and central medulla (c), X 550; 136,
spores, X 1200. All from Overholts
Herb. No. 6191. 137, section of
hymenium showing cystidia (a),
spores (b), and central medulla (c).
X 550; 138, spores, X 1200. All
from Overholts Herb. No. 6042

19271

OVERHOLTS — THE GENUS PHOLIOTA IN THE UNITED STATES 155

with a small hollow, 4-12 en. long, 5-20 mm. thick; spores ellip-
soid or oblong-ellipsoid, smooth, 7-9 (-11) X 4-5 n; cystidia pres-
ent, not conspicuous, pro j ecti ng only slightly if at all, dark brown,
clavate, 20-42 x 5-10 ja.

Habitat: stumps and trunks of deciduous trees.

Distribution: specimens have been examined from Cambridge,
Mass. ; Lake Placid, Bronx P irk, and Wells, N. Y. ; Forked River,
N. J.; State College, Musser Gap, Bear Meadows, and Westport,
Pa.; Greencastle and Putnam Co., Ind.; Edgmont and Falling
Springs, 111.; Oxford, Ohio; St. Louis, Mo.; Fort Dodge, Iowa;
Plumas Co., Cal.; and Seattle, Wash.

Illustrations: Atkinson, Mushrooms, pi. 47 {43); Berkeley, Outl.
Brit. Fungi, pi 8, f. 2; Cooke, 111. Brit. Fungi, pi 395 {358) ;
Hard, Mushrooms, /. 211; Mycologia 1: pi 7, f. 1-2. 1909;
Peck, Rept. N. Y. State Bot 49: pi 46.

The pileus in both fresh an i dried condition is typically golden-
yellow or lemon-yellow as contrasted with the prevailing tawny
or rusty color of P. aurivella, its closest relative. The latter
species has larger and more spot-like scales and the stem is short
and thick and increasingly scaly

downward, while in P. adlposa
the stem is equally scaly t.p 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.

PL. 22.

Pileus 2.5-10 cm. broad, sub-
globose to convex or nearly plane,
light-colored, typically cinna-
mon-buff or cinnamon in dried
plants, viscid, covered with erect,
pointed, cinnamon or tawny scales
that give color to the pileus,
sometimes disappearing on the

Figs. 139-142. P. squarrosoides:

139, section of hymenium showing
cystidia (a), and spores (b), X 550;

140, spores, X 1200. All from Over-
holts Herb. No. 3770. 141, section of
hymenium showing cystidia (a), and
spores (b), X 550; 142, section of
hymenium showing central medulla
tissue, X 550. All from type speci-
mens.

[Vol. 14

156 ANNALS OF THE

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
cm. long, 5-12 mm. thick; spores ellipsoid or oblong-ellipsoid,
smooth, 4-6 X 3-4 \x; 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 \i.

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, /. 216; Harper, Trans. Wis.
Acad. Sci. 17: pi 36, 37; Peck, Ann. Rept. N. Y. State Mus. 54:
pi. 73, /. 6-14; White, Hymen. Conn, pi 21.

The scales on the pileus appear floccose 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 n, the wall is
thin, and the shape more apt to be broadly ellipsoid, while in P.
squarrosa, (> [i 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

Syst. Myc

1821. PL 21.

Pileus 3

cm. broad

ish or yellowish-brown, antimony-yellow to tawny in dried

mens, covered with rather

recurved

yellowish

scales, dry; flesh yellowish, baste mild; gills sinuate-adnate and
often somewhat decurrent, medium-close, 3-6 mm. broad, pallid
then ferrugi
dried plants
from honey-}

lous, in
varying

wn

veil forming a thick,
persistent, floccose
annulus often striate
on the upper surface;
stem central, equal,

**Os

00

d, yellow

or

brown, with conspic-
uous recurved scales
up to the annulus,
solid, 5-12 cm. long,
5-12 mm. thick;
spores oblong or ellipsoid, smooth, 6-8 X 3.5-4

Figs. 143-146. P. squarrosa: 143-145, sections of
three gills from same plant showing various forms of
cystidia (a), and spores (b), X 550; 146, spores, X
1200. All from Overholts Herb. No. 8058.

cystidia

variable, of

(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 \i.

Habitat: on dead trunks or stumps of various trees, both de-
ciduous and evergreen.

Distribution: specimens hnve been examined from Piscataquis
Co., Me.; Stocktenridge, Mass.; Yama Farms and North Elba,
N. Y.; Washington, D. C; S;ate College, Pa.; Jack Brook, Colo.;
also reported from Michigan and Wisconsin by Harper.

Illustrations: Bernard, Champ. Roch. pi. 19, f. 1; Cooke, 111.
Brit. Fungi, pi. 391 (367); Hard, Mushrooms,/. 217; Hussey, 111.

1: pi. 8; Harper j Trans. Wis. Acad. Sci. 17: pi. 35;
L Tab. Anal. Fung. /. 340; Peck, Rept. N. Y. State

Myc

Mus. 55: pi. 79, f.

7.

For detailed data as to the differences between P. squarrosa and
P. squarrosoides , see the latter species.

158

[Vol. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

45. Pholiota rigidipes Peck, Bui. N. Y. State Mus. 157: 31.
1912.

Pileus 4-8 cm. broad, broadly convex, sometimes slightly or
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-

t hilly disappear in

Figs. 147-148. P. rigidipes
menium showing cystidia

■pes: 147, section of hy-

(a) and spores (e), and

central medulla (f), X 550; 148, spores, X 1200. All

from type specimens.

mature plants, dry;
flesh white, tinged
yellow next the gills,
distinctly yellow in
dried plants ; taste
mild ; gills sinuate-
a d n a t e or adnate,
medium-close, 3-7
mm. broad, cinna-
mon or ochraceous-
tawny and retaining
these colors in dry-
ing; veil forming a
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
X 3.5-4.5 \i; 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,
Mo.

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

1927J

OVERHOLTS — THE GENUS PHOLIOTA IN THE UNITED STATES 159

a Flammula. A second collection, perhaps referable here, is
labeled P. terrigena and was collected by Atkinson, October, 1902.
This collection has dirt attached to the basal part of the stem, but
may have been also from buried wood. In 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

1905

Stropharia Schraderi Peck, Bull. Torr. Bot. Club

Pileus 5-8 cm. broad, convex to nearly plane, pallid when
young, ochraceous-buff when mature, dry, fibrillose, squamulose,
or rimose-squamulose on }he
disk; context white, with taste
of radishes; lamellae adns/te,
close, thin, whitish then brown ;
annulus small, lacerate, white,
sometimes evanescent; stem
central, subequal, squamubse
and concolorous with the pileus
below, white and mealy above,
solid, 2-4 cm. long, 8-12 nm.
thick; spores ovoid, not apicu-
late, smooth, 6-8 (-9) X 4-6 n;
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,

D. C.

The plants are considerably like P. fulvo-squamosa but the stem
is less prominently scaly, ths 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

aaa. Pileus 1-8 cm. in diameter (see also bbb, p. 1 62).

Figs. 149-150. P. Schraderi: 149,
section of hymenium showing cystidia
(a-b), and spores (c), X 550; 150, spores,
X 1200. All from type specimens.

160

[Vol. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

47. Pholiota confragosa Fries, Epicr. Syst. Myc. 169. 1836-
38. PL 22, 23.

Pileus 1-2.5 cm. 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

OOP

and taste not marked; gills
adnate or slightly decurrent,
sometimes white-crenulate
on the edge, medium-close,
1-3 mm. broad, rufous to
cinnamon-brown; veil form-
ing a 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-

tral, equal or enlarged

of edge of a gill showing the clavate or lance-
shaped sterile cells (a), and spores (b), X

550; 152, spores x 1200. All from Overholts a t base, concolorous with the

Herb. No. 8061.

p ileus 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 X 4-5 (i ; 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.), N. Y.; also re-
ported from Michigan by both Harper and Kauffman.

Illustrations: Fries, Ic. Hym. pi. 105, f. 2; Harper, Trans. Wis.
Acad. Sci. 17: pi. 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. caverata or P. comosa as seen without, a, lens.

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 cm. broad, hemispheric then convex or nearly
plane, tawny-brown, not changing color in drying, dry, densely
covered with minute, erect, pyramidal, spine-like, or granular,
tawny scales; gills adnexed ^

or adnate, medium-close or Uc^
subdistant when young, 1-2
mm. broad, pallid then cin-
namon-brown; veil forming
a slight, superior, floccose,
evanescent annulus ; stem

nDni , Q i onufll tflwnv with Figs. 153-155. P. erinaceella: 153, section

central, equal, tawny, witn of j^ of giUs showing clavate aterile ce n 8>

numerous floccose Scales or X 550; 154, spores, X 550; 155, spores, X

, , , , , i 1200. All from Overholts Herb. No. 3751.

granules below the annulus,

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 X 4-5 ja; 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: pi 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. PI. 14.

Pileus 1-3 cm. broad, convex to plane, obtuse or often slightly
umbilicate-depressed, golden-brown or tawny-yellowish when
fresh, brown or 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 sinuate-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;

162

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

veil forming an indistinct superior evanescent annulus; stem
central, equal, yellowish but with few, rusty-brown, fibrillose,

sub-erect scales or becoming nearly
glabrous, stuffed then hollow, 3-7 cm.
long, 2-5 mm. thick, sometimes with
a bright yellow mycelium at the base;
spores ellipsoid, smooth, 6-8 X 3.5-4 (ij
cystidia none.

Habitat : on rotting logs of deciduous

Figs. 156-157. P. muricata:
156, section of hymenium show- trees,
ing spores (a), X 550; 157,
spores, X 1200. All from speci-
men at New York from Michi-

gan

Distribution: specimens have been
examined from Ann Arbor, Michigan;
also reported from Illinois by Harper.

Harper, Trans. Wis. Acad. Sci

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 SDecies.

The scales on the

leus are also different. P. erinaceella may not be distinct but
smaller plant, with more spine-like scales and more brown

color.

666. Pileus 2.5-7 cm. broad.

50. Pholiota lucifera (Lasch) Fries, Epicr. Syst. Myc. 167.

1836-38.

Agaricus luciferus Lasch, Linnaea 3: 408
Pileus 2-5 cm. broad, convex or

plane, uniformly yellow, ochra

1828

dried

buff or ochraceous
plants or the cei

more

tawny, viscid, decorated with small
yellowish or tawny squamules;
flesh yellow; gills adnate, close,
2-4 mm. broad, yellow then brieht

rustv-br

white

on

the edge when young ; veil forming
a superior evanescent annulus;
stem central, equal or slightly thickened at the base, solid

Figs. 158-159. P. lucifera: 158, sec-
tion of hymenium with spores (a),
X 550; 159, spores, X 1200. All from
specimens at New York from Bresadola.

19 27 J

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 n; cystidia none.

Habitat: on dead trunks a ad limbs.

Distribution : reported by Kauffman from Michigan.

Illustrations: Bresadola, Fung. Trid. 1: pi. 85; Ricken, Blat-

terp. pi. 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 cm. broad, convex or nearly plane, sometimes um-

bonate, lemon-yellow when f:*esh, retaining the color in the dried
plants or becoming slightly tawny, with scattered, reflexed or
suberect, fibrillose, reddish or tawny
scales, viscid; flesh thin, yellow; gills
sinuate-adnate or slightly adnexed,
close, 2-4 mm. broad, whitish, be-
coming ferruginous, honey-yellow or
cinnamon-buff in dried plants; veil
forming a floe cose, evanescent, yel-
low annulus; stem central, equal, Figs. 160-161. P. limonella: 1 60,

■vpllnwkh with ^rattprpd roourved section of hymenium showing cys-

yeiiowisn, witn scanerea r«,curveu ^ ^ sporeg (b)> &nd centnd
yellow scales, smooth above the an- medulla (c), x 550; 161, spores, x

1 i- i o n 1 o e 1200. All from type specimens.

nulus, solid, 3-7 cm. long, 3-5 mm. J * *

thick; spores ellipsoid or ovsid, smooth, deep brown, 6.5-7.5 X

4.5-5 i&; cystidia none; trama with a distinct central medulla.

Habitat: prostrate trunks of beech and birch.

Distribution: specimens hive 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 hvmenium, and tin 3 plant lacks the yellow pulverulent

164

[Vol. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

appearance of dried plants of that species. It may not be distinct
from P. lucifera.

O

52. Pholiota tuberculosa Fries, Syst. Myc. 1: 244. 1821.

PL 14.

Pileus 2-6 cm. 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
a superior, often evanescent annulus;
stem central, with a prominent bulb
just at base and rooting below, hbrillose
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 x 4-5 n; cystidia none.

Figs. 162-103. P. tuberculosa:
162, section of hymenium with
spores (a), X 550; 163, spores,
X 1200. All from specimens at
New York from Europe by
Bresadola.

Ic.

Habitat : on dead wood of deciduous trees.

Distribution: reported by Harper from Michigan.

Illustrations: Cooke, 111. Brit. Fung. pi. 898a (370); Fries,

Hym. pi 104, f. 2; Harper, Trans. Wis. Acad. Sci. 17: pi 41,
B.

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 [l) are somewhat smaller than I find in European
plants.

53. Pholiota curvipes Fr. Epicr. Syst. Myc. 168. 1836-38.

Pileus 2-5 cm. broad, convex to plane, ochraceous-orange,
more tawny in age and in herbarium specimens, at first innately
floccose or densely silky-floccose, in 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

STATES 165

floccose-crenate, 3-6 mm. broad; veil form
vanesoent. radiate-floccose annulus, or an

nulus none; stem central, equal or
tapering upward, ochraceous-orange or
ochraceous-tawny, clear yellow at apex,
decidedly floccose-fibrillose, hollow, 2-5
cm. long, 2-5 mm. thick; spores ellip-
soid or oblong-ellipsoid, smooth, 7-8.5
X 4-5 u.: cystidia none.

XT , ., . , , j i j -j Figs. 164-165. P. curvtpes:

Habitat : on dead WOOd Ol deciduous 164> section of hymenium with

t rpA (3 spores (a), X 550; 165, spores,

U1 ee& * , X 1200. All from specimen at

Distribution: specimens have been New York from Sebec Lake,
examined from Sebec Lake, Me.; Os- Mame -
ceola, N. Y.; and Creve Coeur, Mo.; also reported from Ann
Arbor, Mich., by Kauffman.

Illustrations: Fries, Ic. Hym. pi 104, /• 3; Cooke, 111. Brit.

Fungi, pi 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

In

hundred collections of Pholiota 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 ths pileus, but darker and more sordid
toward the stem, and the stem 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 mo:*e 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

[Vol. 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. P. lucifera
seems to differ in having a 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. The scales and

much bright

both described as tawny while in my plants they

54. Pholiota angustipes Peck, Rept. N. Y. State Mus

1878

PI

•7 cm. broad, hemispheric becoming

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-

ryik 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. angmtipes: yellow, becoming tawny-brown, but
166, section of hymenium with cinnamon-buff or cinnamon in dried

the inconspicuous sterile cells . ..

(a), and spores (b), x 550; 167, plants; veil forming a slight, usually

spores, X 1200. All from type
specimens.

innulus ; stem centr
downward, whitish

laneous, slightly squamose or fibrillose, stuffed or hollow, 3-7.5
cm. long, 4-12 mm. thick; spores ellipsoid, smooth, dilute-brown
under the microscope, 6-8 X 3-5 fx; 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: pi 84.

The species is a well-marked one in the pallid to brown color-

1927]

OVERHOLTS — THE

167

ation and the minute scales i;hat 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 a 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, Bui. Torr. Bot. Club 30:

95. 1903.

PI. 13.

Pileus fleshy, 6-12 cm. broad, rather thin, convex becoming
nearly plane, dry, covered wii;h 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
to a slight collar around the
stem, whitish becoming pinkish-
cinnamon then dark cinnamon

with a white-crenulate edge; an-
nulus ample, membranous, per- .™&- 168-169. P. fulvo-squamosa:

' 168, section of hymenium with spores

sistent, scaly below, the uoper (a), x 550; 169, spores, x 1200. All
surface striate; stem central, from 0verholt8 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 X 3.5-4.5 n; 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: pi. 60.

I have seen no specimens cf this plant other than the type at

[Vol. 14

168

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Albany. At first glance the dried plant looks much like an
Agaricus, similar to A. subrufescens . The gills are quite narrow
as in P. duroldes. 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, pi. 6.
1828-30.

Pileus 6-15 cm. 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 adnata

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

Fi gB . I7<y-i7i7 P. destruens: no, mm. broad; veil white, soon break-
section of hymenium with sporeB i ng forming an evanescent, white,

(a), X 550; 171, spores, X 1200. *" & ' '

noccose-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
cm. thick, solid; spores ovoid or ellipsoid, smooth, 7.5-9.5 X

4-6 |i; 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, 111.; 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).

All from specimens at New York
from Kansas by Bartholomew.

1927J

OVERHOLTS — THE GENUS IHOLIOTA IN THE UNITED STATES 169

Illustrations: Bresadola, Fung. Trid. pi. 84; Hard, Mushrooms,
/. 214; Harper, Trans. Wis. Acad. Sci. 17: pi. 45-47; Kalchbrenner,
Ic. Hym. Hung. pi. 13, f. 1 (as P. comosa).

Bresadola considers P. comosa and P. heteroclita as synonyms
of this species, and he has bee: a 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 bo 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 :iear St. Louis by Glatfelter describe
the taste as sweet at first, becoming bitter.

Supplementary Key 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

Spores smooth at all stages 9

2. Wood-inhabiting; 1 fresh plants bright-colored, i.e., some shade of yellow,

green, or red; gills frequently bi-ight-colored; not viscid; spores 6-10 \x
long; cystidia not noteworthy 3

Wood-inhabiting; 1 gills not bright -colored but cinnamon; spores mostly
6-10 m long; cystidia present though not always abundant, flask-
shaped with a projecting neck 6

Ground-inhabiting, in woods; spores variable, but in all species except

one more than 10 y. long 7

Among Polytrichum moss on the ground; very small plants, less than 1
cm. broad; stem 1-2 mm. thick P.

minima

3. Pileus with distinct shades of greer or ashy-green when young or on being

handled; flesh green-tinged; gilbi bright-colored (yellow to ochraceous-
orange) and remaining so in dried plants; pileus scaly with fibrillose
scales P. aeruginosa

Plants not entirely as above 4

4. Pileus 2-5 cm. broad, typically pinkish-red in color, squamulose with in-

nate scales (i.e., not formed by the separating of a fibrillose cuticle on
the pileus); flesh usually pale avender; gills bright-colored in dried

Plants P. luteofolia

Plants not entirely as above 5

5. Pileus 4-15 cm. broad at maturity, 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. spectabili

u

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.

[Vol. 14

170 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Pileufl hygrophanous, glabrous, otherwise as in P. spectabilis P. cerasina

6. Plants hygrophanous (never viscid), watery-brown and striatulate on the

margin when moist, ochraceous on drying, cinnamon to tawny in her-
barium specimens, not at all scaly; gills simple; annulus soon evanescent
or leaving only a band on the stem P. marginata

Plants as above except pileus viscid, annulus sometimes persistent, and

dried plants cinnamon-buff to cinnamon P. discolor

Plants as in P. marginata except annulus membranous, persistent, con-
spicuous P. unicolor

Plants as in P. marginata except gills distinctly forking P. furcata

7. Spores broadly-ovoid to subglobose, 7-9 X 5-6 m, distinctly verrucose;

pileus entirely covered with matted floccose fibrils, or these collected
into squamules at the center of the pileus; plants with somewhat the

aspect of Cortinarius P. trachyspora

Spores ellipsoid or elliptic, more than 12 /x long, distinctly verrucose 8

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,
particularly in dried plants; plants eastern P. caperata

9. Plants growing on the ground 1 and with either pileus or stem or both scaly. ... 10
Plants growing on the ground 1 or among moss and with neither pileus nor

stem scaly, or at most only floccose or scurfy 17

Plants growing on wood 1 and with either pileus or stem or both scaly or with

floccose patches 35

Plants growing on wood 1 and with neither pileus nor stem scaly 52

10. Stem scalv 13

Stem not scaly or at most only strongly floccose or scurfy 11

11. Spores constantly 6 ^ or less long; annulus membranous, disappearing

P. duroides

A considerable number of spores in each mount measuring as much as 8

or 9 m long; annulus a persistent cottony roll on the stem 12

Spores 9-12 p. long; annulus membranous, disappearing P. aurea

12. Pileus brown, ochraceous-brown, or ochraceous-tawny, minutely scaly

all over; annulus slight, evanescent P. angustipes

Pileus ochraceous to pallid, minutely scaly only at the center; annulus a

persistent cottony roll on the stem P. Johnsoniana

13. Sterile organs of conspicuous form or color entirely lacking from the hy-

menium; pileus and stem covered with a fibrillose cuticle that soon

separates into soft silky scales on both pileus and stem 14

Sterile organs present in the hymenium, sometimes projecting but some-
times with the aspect of over-mature basidia, and brown in color and
not projecting; plants not otherwise as above 15

14. Spores 6-8 m long, rather strongly apiculate P. fulvo-squamosa

Spores 8-9 m long, not apiculate P. terrigena

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.

1927]

OVERHOLTS — THE GENt S 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, snail, and appearing as black dots to the
unaided eye, or medium-sized and more conspicuous; cystidia not as
above 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 cm. broad. .P. angustipes
Cystidia abundant, projecting, sharp-pointed, many hyaline or with
rounded brownish globules inside; plants dark brown, 1-3 cm. broad

P. terrestris

Cystidia rather numerous, projecting, fusoid, hyaline; plants 5-8 cm. or

more broad when mature, ochraceous-buff P. Schraderi

17. Spores ovoid or narrow-ovoid with a truncate apex and base more or less

obtuse or rounded though sometimes apiculate 18

Spores variously shaped, not truncate at the apex but sometimes apiculate

at the base 27

18. Spores narrow-ovoid, 4-5 m broad, or else annulus median, conspicuous,

striate on the upper side; cyst dia none; slender plants not more than

4 cm. broad 19

Plants not entirely as above 23

19. Plants growing in wet places an.ong Sphagnum moss P. mycenoides

Plants not as above 20

20. Plants watery-brown, ochraceous in herbarium specimens; stem 2-10 cm.

long

Plants yellowish-red or dark feiruginous and retaining these colors in

21

herbarium specimens; stem 1-S .5 cm. long P. rugosa

21. Spores 6.5-9 /i long P. blattaria

Spores 9-10.5 m long 22

22. Stem 2 mm. or more thick; medk m-slender plants more than 1 cm. broad

P. togularis

Stem about 1 mm. thick; very slender plants less than 1 cm. 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

carried into the woods 24

Plants growing on the ground in dense woods, or growing on wood 26

24. Spores 10-14/2 long; cystidia scarcely projecting, the apex broadly round-

ed; pileus white, soon areolate P. vermiflua

Plants not entirely as above; spores 8-10.5 m long; cystidia flask-shaped

and projecting conspicuously if present 25

25. Spores 4-6 /x broad; pileus white or tinged with yellow or tan; gills narrow

or medium-broad P. praecox

Spores 6-7 m broad; pileus ochracoous-yellow; gills very broad. ... P. temnophylla

26. Plants growing on wood or on rioh humus; pileus rugose at times. P. Acericola
Plants growing on the ground; pileus never rugose P. Howeana

27. Spores up to 9 m long 28

Spores more than 9 m long 33

172 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

28. Plants growing among Polytrichum moss, small, less than 1 cm. broad;

spores finely verrucose under the oil-immersion lens P. minima

Plants not entirely as above 29

29. Plants small or medium-sized, less than 4 cm. broad; cystidia none or not

noteworthy 30

Plants larger; cystidia various, sometimes absent 31

30. Annulus median, long-persistent, striate on the upper surf ace . . . . P. blattaria
Annulus superior, evanescent P. anomala

31. Plants western; pileus slimy-viscid; stem conspicuously white-tomentose

just below the annulus P. albivelata

Plants eastern; pileus dry; stem not white-tomentose 32

32. All spores 6 xx or less long; cystidia rather abundant but not conspicuous,

mostly with a mucronate tip; annulus membranous, disappearing. P. duroides
Some spores in every mount as much as 8 or 9 m long, mostly 5-9 /x; cys-
tidia quite rare and scarcely noteworthy, or abundant and conspicuous;
annulus a cottony roll on the stem P. Johnsoniana

33. Plants small, less than 2 cm. broad P. subnigra

Plants larger, 2-10 cm. broad 34

34. Plants 2-5 cm. broad; gills adnate or slightly decurrent; stem fibrillose

below the annulus P. erebia

Plants as above, but stem concentrically white-zoned from the veil . P. platyphylla

Plants 4-10 cm. broad; gills distinctly decurrent P. ombrophila

Plants 10-15 cm. broad; gills 7-20 mm. broad, rounded behind P. aurea

35. Spores 9 jx or less long, or if somewhat longer, then pileus glabrous 36

Spores 9-11 /x long, oblong-ellipsoid; brown sterile organs present in the

hymenium; plants western; pileus scaly P. aurivelloides

Spores 11-14 /x long, broadly-elliptic or fusoid-elliptic; brown sterile

organs absent from the hymenium; plants eastern; pileus scaly . P. albocrenvlata

36. Stem only scaly and spores with a truncate apex; small plants up to 3 cm.

broad; no sterile organs in the hymenium P. mutabilis

Plants not entirely as above 37

:«

39

37. Spores 3-6 xx long

Spores 6 /x and more long

38. Pileus pallid to cinnamon when fresh, viscid, densely scaly with erect or

sub-erect, pointed, concolorous scales; stem with concolorous scales

P. squarrosoides

Pileus lemon-yellow to tawny or fiery-yellow when fresh, dry, with fibril-
lose, superficial, sulphur-yellow scales; the stem with yellow floccose
scales or scarcely more than yellow flocci P. flammans

39. 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 1 in the hymenium; spores 6-8 /x long.

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 m long P. oregonense

1 But there are usually present more inconspicuous bodies, not projecting, and
with the appearance of old basidia, irregular in shape. These are apt to show up
particularly well in permanent sections in glycerine.

1927J

OVERHOLTS — THE GENUS PHOLIOTA IN THE UNITED STATES 173

Plants not entirely as in either of the above 40

40. Hymenium with numerous sterile brown organs, imbedded or projecting;

all species bright-colored, yellow, reddish, or tawny, and with both

pileus and stem scaly, and pile is more than 5 cm. broad 41

Hymenium with distinct projecting hyaline fusoid 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 variable 44

41. Pileus with large appressed spot-like scales; stem increasingly scaly down-

ward P. aurivella

Plants not entirely as above 42

42. Pileus viscid P. adiposa

Pileus dry 43

43. Pileus with appressed scales; stem only inconspicuously scaly, 4-6 mm.

thick P. rigidipes

Pileus with recurved scales; stem conspicuously scaly with recurved scales

and 5-12 mm. thick P. squarrosa

44. Large species, 6-15 cm. broad, white to pallid or wood-brown, with large

white floccose spots or patches on the pileus; spores 7.5-9.5 m long. . P. destruens
Large species 6-12 cm. broad, wi h. fibrillose scales; plants growing at the
base of trees or attached to bjried wood; spores 6-8 m long, strongly

apiculate P. fulvo-squamosa

Plants not entirely as above 45

45. Both pileus and stem scaly 46

Pileus only scaly 49

46. Pileus deep rich brown or golden brown to tawny 47

Pileus brighter-colored, lemon-yellow to ochraceous-orange 48

47. Both pileus and stem densely covered with a sheath of small erect, conical,

superficial scales P. erinaceella

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
fibrillose, not sheathed P. muricata

48. 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 P. lucifera

Stem not with a bulb at the base; not sheathed but scaly P. limonella

49. Pileus 1-2.5 cm. broad, hygrophunous; gills cinnamon; annulus distinct,

persistent, membranous P. confragosa

Pileus 2-10 cm. broad; gills bright ochraceous-orange; annulus fibrous-
lacerate and evanescent 50

50. Pileus viscid P. lucifera

Pileus dry 51

51. Pileus soon areolate and often with greenish tints; edge of gills concolorous

P. aeruginosa

Pileus not areolate, never greenish; edge of gills conspicuously white-

crenate P. curvipes

52. Pileus floccose or silky-floccose, 2-5 cm. broad; gills, pileus, and stem

bright ochraceous-orange P. curvipes

Plants not entirely as above 53

53. Plants uniformly less than 4 cm. broad 54

174 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

Plants at maturity more than 4 cm. broad

56

54. Spores with a truncate apex, 6-8 n long; cystidia none; veil evanescent

P. marginella

Spores not truncate, 7.5-9.5 n long; flask-shaped cystidia present and

projecting, though not abundant 55

55. Pileus viscid; annulus subpersistent P. discolor

Pileus dry; annulus at first funnel-shaped, persistent and conspicuous

P. unicolor

Pileus dry; annulus usually persistent but not funnel-shaped P. marginata

56. Pileus ochraceous to tan; annulus long-persistent, membranous 57

Pileus bright-colored, ochraceous-buff to tawny; annulus soon evanescent;

gills rather distant and the gill cavities strongly veined P. oregonense

57. Cystidia rather abundant, flask-shaped, strongly projecting; pileus often

rugose; on rotten wood or on humus P. Acericola

Cystidia present and rather abundant but projecting only slightly and
more fusoid than flask-shaped; on recently felled logs or from wounds
of living trees P. Aegerita

Synonyms and Doubtful and Excluded Species

Pholiota aggericola Peck is P. erebia of this paper. See p. 119.

Pholiota appendiculata Peck, Bui. 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.) Sacc. 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. See p. 161.

Pholiota dura (Bolt.) Quel. Champ. Jura Vosg. 91. 1872
(Agaricus durus Bolt. Hist. Fung. pi. 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.) Sacc. Syll. Fung. 5: 754. 1887

1927]

OVERHOLTS — THE GENUS PHOLIOTA IN THE UNITED STATES 175

(Agaricus hormophorus Mcnt. 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 P. ombrophila Fries. See p. 121.

Pholiota lutea Peck is here treated as a 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. aa-uginosa 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, Bui. N. Y. State Mus. 122: 151. 1908
(Agaricus ornellus Peck, Rept. N. Y. State Mus. 34: 42. 1883).
Not distinct from Flammulc, polychroa Berk.

Pholiota radicosa (Bull.) Quel. Champ. Jura Vosg. 92. 1872
(Agaricus radicosus Bui. Herb. Fr. pi. 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, Eul. Torr. Bot. Club 23: 414. 1896.
Described as growing in sandy soil in Alabama. The spores are
rough-walled, 8.5-9.5 X 5-3 ;x. Flask-shaped cystidia project
from between the basidia. Both 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 for the recognition of the species. If
the spore measurements (5 X 4.5 v.) are correctly recorded it
would fall in the neighborhood of P. duroides, with which it seems
to have other characters in oommon.

Pholiota sphaleromorpha (Bull.) Quel. Champ. Jura Vosg. 91.
1872 (Agaricus sphaleromorphus Bull. Herb. Fr. pi. 640. 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

176

[Vol. 14

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.

Pholiota subsquarrosa (Fries) Sacc. Syll. Fung. 5: 750. 1887.
Reported by Mcllvaine. I have seen no specimens so referable,
and Mcllvaine records that the plants seem different from the
European species. See p. 151, 156.

Pholiola terrigena (Fries) Sacc. Syll. Fung. 5: 737.

1887

The

>cies has been reported from the United States but I have
material that corresponds to specimens from Bresadola.

The
orded should be compared with P. terrestris Overholts,

hich has smaller spores and prominent
Pholiota ventricosa Earle is P. spectabi
Pholiota mllosa (Fries) Sacc. Syll. Fun

See J
See p

1887

Speci

mens

determined by Peck and similar collections from the
ast seem referable to P. svectabilis.

Pholiota washingt

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 italics; and previously published valid names in ordinary type.

Page

Agaricus

Aegerita 139

aureus 122

aurivellus 152

autumnalis 134

caperatus 123

dactyliotus 174

detersibilis 161

flammans 148

hormophorus 175

Johnsonianus 130

luciferus 162

marginatus 132

mutabilis 138

praecox 106

radicosus 175

temnophyllus 112

togularis 114

unicolor 134

Flammxda

unicolor 136

viridans 148

Hebeloma

hortense 105, 107

Page

Hypodendrum

oregonense 140

Pholiota

Acericola 108

adiposa 154

Aegerita 139

var. strobiloidea 140

aeruginosa 147

aggericola 119, 121, 174

albivelata 128

albocrenulata 150

angustipes 166

anomala 127

appendiculata 174

aurea 121

aurivella 152

aurivelloides 151

autumnalis 134, 174

blattaria 113

caperata 123

cerasina 141

comosa 169, 174

confragosa 160

curvipes 164

1927]

OVERHOLTS — THE GENUS PHOLIOTA IN THE UNITED STATES 177

Page
Pholiota (continued)

dactyliota 1 74

destruens 1 38

detersibilis 161, 1 74

discolor 1 J5

dura 105, 174

var. xanthophylla 1)5

duroides 1 29

erebia 118

erinaceella 131

filaris 116

flammans 1 18

f ulvo-squamosa 1 37

furcata 136

heteroclita 169, 174

hormophora 1 74

Howeana 110

indecen8 121, 1 75

Johnsoniana 1 JO

limonella 1 33

lucifera 1 32

lutea 145, 1 75

luteof olia 1 13

luxurians 1 75

marginata 1 32

marginella 157

Mcmurphyi 1 24

minima 1 25

mollicula 1 75

muricata 131

mutabilis 1 38

mycenoides 1 15

ombrophila 1 20

Page
Pholiota (continued)

oregonense 140

ornella 175

platyphylla 117

praecox 106

radicosa 175

rigidipes 158

rubecula 142

rugosa 115

sabulosa 175

Schraderi 159

8peciosa 175

spectabilis 144

sphaleromorpha 109, 111, 175

squarrosa 157

squarrosoides 155

subnigra 120

subsquarrosa 151, 156, 176

temnophylla 112

terrestris 126

terrigena 127, 176

togularis 114

var. filaris 116

trachyspora 125

tuberculosa 164

unicolor 134

ventricosa 146, 176

vermiflua 104

villosa 176

washingtonensis 119, 121 F 176

Stropharia

Howeana 110

Schraderi 159

178

[Vol. 14, 1927]

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 8
P. vermijlua. Photo of specimens in Overholts Herb. 3248, X 2/3. Original

Ann. Mo. Hot. Gard., Vol. 14. L927

Plate s

OVERHOLTS— PHOLIOTA IX THE rxiTED STATES

[Vol. 14, 19271

180 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 9

P. praecox. Photo of specimens in Overholts Herb. 3318. Upper half, X 3/5,
lower half, X 4/5. Original.

Ann. Mo. Bot. Gard., Vol. 14, 1927

Platk 9

OVERHOLTS— PHOLIOTA IX THE UNITED STATES

[Vol. 14, 1927|

182 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 10

P. Accricola. Photo of specimens in Overholts Herb. 3911, X 3/4. Photo by
E. T. Kirk.

»

Ann. Mo. Bot. Gard., Vol. 14, 1927

Plate 10

O
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R

X

o

R

X

c

tr 1

C

2!

a

H

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R

[Vol. 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.

Ann. Mo. Hot. Gard., Vol. 14, 1927

Plate 11

C

H

C

-

Tf.

13

C

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3

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v.

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■

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4

s

.

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[Vol.. 14, 1927J

180 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.

Ann. Mo. Bot. Gaed., Vol. 14. 1!)27

Plate 12

C
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C/2

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c

c

H

s

X

[Vol. 14, 1927]

188 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 13

P. julvosquamosa (left half). From Harper, Trans. Wis. Acad. Sci. 17: pi. 60
1913. About natural size.

P. caperata (right half). Photo of specimens in Overholts Herb. 7967, X 1
Small specimen. Original.

Ann. Mo. Hot. Gaud.. Vol. 11. 1027

Platk Vi

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7:

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2;

CI
2

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j

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[Vol. 14. 19271

190 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 14

Fig. 1. P. marginata. From Harper, Trans. Wis. Acad. Sci. 17: pi. 54, fig. C.
1912. About natural size.

Figs. 2-3. P. muricata. From Harper, Trans. Wis. Acad. Sci. 17: pi 52 , fig.
A, B. 1912. About natural size.

Fig. 4. P. tuberculosa. From Harper, Trans. Wis. Acad. Sci. 17: pi 41, fig. B.
1912. About natural size.

/

Ann. Mo. Bot. Gard., Vol, 14, 1927

Plate 14

OVERHOLTS— PHOLIOTA IX THE UNITED STATES

[Vol. 14, 1927]

192 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 15

P. discolor (lower half). Photo of specimens collected in South Boulder Canon,

Colorado, July 18, 1923, elev. 10,000 ft, About natural size. Original.

P. rugota (upper, left). Photo of specimens in Overholts Herb. 3623, X 1.
Original.

P. togularis (upper, right). From Harper, Trans. Wis. Acad. Sci. 17: pi. 59. 1913.
About natural size. (Published as P. blattaria.)

Ann. Mo. Hot. Gaud., Vol. 14, 1927

Plate 15

OYERHOLTS— PHOIJOTA IX THE UNITED STATES

(Vol. 14, 1927|

194 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 16

P. speclabilis. Photo of specimens in Overholts Herb. 9SX9. Reduced somewhat
Photo bv L. W. Brownell.

Ann. Mo. Hot. Gard., Vol. 14, \ l .Y27

Plate 10

C
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tr 1

j

[Vol. 14, 1927]

196 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 17
P. apeclabilis. Reduced somewhat. Photo by L. W. Brownell

Ann. Mo. Bot. Gard., Vol. 14, 1927

Plate 17

3

X

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3

X

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>

[Vol. 14, 1927]

198 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 18
P. aeruginosa. About natural size. Photo by Burtt Leeper.

Ann. Mo. Hot. Card., Vol. 14, 1027

IYatk 18

C

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[Vol. 14, 1927]

200 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 19

P. albocrcnulata (upper part). From Harper, Trans. Wis. Acad. Sci. 17: pi. 42.
1912. About natural

fiammans (lower part). Right, from Fries, Ic. Hym. pi. 104, fig- 1; left, from
»er, Trans. Wis. Acad. Sci 17: pi. 41, fig. C. Both about natural size.

Ann. Mo. Bot. (Jard., Vol. 14, 1927

Plate 19

OVERHOLTS— PHOLIOTA IX THE UNITED STATES

[Vol. 14, 1927]

202 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 20
P. adipoaa. About natural size. Photo by L. W. Brownell

Ann. Mo. Bot. Gard., Vol. 14, 1927

Plate 20

Z

z

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F

3

-J

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[Vol. 14, 1927]

204 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 21

P. xquarrosa (upper half). About natural size. Photo by L. W. Brownell.
)\ adiposa (lower half). About natural size. Photo by L. W. Brownell.

Ann. Mo. Bot. Gard., Vol. 14, 1927

Plate 21

<>\ r ERHOLTS— PUOLIOTA IX THE TXITED STATES

IVol. 14, 1927]

206 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 22

P. squarrosoide* (large cluster). From Harper, Trans. Wis. Acad. Sci. 17: pi. 86
1912. Slightly reduced.

P. confragosa (inserts). From Harper, Trans. Wis. Acad. Sci. 17: pi. 41, fig. D
1912.

Ann. Mo. Hot. Card., Vol. 14, l'.l'JT

Plate 22

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(Vol. 14, 1927J

208 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 23

P. angustipes (except lower left corner). From Harper, Trans. Wis. Acad. Sci.
17: pi. 34. 1912. About natural size.

P. confragosa (lower left corner). Perhaps slightly enlarged. Photo presented
by E. T. Harper.

Aw. Mo. Bot. Gard., Vol. 14, 1027

Plate 23

( ) \ E RHOLTS— PH ( ) L I ( )T A IX THE UNITED STATES

[Vol. 14, 1927]

210 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 24

P. aurivella. Somewhat reduced. Photo by L. W. Brownell

Ann. Mo. Hot. Gard., Vol. 14, 1927

Plate 21

X
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Annals

of the

Missouri Botanical Garden

Vol. 14 SEPTEMBER, 1927 No. 3

A MONOGRAPH OF THE SECTION OREOCARYA OF

CRYPTANTHA 1

EDWIN BLAKE PAYSON*

Professor of Botany, University of Wyoming
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.

a Since the receipt of the manuscript of this paper, Dr. Payson passed away on
May 15, 1927.

Ann. Mo. Bot. Gard., Vol. 14, 1927

(211)

[Vol. 14

212 ANNALS

these

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 piny on (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-
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-

i

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. If 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.

The 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. The 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 a 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. The 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. The
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.

[Vol. 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. Jamesii 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. confer tiflora and C. flava
are very closely related and have been confused in the past with
C. leucophaea. The 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 cither case.

C. confertiflora C. flava C. leucophaea

Crests usually evi- Crests usually ob- Crests usually evi-
dent solete dent

Conspicuously het- Conspicuously het- Less heterostyled

erostyled erostyled

1927J

PAYSON — SECTION OREOCARYA OF 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. C. leucophaea seems to be less distinctly di-
morphic 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.
Jamesii var. multicaulis when contrasted with C. oblata and C.
P aimer i.

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 in 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. The 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

[Vol. 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 polyphyletic. C. Palmeri, oblata, confertiflora
and flam belong to the series of which C. Jamesii is the primitive
member. C. 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 Jamesii group. The 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.

The 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. The 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 a 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 a 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. The 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 a 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. The 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 cymes. As development proceeded the cymes became shorter
and the inflorescence tended to extend over more and more of the
stem.

218 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

1 1 . The rather slender imbranched 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. The linear-oblanceolate leaf is more primitive than the
broader ovate or spatulate types.

II. THE GENUS CRYPTANTHA AS A 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. All these differences go to show that Oreocarya
is, as a 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

specialized Oreocarya species or Oreocarya

cestor.

1927]

PAYSON — SECTION OREOCARYA OF CRYPT ANTHA 219

The other two sections of Cryptantha, Geocarya and Eucryp-
tantha, are, as it was stated before, characterized by the presence of
cleistogamous flowers. In 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.

The 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. The 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 a 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.

The 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

220 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

cleistogamous flowers common to the species. Eucryptantha,
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. The 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. confertiftora or C. setosissima

scarcely seem to be large or efficient enough to be of real value to

the species. The 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] 001

PAYSON — SECTION OREOCARYA OF CRYPTANTHA L£\.

In the

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 ne
teleological about this mode of development,
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

222

[Vol. 14

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 parents, 1 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 genua Lesquerella. Ann. Mo. Bot.
Gard. 8: 104. 1921.

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 223

North America, and the most primitive species is in the southern
part of the sectional range. The 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. The most primitive of
these invaded North America as Cryptantha in the limited sense
of recent American floras. The 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.

[Vol. 14

224

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Species and subordinate

Species and subordinate

Species and subordinate

groups that the author

groups as recognized in

groups as they might be

would recognize if a broad-

the present study

recognized if the specific

er specific concept were

concept were smaller and

employed

*

the presence of intermedi-
ates ignored

Jamesii

Jamesii

multicaulis

multicaulis

multicaulis

laxa

laxa

cinerea

cinerea

disticha

disticha
Jamesii

typica

typica

abortiva

abortiva

abortiva

pustulosa

pustulosa

pustulosa

Pdlmeri

Palme ri

Pulmeri

oblata

oblata

oblata

leucophaea

confer ti flora

confert iflora

confertiflora

flam

flava

flava

tyjrica

leucophaea

leucophaea

salmonensis

salmonensis

salmonensis

stricta

stricta

stricta

f nulngena

nubigena

nubigena

Clemensae

Clemensae

Clemensae

setosissima

setosissima

setosissima

virgata

virgata

virgata

h u m His

insolita

insolita

insolita

virginensis

virginensis

virginensis

tumulosa

tumulosa

tumulosa

modesta

modesta

modesta

typica

humilis

humilis

t nubigena

caesjntosa

caespitosa

caespitosa

thyrsiflora

thyrsiflora

thyrsiflora

t elata

elata

elata

sericea

sericea

tyjrica

typica

perennis

perennis

t aperta

aperta

aperta

Bradburiana

t elata

t aperta

rugulosa

rugulosa

rugulosa

interrupt a

interrupta

interrupta

sfriculifera

spiculifera

spiculifera

celosioides

celosioides

celosioides

Sheldonii

Sheldonii

Sheldonii

M acounii

Macounii

Macounii

sobolifera

sobolifera

sobolifera

typica

Bradburiana

Bradburiana

nana

nana

com mixta

co m mixta

commixta

Shantzii

Shantzii

Shantzii

ovina

ovina

typica

typica

nana

cana

cana

cana

propria

propria

propria

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA

225

Species

and
that

subordinate
the author

groups

would recognize if a broad-
er specific concept were
employed

fulvocanescens

breviflora

typica

echinoides
Jonesiana
longiflora

Wetherillii

typica

? tenuis
Osterhoutii
paradoxa
Bakeri
flavoculata

mensana

typica

Species and subordinate
groups as recognized in
the present study

breviflora
fulvocanescens
echinoides
Jonesiana

Wetherillii

longiflora

tenuis

Osterhoutii

paradoxa

Bakeri

mensana
flavoculata

Species and subordinate
groups as they might be
recognized if the specific
concept were smaller and
the presence of intermedi-
ates ignored

breviflora
fulvocanescens
echinoides
Jonesiana

Wetherillii

longiflora

tenuis

Osterhoutii

paradoxa

Bakeri

mensana
flavoculata
t cristata
t Shockleyi

TAXONOMIC CHARACTERS OF SPECIFIC VALUE IN THE SECTION

OREOCARYA

Nutlets. — Nutlet differences are remarkably constant betweei

individuals of any species in this

With rather few

ceptions, it is possible to determine any species by means of
mature nutlet. Differences in the character of the surface a
most important, although size, form of the scar, and outline a

usually characteristic. Were the

peculiarities the only

differences that exist, the species would be doubtfully tenable

species

However, when the plants are well known certain other

characters are evident that are correlated with nutlet characters
as well as with eeoeraphical distribution.

Corolla. — Certain differences

portance.
is very cc

the corolla are of great ira-
te corolla tube and calyx lobes
es. The form of the fornices,

absence of crests at the base of the tube

able diagnostic characters. Two species have yellow corollas

white

the corolla tube

pale yellow. In most species the fornices are distinctly

Calyx

The sepals offer a few valuable characters

The

amount of elongation of the lobes after anthesis is com

226

[Vol. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

x

JJL

Fig. 1. Comparison of floral and fruit details of four species of Cryptaniha: I. C.
virginensis, section Oreocarya; II. C. capituli flora, section Eucryptantha; III. C. Kingii,
section Geocarya; IV. C. ambigua, section Krynitzkia.

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 227

small in a 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 in which the flowers are borne. A 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. A series of careful
drawings of the gynobase was made and these are available in
the Rocky Mountain Herbarium.

[Vol. 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. The 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 a single mor-
phological character that will definitely separate Oreocarya and

[Vol. 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); Atamisquaea 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 are, 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. In 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
Oreocarya 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 Oreocarya 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 re-
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 in the 'Prodromus' transferred Bradburiana and
leucophaea to the genus Eritrichium in 1846 and so started a
treatment that persisted until 1885. During this period four
species and two varieties were described in the group. The most
important publication affecting the genus was by Gray in Proc.
Am. Acad. 10: 61. 1875. As a part of a revision of Eritrichium
Gray grouped the known species and varieties together under the
section Pseudo-Myosotis 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. The species are here
transferred to the section P seudokrynitzkia 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.

[Vol. 14

232 ANNALS OF THE MISSOURI BOTANICAL GARDEN

The 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.
In 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. M. E. Jones and K. 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 tlnited States, and this was fol-
lowed by a flood of floristic works on the plants of the regions so

[Vol. 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 OF 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. To 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:

[Vol. 14

236 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Plant producing only chasmogamic flowers.

Perennial or biennial herbs, native to North America Section I. Oreocarya.

Annual (or rarely perennial) herbs, native to North and South America

Seel ion 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 the cauline leaves

and frequently in the spikes above Section. III. Eucryptantha.

Cleistogamic flowers highly specialized, becoming lenticular structures borne
at the base of the stem 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.

2. 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.

3. 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, C. 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 C. 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

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 237

that a perfectly phylogenetic classification would result from
dividing the sections strictly according to morphological char-
acters, then there would be no question as to the preferable
procedure. As it is, the sections are convenient, mainly mono-
phyletic, 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. PI. 2: 850. 1876; Gray, Syn. Fl. N. Am. 2 1 : 196-197. 1878.

Krynitzkia, section Pterygium Gray, Proc. Am. Acad. 20: 276.

1885, in part.

Krynitzkia, section P seudokrynitzkia 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; Giirke 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, 111. 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.

or biennial herbs with a conspicuous setose, hirsute
indument (except in C. pustulosa). Leaves entire,
, spatulate or linear. Stems solitary from the root
or caespitose, commonly unbranched below the inflorescence,

oblanceolate

9 dm. high

glomerate

of elongating or reduced, simple or branched, bracteate or nearly
ebracteate, two-ranked, unilateral, scorpioid cymes. Sepals
distinct, usually conspicuously accrescent. Corollas white or
vellow. salverform: limb 4-12 mm. broad; tube equalling or ex-

238 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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 Vj-Vb °f 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 Key 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.
c. Corollas yellow, individuals distinctly dimorphic as regards stamen

southern California.

Wyoming

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

dd. 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 and

eastern Utah 6. C. flava

cc. Flowers white; individuals less evidently dimorphic as regards stamen

insertion; native to Washington and (?) Oregon 7. C. leucophaea

bb. Nutlets more or less roughened or wrinkled on dorsal surfaces; leaves
various.

c. Nutlets uniformly muricate or papillose, not evidently tuberculate,
rugose or wrinkled.

d. Leaves silky-strigose, pustulate hairs small or lacking; fornices
elongated.

e. Muriculations usually acute and sometimes minutely setose;
eastern Utah, adjacent Colorado to northern New Mexico and

northeastern Arizona 36. C. Julvocanescens

ee. Muriculations conspicuously 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.

38. C. Jonesiana

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- 0. oblata

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 239

dd. Inner surfaces of the nutlets distinctly roughened.

e. Nutlets conspicuously muricate, some of these confluent into
rugae; scar narrow, straight, open, no indication of an elevated

margin 98. & Jonesiana

ee. Nutlets rugose or tuberculate, 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

leaf surfaces 43. C. paradoxa

ff. Margins of the nutlets in contact.

g. Scar of the nutlets conspicuously open and surrounded by a

definite elevated margin 46. C. flavoculata

gg. Scar of the nutlets closed 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 rugose 41 > C. tenuis

jj. Leaves obovate or broadly oblanceolate; nutlets with
rounded rugae and tubercles; fornices long-papillose.
39. C. Wetherillii

hh. Leaves conspicuously pustulate ventrally, especially in

age; corolla tube 1 cm. or more long; Colorado. .40. C. 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-
culate.
c. Fruit depressed, globular, nutlets not in contact by their margins.

d. Leaves conspicuously pubescent on both surfaces.

e. Crests at base of corolla tube conspicuous; calyx not conspicu-
ously accrescent !• C- Jo/mesii

ee. Crests at base of corolla tube obsolete; calyx conspicuously ac-
crescent 3. C. Palmen

dd. Leaves glabrous above, sparingly hairy below 2. C. pustulosa

cc. Fruit conical or ovoid, nutlets in contact by their margins.

d. Stout, strictly erect plants with many elongated and conspicuous
bracts in the inflorescence; eastern Colorado and southeastern

Wyoming 19* C. virgata

dd. Smaller, somewhat caespitose plants with few or inconspicuous

bracts in the inflorescence; central Idaho 8. C. salmonensis

bb. Nutlets more or less roughened (muricate, wrinkled, tuberculate), 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 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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-
jacent Colorado (?) 35. C. breviflora

ff. Plants densely caespitose, caudex multicipital; leaves linear-
oblanceolate; native to eastern Wyoming and adjacent Ne-
braska and Colorado 33. C. cana

ee. Leaves with two distinctly different kinds of trichomes 32. C. nana

dd. Leaves distinctly subtomentose as well as appressed-setose . .32. C. nana
cc. Nutlets 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
12. 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 . . . AS. 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 Mexico
20. C. thyrsiflora

gg. Inflorescence narrower; nutlets usually with an evident

wing-margin (except in C. rugulosa); native west of the con-
tinental divide.

h. Nutlets with evident transverse rugae, these more con-
spicuous than the tubercles that may be present,
i. Nutlets scarcely or not at all muricate between the
rugae,
j. Strictly erect, conspicuously setose perennial of

Colorado 9. C. stricta

jj. Caespitose, less conspicuously setose perennials of high
elevation in Idaho, Nevada, Oregon, and California.
10. C. nubigena

ii. Nutlets distinctly muricate between the rugae and near
the margins of the nutlets.

j. Erect perennials from western Utah 24. C. rugulosa

jj. Caespitose and more or less soboliferous perennials

from Montana 30. C. sobolifera

hh. Nutlets with distant tubercles and no conspicuous rugae;

native of the southern Sierra Nevada 11. C. Clemensae

dd. Inner surfaces of the nutlets conspicuously and definitely rugose,
tuberculate or 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 leaf -surf aces 4&- C. mensana

gg. Elevated margin indefinitely limited; pustules present only

on the lower surface of the leaves 42. C. Osterhoutii

ff. Scar triangular and not constricted below the middle,

g. 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 open 14* C. insolita

hh. Cymules shorter and inflorescence narrow; long-lived
perennials; nutlets with only a slight dorsal ridge, if any.

i. Nutlets indefinitely tuberculate and rugose; California

16. C. tumulosa

ii. Nutlets definitely tuberculate or rugose; Utah and

eastern Nevada 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
caespitose perennials of eastern Oregon and adjacent

Idaho 34. C. propria

ii. Leaves setose and tomentose, less densely caespitose.

18. C. humilis

hh. Style not exceeding the mature nutlets by more than 0.5
mm.; densely caespitose perennials of southern Wyoming.
19. C. caespitosa

ee. Scar of the nutlets closed or nearly so, no conspicuous triangular,

open area near the base.
f. Upper surface of the leaves uniformly appressed-strigose and

without pustules.

g. Nutlets sharply rugose and tuberculate; scar surrounded by

an elevated margin 44- C. Bakeri

gg. Nutlets not so sharply rugose or tuberculate; scar not sur-
rounded by an elevated margin,
h. Densely caespitose from a multicipital caudex; native to

eastern Oregon and western Idaho 34* C. propria

hh. Less evidently or not at all caespitose; native to Utah,

Colorado, and Wyoming 22. C. sericea

ff. Upper surface of the leaves with two distinct kinds of hairs,
pustulate.

g. Densely caespitose perennials; style not over 0.5 mm.
longer than mature nutlets; stems not over 1.5 dm. high.
19. C. caespitosa

gg. Caespitose or solitary plants; stems usually more than 1.5
dm. high.

h. Perennials; mostly native in the Rocky Mountains or
west of them.

242 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

i. Mature sepals exceeding the nutlets by 2-3 mm.; in-
florescence broad-topped; western Colorado 21. C. elata

ii. Mature sepals exceeding the nutlets by 4-8 mm.
j. Nutlets tuberculate, scarcely if at all rugose.

k. Ventral surfaces of the nutlets nearly smooth;

Montana SO. C. sobolifera

kk. Ventral surfaces of the nutlets distinctly rough-
ened.
1. Stems 1-2 dm. high; inflorescence spreading;

western Colorado 23. C. aperta

11. Stems 2-9 dm. high; inflorescence usually nar-
rower; Nevada to Montana,
m. Nutlets broadly lanceolate; murications
lacking or indefinite; Oregon, Washington,

and Montana 28. C. Sheldonii

mm. Nutlets narrowly lanceolate; murications

very definite 25. C. interrupta

j j . Nutlets more or less rugose.

k. Scar of the nutlets somewhat open at the base;

Nevada and California 18. C. humilis

kk. Scar closed or very nearly so; northerly in range.
1. Nutlets 4-5 mm. long; along the Columbia River
and tributary streams in Oregon and Washing-
ton 27. C. celosioides

11. Nutlets less than 4 mm. long.

m. Leaves narrowly oblanceolate, strongly hir-
sute-ciliate.
n. Stems slender; nutlets 3-4 mm. long; Idaho

and Washington 26. C. spicnlifera

nn. Stems slender; nutlets 2-3 mm. long;

Canada, Washington, Wyoming. .29. C. Macounii
mm . Leaves broadly oblanceolate or spatulate.
n. Ventral surfaces of the nutlets nearly

smooth; Montana 30. C. sobolifera

nn. Ventral surfaces of the nutlets distinctly

rough; Oregon to Montana 28. C. Sheldonii

hh. Biennials; native to the plains east of the Rocky Moun-
tains and to the foothill region east of the continental
divide 31. C. Bradburiana

Series 1. Jamesianae. Nutlets smooth on all surfaces, or
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. Lye. N. Y. 2: 225. 1827, not
Cryptantha suffruticosa Piper, Proc. Biol. Soc. Wash. 32: 42. 1919.

1927]

PAYSON — SECTION OKEOCARYA OF CRYPTANTHA 243

Perennial; leaves linear-oblanceolate, obtuse or acute, 3-15
cm. 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 than 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 l /, to l / 4 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; leaves
tufted at the base; from southern Colorado and Utah to northern Mexico.
la. var. multicaidis

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 Texas lb. var. laxa

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 lc. 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
Arizona Id. 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
eastern Wyoming to eastern Colorado le. var. typica

Stems branched from the base, usually simple upwards, prostrate or nearly so,
0.6-1.5 dm. long; leaves somewhat tufted at the base; eastern deserts of
southern California and adjacent Nevada If. var. abortiva

244 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

la. Var. multicaulis (Torr.) new comb. Plate 25, figs. 1-4.

Eritrichium multicaule Torr. in Marcy, Expl. Red River, 262.
1854.

Oreocarya multicaulis (Torr.) Greene, Pittonia 3: 114. 1896.

Krynitzkia multicaulis var. setosa Jones, Contr. West. Bot. 13:
4. 1910. (Type: "near Ft. Cove, Utah, June 27, 1901, growing
under junipers." M. E. Jones.)

Oreocarya sujjruticosa var. multicaulis (Torr.) Payson, Univ.
Wyo. Publ. Bot. 1: 171. 1926.

Perennial from a woody root; stems branched from the base,
simple above, erect, x&ther 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 34, 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 335
(Mo., Gray) ; near Shattuck, Ellis Co., May 10, 1914, Clifton 3039
(Mo.) ; near Shattuck, May 16 and 17, 1914, Stevens 3039, 3024
(Gray). _ '

Texas: sandhills on 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, R. 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 Pecos,
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., Sept. 29-Oct. 1, 1919, Eggleston 16184 (Field); near Roswell,
June, 1914, Wooton (U.S.); Dona Ana, April, 1851, Thurber 293
(Gray); Jornada del Muerto, April, 1851, Thurber 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
(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,
R. 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 (R. 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 5291 % 5298b
(U.S.) ; canyon above Tropic May, 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 part); Skull
Valley, April 28, 1903, Jones (Pomona) ; Flagstaff, Aug. 6, 1884,
Jones 4007 (Pomona, R. 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
A140 (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 404? (Po-
mona, U.S.); Santa Rita Forest Reserve, March 31-April 23,

IVol. 14

246 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1903, Griffiths 4268 (U.S.); Alpine, July 31, 1912, Goodding 1264
(R. Mt., Calif., U.S.) ; Outlaw Canyon, Chiricahua Mts., July 30,

1907, Goodding 2349 (R.Mt., Gray).
Nevada: Ely, June 26, 1907, Jones (Pomona).
Chihuahua: Mexican boundary near White Water, June 18,

1892, M earns 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.).

lb. Var. laxa (Macbr.) new comb.

Oreocarya multicaulis var. laxa 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 cm. 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 \i-Y2 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.S.).

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).

lc. 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

hirsute : leaves somewhat tufted at the base

cm

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 y z or % 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. Type: "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 (U.S.);
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-Sept. 5, 1897,
Hough 8 (U.S.) ; Grand Canyon, June, 1915, Macbride & Payson
950 (R.Mt., U.S., Gray) ; vicinity of Flagstaff, July 2, 1898, Mac-
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,

|Vol. 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 875 (Gray); Peach Springs, May 26, 1884,
Jones 6684 (Pomona, R.Mt., Calif.).

Nevada: Ely, Aug. 11, 1913, Hitchcock 1193 (U.S.); Mt.
Magruder, May-Oct., 1898, Purpus 6068 (Pomona, U. S.).

Id. 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 cm. 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: "on 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 (U.S.); Barton
Range, s.e. Utah, July 13, 1895, Eastwood (Calif., U.S., Mo.,

Gray).

Arizona: Laguna Canyon, July 10-11, 1920, Cluts 37 (U.S.,

R.Mt., Gray).

le. Var. typica n. var.

Myosotis sujjruticosa Torr. Ann. Lye. N. 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 sujjruticosa (Torr.) Greene, Pittonia 1: 57. 1887.
Perennial, but probably shorter-lived than in the varieties
cinerca and multicaulis; stems branched upwards as well as from

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 249

the base, more or less decumbent, 10-30 cm. long, rather slender,
strigose and setose-hirsute; leaves usually not tufted at the base,
linear-oblanceolate, 3-8 cm. 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 onen. mostlv con-

fined to the upper %-% of the stem, cymes elongating, bracts of

the lower part of the inflorescence elongated and leaf-like; 2-4

com

Distribution : Upper Sonoran Zone in the western part of South
Dakota, Nebraska, and Kansas, northwestern Oklahoma and
Texas, southeastern Wyoming, and eastern Colorado. Type:
"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.); Al-
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
(N. Y., Mont.); Hamilton Co., Aug. 3, 1895, Hitchcock 347 (R.
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.S.) ; 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

omona) ; Sterling

Co., June 12, 1896, Osterhout (Minn.); Ft. Lupton, July

[Vol. 14

250

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, El 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, May
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.).

If. 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 cm. long, strigose and hirsute; leaves somewhat tufted at
the base, 5-10 cm. 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 % 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 OF CRYPTANTHA 251

Mts., June 21, 1922, Pierson 3103 (Pomona); Bear Valley, San
Bernardino Co., July 19, 1900, Jones 5315/ (Pomona); Bear
Valley, June 16-20, 1895, Parish 3694 (Calif., Gray); Bear
Valley, May, 1882, Parish 1480 (Calif., Mo., U.S., Gray);
Bear Valley, 6500 ft., June 15, 1894, Parish 3238 (U.S.).

C. Jamesii, in the aggregate sense, is the most widely distrib-
uted and most heterogeneous species in the genus. As a 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

be expected

the

This is ereatlv 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-

252 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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-

th appressed

Cine

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 laxa 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

adjacent Arizona. The character of the
is certainlv of no value. On Dart of the

Utah and
maturing

preserved in the herbarium of the University of California

some fruits were found

another

of similar general appearance four mature nutlets were found
Other varieties may need to be characterized from time
time. A study of the different varieties in intermediate ran
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
cm. long, acute or obtuse, glabrous and without pustules above,
conspicuously and densely pustulate and short-hairy below; in-

confined

the stem

g and probably lax in age, foliar bracts inconspicuous;
iceolate, acute, about 4 mm. long in anthesis, probably
>icuously elongated in fruit, strigose and rather sparsely
th 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 Y± their length; mature
fruit not seen, presumably very like C. Jamesii.

Distribution: Upper Sonoran Zone, southeastern Utah. Type:
" Hammond Canyon, Elk Mountains, July 31, 1911, Rydberg &

Garrett 9820"

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. The 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.
The specimen available of this species has more the appearance
of a Heliotropium or of a Plagiobothrys than a Cryptantha but
there seems no doubt that it is correctly placed in the latter genus.
The variety disticha of Jamesii is probably something of a con-

between Jamesii var. multicaulis and

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 cm. 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 \i 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

[Vol. 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 \

3

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 3 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. The 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. The 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
corolla tube with no crests at the base

(2) the

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 0. hispidissima (Torr.) Rydb.

1927)

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 255

0. oblata (Jones) Macbr. Proc. Am. Acad. 51: 548. 1916.
0. 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 Yi 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 l /i 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 El Paso. Type: "no. 3759 [M. E.
Jones], El Paso, Texas, April 23, 1884."

Specimens examined:

Texas: Marfa, March 26, 1919, Hanson 899 (U. S., Gray) ; near
J. Davis' R. C, West Texas, Sept. 1883, Havard (U.S.); Fort
Bliss, April 22, 1915, Carlson (Gray); El Paso, April, 1881, Vasey

250 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

(Mo.); near El Paso, March, 1851, Thurber 147 (Gray); El Paso,
April 23, 1884, Jones 3759 (Pomona, type, R.Mt., U.S.);
northern base of Chenate Mts., Sept. 1892, Nealley 167 (Field,
N.Y., U.S.).

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, Beats (U.S.); Berendo Creek, Sierra Co.,
May 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 a specimen from Peach Springs,

Arizona. This proves on examination to be C. Jamesii var.
cinerea.

C. oblata is most closely related to C. Palmeri and to C. Jamesii
var. multicaulis. It differs from both in having exserted corolla
tubes and somewhat roughened nutlets that are quite distinctly
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. (Type: "No. 52891, May 23, 1894, Johnson,
Utah, 5,000 ft. alt., on sandstone cliffs," M. E. Jones).

Oreocarya 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. (Type: "no. 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.

Long-lived perennial from a woody root; stems few to many

1927]

PAYSON — SECTION OREOCARYA OF 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 Yi 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 J4 their length, about % 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 j (Pomona, U.S.) ; Wa Wa, 30 miles west of

[Vol. 14

258 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Frisco, June 25, 1906, Jones (Pomona); Vermilion, June 4, 1901,
Jones (Pomona); canyon south of Glenwood, May 25, 1875,
Ward 108 (Mo., U.S., Gray) ; Johnson, May 23, 1894, Jones 5289t
(Pomona) ; Johnson, June 20, 1890, Jones (Pomona) ; Silver Reef,
May 3, 4, 1894, Jones 5144 (Pomona, R.Mt., Calif., Mo., U.S.);
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 (N. Y., U.S.); Red Canyon trail, Grand
Canyon, June 10, 1901, Ward (U.S.).

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, K. Brandegee
(Calif.) ; Goldfield, May 14, 1909, Heller 9619 (Phila.) ; Esmeralda
Co., ShockUy (Calif.).

California: Andrew's Camp, Bishop Creek, Inyo Co., July, 1911,
Davidson 2722 (Gray); Andrew's Camp above Bishop Creek,
July, 1913, K. Brandegee (Pomona, Calif.); Bishop Creek, May
31, 1906, Hall & Chandler 7234 (Calif.); northern slope of San
Bernardino Mts., May, 1882, Parish 1319 (Calif.) ; 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 ('98?), Purpus 5323 (Calif., Mo., U.S., Gray);
White Mts., 1898, Purpus 5802 (Calif., U.S.); Silver Canyon
east of Laws, May 9, 1906, Heller 8211 (Calif., Mo., Phila., U.S.,
Gray); south of Mono Lake, July 8, 1863, Brewer 1822 (U.S.);
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 (Calif.).

C. confer tiflora and its close relatives, C. flava and C. leucophaea,
are readily distinguishable from all other species of the section

1927]

PAYSON — SECTION OREOCARYA OF 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. The 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 lulea
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 Monographic der Gattung
Oreocarya den Namen 0. lutea Greene als Synonym zu 0. con-
fertiflora. Aber abgesehen von der goldgelben Bliitenfarbe sind
bei 0. lutea die Staubbliitter an der Spitze des Kronentubus dicht
unter den Hohlschuppen eingef iigt, wahrend sie bei 0. confertiflora
(0. 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 taxonomic 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 (A. Nels.) new comb. Plate 25, figs. 11-13.

Oreocarya flava A. Nels. Bull. Torr. Bot. Club 25: 202. 1898.

0. lutescens Greene, Pittonia 4: 93. 1899. (Type: "common
on hills about Aztec, New Mexico, 25 April, 1899," C. F. Baker.)

Very similar in habit of growth to C. confertiflora; stems 15-35
cm. tall, bristles somewhat coarser and more widely spreading
than in confertiflora; leaves linear or linear-oblanceolate, acute,
3-9 cm. 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

260 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

-% of the stem, scarcely glomerate, indument very similar to
that in confertifiora, 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 confertifiora, limb 8-9 mm. broad,
lobes united for about 34 their length, less than Yi as l° n g 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 confertifiora, margins in
contact, acute, not wing-margined, surfaces of nutlets glossy,
smooth, dorsal somewhat more rounded crosswise than in con-
fertifiora, 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, E. 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, Pay son & Pay son
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 10, 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.,
U.S.) ; 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.,
U.S.); 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 confertifiora 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. confertifiora.

[Vol. 14

262 ANNALS OF 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, Proc. Am. Acad. 20: 280.

1885.

Oreocarya leucophaea (Dougl.) Greene, Pittonia 1: 58. 1887.

Very similar in habit of growth to C. confer h 'flora; stems 15-40

cm. tall, white-hairy at the base, densely strigose but scarcely

setose upwards except in the inflorescence ; leaves linear to linear-

oblanceolate, acute, 3-9 cm. long, dorsal surface densely strigose

and with numerous weak bristles with pustulate bases, ventral

surface uniformly strigose, pustules few or none, leaf-bases and

inflorescence mainly restricted to upper l A 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 ^ 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)

mm. lone, 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. PI. 2: 338. 1884, and Henry, Fl. Southern British
Columbia, 254. 1915), and probably to be found in northern
Oregon. Type: "arid barrens of the Columbia, and of its north-
ern and southern tributaries." Collected by Douglas.

Specimens examined:

Washington: "ex 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 CRYPTANTHA 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 S921 (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. conferti flora. 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 ^2 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
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.

(

[Vol. 14

264 ANNALS OF THE MISSOURI BOTANICAL GARDDN

Distribution: Upper Sonoran areas in central Idaho. Type:
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. striata (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 a 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 ^ or % 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 x /i^° A 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 OF CRYPTANTHA 265

Distribution: Upper Sonoran Zone, northwestern Colorado.
Type: collected "some distance south of the Yampa or Bear
River in Moffat County (Colorado) along the Victory Highway."

Specimens examined:

Colorado: some distance 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. The 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. C. stricta may
represent a separate line of evolution from C. Jamesii 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 a 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 %, 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, crests at base of the tube evident, fornices prominent,
broad, papillose, yellow, about 0.5 mm. high, limb 4-7 mm. broad,
lobes slightly shorter than the tube (measuring from the fornices) ,

united for about y z 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 narrowlv

[Vol. 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. Type: "On 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 13435
(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. The 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 geographic varieties
when it is better known. Perhaps it is simply an especially
Dolvmornhic unit and anv one locality will give great extremes of

variation.

confused

lationship to other species is not clear,
and habitat, nubigena is most likely to be
That it is reallv closelv related to it is

not so certain. The outstanding peculiarities of nubigena are its
greatly elongated nutlets that are nearly smooth on the inner sur-

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 267

faces. A satisfactory concept of this species must await further
study and exploration.

mm

11. C. Clemensae Payson 1 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 cm. 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

; 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 H 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; inflorescentibus subcapitatis;
sepalis setosis linearo-lanceolatis 3-4 mm. longis, fructiferis ca. 7 mm. longis, quam
nuculae 3-4 mm. longioribus; corolla alba, tubo 2.5 mm. longo, quam sepalis brevi-
oribus, limbo ca. 4 mm. lato; stylo nuculas 0.5-1 mm. superante; nuculis linearo-
lanceolatis acutis 3 mm. longis anguste alatis, facie exteriore sparse tuberculosis,
faciebus ventralibus fere laevibus, sulco recto angusto, margine non edito. — Col-

Mrs

type).

[Vol. 14
268 ANNALS OF 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
(U.S.); Mt. Whitney, alt. 13,000 ft., Aug. 17, 1899, Copeland 52

(U.S.).

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,
C. Clemensae is very different in general appearance,
green and rather flaccid leaves eive it an annearanoe th

The thin

winged

the genus. The small, rather turgid but

nutlets are also quite characteristic. It seems as distinct
a 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. A study of the flowers and fruit seems to
make it cerlain 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. Virgatae. 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. rirgata — quite smooth. Species 12-13.

12. C. setosissima (Gray) new comb. Plate 20, 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 cm. long, somewhat reduced un-

1927]

PAYSON — SECTION OREOCARYA OF 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 Yi oi 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 l / ( 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. Type: "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.S.); 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 357 (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, Tourney 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-Oct. 1900, Purpus 8048 (Pomona, Calif., Mo., U.S.);
Flagstaff, Aug. 1883, Rusby 748 (Calif., Phila., U.S.); Mt. Hum-

270 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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 5800 (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 C. 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 0. holoptera (Gray) Greene; now treated as Cryptantha
holoptera (Gray) Macbride.

13.

rter) new comb. Plate 26, figs. 32-34

um Porter, Hayden Rept. 479. 1870.

r. virgatum Porter in Porter & Coulter, Syn

Fl. Colo. 102. 1874.

Krynitzkia virgata (Porter) Gray, Proc. Am. Acad. 20: 279.

1885.

Oreocarya virgata (Porter) Greene, Pittonia 1: 58. 1887.

O. spicata Rydb. Bull. Torr. Bot. Club 36: 678. 1909. (Type:
Artist's Glen, Pikes Peak, Colorado, Aug. 1, 1901, Clements 102.)

0. 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 loner, stout, divaricate bristles, hirsute: leaves narrowlv

1927)

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 271

oblanceolate, obtuse, 3-12 cm. long, hirsute and setose, appressed
pubescence lacking, pustulate hairs numerous on both surfaces
but somewhat more numerous dorsally; inflorescence usually
extending along at least % 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 % 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 Ve 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. Type: "near 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 231 (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 4%o~3
(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

[Vol. 14

272 ANNALS OF THE MISSOURI BOTANICAL GARDEN

(R.Mt.); Front Range, July 2, 1896, Crandall (R.Mt., Wash.,
Calif., 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 AA07 (R.Mt.); near Boulder, July, 1902, Tweedy 5223,
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. The 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 O. spicata Rydb. is
not even deserving of varietal rank. It is simply to be regarded
as a 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 OF 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 cm. 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

rcely ciliate; inflorescence mostly confined to the u]
of the stem, cymes rather few (for the genus) and

or

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 papillose, slightly emarginate,
limb about 7 mm. broad, lobes and tube subequal, lobes united for

about 34 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. Type: Las Vegas, Nevada, May 4, 1905, L. N.
Goodding 2286.

Specimens examined:

Nevada: Las Vegas, May 4, 1905, Goodding 2286 (R.Mt.,
Minn., Mo., Gray, type); Las Vegas, K. Brandegee (Calif.);
Las Vegas, April 29, 1905, Jones (Pomona).

C. insolita is probably most closely related to C. virginensis and
C. tumulosa. The 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:

[Vol. 14

274

ANNALS OF THE MISSOURI BOTANICAL GARDEN

C. tumulosa
Perennial.
Stems 1-2.5 dm.
high.

C. insolita
Biennial.
Stems 1.5-4 dm.
high.

C. virginensis
Biennial.
Stems 1.5-4 dm.
high.

Leaves setose below, Leaves inconspicu- Leaves conspicuous-

with subappressed ously setose below

setose

below

bristles.

with
bristles.

appressed

with
bristles.

spreading

Inflorescence narrow, Inflorescence broad, Inflorescence broad,

cymes short.

cymes elongating, cymes elongating,

very numerous,
flowers crowded.

few, and flow r ers
rather distant.

Nutlets slightly Nutlets distinctly Nutlets distinctly

carinate.

carinate.

carinate.

Dorsal surface with Dorsal surface with Dorsal surface w T ith
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.

1916.

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, obtu

12 cm. long,

rather sparsely setose, slightly hirsute and distinctly subtomentose

on

the d

surface, abundantly pustulate, ventral surface

setose and subto-

rather sparsely pustulate, weakly appressed-setose
mentose, at times subsericeous ; inflorescence extend
or more of the stem, very floriferous, not interrupted, becoming i
broad thyrsus in which the individual cymes are much elongated
densely setose, in age usually fulvous, lower foliar bracts ofter
conspicuous and somewhat reflexed; calyx densely setose, sepal
in anthesis lanceolate, about 4 mm. lone, acute, in fruit linear

1927]

PAYSON — SECTION OREOCARYA OF 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 \4 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. Type: La
Verken, Utah, May 8, 1894, M. E. Jones 5195a.

Specimens examined:

Utah: Valley of the Virgin near St. George, 1874, Parry 173
(Field, N.Y., Gray); Diamond Valley, May 16, 1902, Goodding
830 (R.Mt., Pomona, Mo., U.S., Gray); Diamond Valley, April
28, 1894, Jones 5125 (R.Mt., Pomona, Calif., Mo., U.S.); La
Verken, May 8, 1894, Jones 5195a (Pomona, type, R.Mt., Calif.,

Mo., U.S.).

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,

K. Brandegee (Pomona) ; Gold Mountain, 1898, Purpus (Calif.) ;

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).

C. mrainensis is in general appearance more like C. thyrsiflora

of

be

fused with that species because of the great distance between

their

of the two species are entirely dif

ferent, and it seems quite doubtful if they

[Vol. 14
276 ANNALS OF THE MISSOURI BOTANICAL GARDEN

related. Its nearest relatives are probably C. twnulosa, C. mo-
desta, and C. insolita. The distinguishing characteristics of these

contrasted under C

16.

comb. Plate 27, fies. 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 cm. 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 sliehtlv more pus-

than the ventral; inflorescence extending over 34 to

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.

8-10 mm. long, exceeding the nutlets by 4-6 mm.;

fruit

a white, tube 3.5-4 mm. long, equalling or slightly shorter
the sepals, crests at the base of the tube evident but not
icuous, fornices conspicuous, probably yellow, about 1 mm.
limb 7 mm. broad, limb and tube subequal, lobes united

for about y% 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, T. S. Bran-
degee (Calif., type) ; Providence Mts., May, 1892, T. S. Brandegee

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 277

(Phila.) ; Barnwell, May 14, 1911, K. 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 C. humilis and C. nubigena.
The nutlets are quite different from either of those species. Its
nearest relatives are probably C. virginensis, C. insolita and C.
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

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 cm. long,
closely tomentose and setose with weak appressed bristles, pus-
tulate on both surfaces, more abundantly on dorsal, petioles about
as lone as the blades, conspicuously ciliate near the base; inflores-

cence extending over ]/ 2 to % oi 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 M 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)

278 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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. Type: Aurum, Nevada, alt. 7,300 ft., June 20,
1893, M. E. Jones

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 6718 (Pomona).

This species is perhaps most closely related to tumulosa, vir-
ginensis, and insolita. In general appearance it is not unlike

some of the varieties of C

18. C. humilis (Greene) new comb

47-49

ium glomeratum var. (?) fulvocanescens Wats. Bot.

King's Exp. 243. 1871.

E. 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.

0. Macbridii Brand, Fedde, Rep. Sp. Nov. 19: 73. 1923.
(Type: Mt. Jarbidge, Nevada, July 6, 1912, Nelson & Macbride
1960.)

Caespitose perennial ; stems 1-several from the woody caudex,
rather stout or slender, 10-30 cm. high, setose with rather weak
bristles; leaves obovate-spatulate to oblanceolate, 3-8 cm. 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 OF CRYPTANTHA 279

or % 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, papillose,
probably pale yellow, limb 8-10 mm. broad, lobes and tube
subequal, lobes united for about }/i 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. Type not desig-
nated: "Frequent in the mountains of Nevada and adjacent
eastern California: the Calif ornian 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 (Po-
mona); Monitor Valley, July, 1868, Watson 853, 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., May 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-

280 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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
529/f (U.S.); 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. No 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. C. 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 Truckee 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

rather weak. The rugae tend to be
reduced to scarcely continuous tubercles or indistinct wrinkles.

seems

of these forms that Macbride classified

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 a
new species for it — 0. Macbridii. An examination of a 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. commixta 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. C. 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

confined to upper Yr^A of the stem, foliar bracts

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 3 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 }4r l A their length; fruit

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

(Vol. 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. Type (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 6448 (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, Tweedy 4260 (U.S.); Ft. Steele, June 16, 1900, Nelson 7255
(R.Mt., Gray); Point of Rocks, June 15, 1898, Nelson 4749
(R.Mt., type, U.S.); Point of Rocks, June 19, 1901, Merrill &
Wilcox 617 (R.Mt., U.S., Gray) ; Red Desert, June 3, 1897, Nelson
8120 (R.Mt.); Bush Ranch, Sweetwater Co., June 10, 1900,
Nelson 7078 (R.Mt., Pomona, Minn., Mo., 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 on Gros
Ventre Fork, June 10, 1860, Hayden (Mo.); between Eden & Big
Piney, July 6, 1922, Payson & Payson 2578 (R.Mt., Pomona,
Mo., Gray) ; between Opal and Kemmerer, June 19, 1923, Payson
& Armstrong 8223 (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. C. 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. Bradburianae. Nutlets always roughened dorsally
and distinctly rugose or tuberculate, or both, and often muricate

1927]

PAYSON — SECTION OREOCARYA OF 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.

0. 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).

0. 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
1-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
i 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.

i

[Vol. 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. Type: "On 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 (Mo.) ; 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 (R. 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,

& Vreeland 5704

Sheldon 526 (U.S

na Vista, July 6,
2, 1919, Eggleston

15379 (Field); Trinidad, June 28, 1898, Osterhout (R. 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. G. 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 in
Bradburiana. The 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. Sheldonii.

A 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 cm. 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 3^ or

£ the length of the stem, upper cymes elongating, the lower re-
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-

[Vol. 14

286 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ceeding the nutlets by 2-3 mm. ; corolla white, tube about 4 mm
long, equalling the sepals, crests at the base of the tube wel
developed, fornices nearly 1 mm. high, distinctly papillose
probably yellow, limb about 9 mm. broad, lobes and tube sub

lobes united for about % 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. Type: "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."

Specimens examined:

Colorado: Grand Junction, May 15, 1892, Eastwood (N.Y.,
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. In 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 % to % the length

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 287

of the stem, in flower narrow, somewhat glomerate, abundantly
setose, foliar bracts 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 % 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 of the cy-
mules; leaves oblanceolate; nutlets 3-3.5 mm. long 37a. var. typica

Inflorescence usually narrow, even in age; leaves narrowly oblanceolate; nut-
lets 2.75-3 mm. long 37b. var. perennis

Var

Plate 28, figs

Eritrichium glomeratum var. humile Gray, Proc. Am. Acad. 10:
61. 1875, in large part.

Krynitzkia sericea Gray, Proc. Am. Acad. 20: 279. 1885, in
part.

Oreocarya sericea Greene, Pittonia 1: 58. 1887.

0. argentea Rydb. Bull. Torr. Bot. Club 31: 637. 1904. (Type:
Rifle, Garfield Co., Colo., 1900, Osterhout 2122).

0. 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

[Vol. 14

288 ANNALS OF 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, type, Mo.); Pass
Creek to Bridger's Pass, 1859, H. Engelmann (N. Y., Gray); 11
miles north of Encampment, July 2, 1922, Pay son & Pay son
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.); Glen wood 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.,

U.S.).

22b. Var. perennis (A. 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 cm. 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, Pay son
& 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., type, Mo., Gray); Green River, May
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

(U.S.).

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. The evidence
on which this conclusion is based is presented in detail in the
following paragraphs.

[Vol. 14

290 ANNALS OF THE MISSOURI BOTANICAL GARDEN

First of all it is useful to summarize the criteria that have

governed the

specimen on which

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

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. The type should be represented by a 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. The 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 is a 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 OREOCAKYA OF CRYPTANTHA

291

mountains from Colorado and Utah to Oregon and Montana."
6. Dr. Gray says, "I 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.

The next point of attack is 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

b. W. side of Wasatch Mts., 1844, Fremont

0. argentea Rydb.
0. humilis Greend, K. de-
pressa Jones, or sotne form
of 0. nana Eastwood. Too
immature for certaih iden-
tification.

c. Clover Mts., Nevada, 1868, Watson 853

0. humilis Greene

de-

pressa Jones, or some form
of 0. nana Eastwood. Too
immature for certain iden-
tification.

d. Mountain Hot Springs, Yellowstone Park, 1885,

Tweedy 816 = C. Bradburiana.

Sheet 2 contains 3 specimens:

a. Montana Terr. 1867

It also bears the two separate notations in
Dr. Gray's handwriting "E. glomeratum" and
"Probably dwarf K. glomerata, JJ

b. Summit, California, 1871, Bolander

C. Bradburiana.

c. Grass Valley, Utah, 1875, Ward 49

humilis or possibly nubi-
gena. Specimens immature.
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 identification
practically certain.

b. A specimen of Bradburiana without data. A note by Dr. Gray says " origin un-

certain.

>»

292 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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 sheet 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 note in Dr. Gray's
handwriting: "Two more dwarf and very sericeous-canescent 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 OF CRYPTANTHA 293

Gray containing this description were presented to the Academy
May 12 and Oct. 13, 1874." We find Dr. Gray's notation

. t

(I

E. 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 "W. 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." The nutlets were unknown to Dr. Gray, and if they
had been known would not have been described as "subrugose-
tuberculate." The range given by Gray for sericea is scarcely
large enough to include cana. Scott's Bluffs, from where he had
a specimen of cana, is in Nebraska and so hardly to be described
as " 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

[Vol. 14

294 ANNALS OF 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. The 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. The 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 Oreocarya 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. The 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 0. argentea
Rydb.

It is possible that further collections of 0. procera Osterhout
and of Jones' plant from Utah may prove them worthy of varietal
rank. The 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. Dr. 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 OF CRYPTANTHA 295

23. C. 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, May 17, 1892, Eastwood (Calif.).

C. aperta is 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.

[Vol. 14

296 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Short-lived perennial; stems few to many, unbranched above
the base, rather slender, 15-30 cm. high, rather sparsely but con-
spicuously setose; leaves narrowly oblanceolate or spatulate,
obtuse, 2-4 cm. 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 ^ or 3^ 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 }4 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. Type:

Fish Springs, Utah, M. E. Jones.
Specimen examined:
Utah: Fish Springs, June 4, 1891, Jones (R.Mt., type, Pomona,

Calif.).
The 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.

comb. Plate 28, figs. 68-70

Oreocarya interrupta Greene, Pittonia 3: 111. 1896.

Long-lived, caespitose perennial; stems few to many from
branching caudex, rather slender, 3-9 dm. high, rather spar
strieose and setose with slender white trichomes: radical le;

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 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 Yi-Yz 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 3 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 \i 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: "in 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

and

has not been possibl

to locate the type but specimens from near the type locality ag
very well with the original description and there is probably

[Vol. 14

298 ANNALS OF THE MISSOURI BOTANICAL GARDEN

doubt but that the species has been correctly identified. No
specimens have been seen that were so high as the plants de-
scribed by Greene. He says "1^ to 3 feet high." The speci-
mens at hand are ail 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.

0. cilio-hirsuta Nels. & Macbr. Bot. Gaz. 55: 378. 1913.
(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 cm. 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 on the upper 3^ or ^ 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 l /i 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 3228 (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., type, Gray).
C. spiculifera is a 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. Macounii, 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, Macounii, spicu-
lifera, interrupta, rugulosa, Bradburiana, sobolifera — that it
seemed unwise. C. spiculifera has some contact, too, with
C. nana var. Shantzii. A specimen from southeastern Idaho
was examined that looked remarkably like spiculifera but the
nutlets were uniformly muricate as in the var. Shantzii. 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 with 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 onlv the petioles of

—or only the petioles of former leaves remaining to
j the basal tuft — spatulate to oblanceolate, usually obtuse,
cm. long, densely setose with spreading white bristles and

300 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

subtomentose, densely pustulate on both surfaces, petioles con-
spicuously ciliate; inflorescence extending over upper y% to %
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 l /i to % 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. Type: " 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, W kited (Wash.) ; Wenatchee, May 28, 1899, W kited 1099
(Wash., U.S., Gray).

Oregon: Dalles, May, 1882, Howell (Phila.); Mitchell, Aug. 1,
1917, Lawrence 1030 (U.S.); near Fossil, Gilliam Co., May 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. Macounii. 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. The 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. sviculifera.

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.

0. celosioides Macbr. Contr. Gray Herb. 48: 29. 1916, in part,
as to specimens cited, not 0. celosioides Eastw.

0. glomerata Standley, Contr. U. S. Nat. Herb. 22: 401. 1921,
not Cynoglossum glorneratum Pursh.

0. Sheldonii Brand, Fedde, Rep. Sp. Nov. 19: 73. 1923.

Definitely perennial, rather loosely caespitose; stem rather
stout, 2-3 cm. high, abundantly setose with rather slender, di-
varicate hairs; basal leaves numerous, spatulate to oblanceolate,
usually obtuse, 2-4 cm. 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 14 to Va 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 Yz their length ; fruit ovoid or lanceolate-
ovoid, all four nutlets commonly maturing, style exceeding the

302 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

nutlets by about 2 mm.; nutlets lanceolate, acute or subacute,
3-4 mm. long, margins in 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. Type:
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,
Standby 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, E. N. Brandegee (Calif.); Hallett's
Ranch, Aug. 14, 1892, E. N. Brandegee 17 (Calif.); Helena, June
& July, 1892, Alton 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, A. & E. 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, Alton 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, Turesson (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., U.S.).

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.

19271

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. C. Macounii is the other perennial
relative of C. Bradburiana. That plant is characterized by nar-
row, strongly setose leaves, while Sheldonii 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. To 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
0. sericea which must be used to designate a very different plant
of the Rocky Mountains. " With sericea now identified with
Rydberg's argentea y 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.
1913.

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
(particularly on the petioles), leaf-surfaces strigose, more densely
pustulate below than above; inflorescence narrow, mainly

limited to upper % or ^ 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,

304 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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 \i 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 a 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. Type: 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., type, 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
7674V (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. Sheldonii.

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 305

Sept. 10, 1909, Jones 1 (Pomona); Lewis & Clark Nat'l. Forest on
Beaver Creek, Aug. 6, 1912, Saunders 1 (U.S.); near Helena, June,
1921, Wooton 1 (U.S.); Helena, June, 1893, Start* (Mo.); Helena,
1886, Kelsey 102 1 (Pomona); Helena, June, 1888, Kelsey 1 (U.S.);
plains about Helena, June, 1887, Anderson 1 (Calif., Mo.); Mid-
vale, June 24, 1903, Umbach 149 1 (U.S.) ; Garrison, July 10, 1909,
Jones 1 (Pomona); Armstead, June 20, 1920, Payson & Payson
1746 2 (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 359 1 (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 3 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 Macounii
on the basis of a more robust habit.

2 This is not a typical C. Macounii, but is intermediate to C. Sheldonii.

3 Cryptantha sobolifera sp. nov., perennis; caulibus sterilibus prostratis ad apicem
foliatis, caulibus floriferis 15-18 cm. altis hirsutis et setosis; foliis obovato-spathulatis
obtusis 1.5-3.5 cm. longis dense strigosis aut subtomentosis et insigniter setoso-
hirsutis supra et subter 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. longioribus; corolla alba, tubo 3-4 mm. longo,
calycem non superante, limbo 7 mm. lato; stylo nuculas 1.5-2 mm. superante; nuculis
non nitidis ovato-lanceolatis subacutis 3 mm. longis, 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).

[Vol. 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 cm. 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 ^ or Y2 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 in anthesis, 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 %-
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. Type:
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. The 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 OF 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 denned by Jones as the spruce belt. The 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,
>t Cryptantha glomerata Lehm.

Myosotis glomerata (Pursh) Nutt. FL Am. Sept. 2: 729. 1814.
Rochelia glomerata (Pursh) Torr. Ann. Lye. N. Y. 2: 226. 1828.
Eritrichium glomeratum (Pursh) DC. Prodr. 10: 131. 1846.
Krynitzkia glomerata (Pursh) Gray, Proc. Am. Acad. 20: 279.

1885.

Oreocarya glomerata (Pursh) Greene, Pittonia 1: 58. 1887

0. affinis Greene, Pittonia

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 0. 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 cm. 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
of the stem, foliar bracts near the base of the inflorescence

nspicuous and longer than the cymules in the young inflores-

densely setose and hirsute, seps
hesis about 4 mm. loner, in fruit

mm

ceeding the nutlets about 5 mm.: corolla white, tube 3-4 mm

[Vol. 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 % their length; fruit ovoid

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: "in
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 431 (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 (U.S.);
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 (U.S.).

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., May 15-31, 1912, Eggleston
7838 (U.S.); Livingston, June 7, 1901, Scheuber (Minn., U.S.);
Bozeman, May 19, 1900, Moore (Calif.); Bozeman, June 16,
1905, Blankinship 373 (U.S.); 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 4882 1 (R.Mt., Wash., Minn., U.S., Mont.,
Gray); Spanish Basin, Gallatin Co., June 23, 1897, Rydberg &
Bessey 4883 1 (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 (Mo.); 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 941 1 (R. Mt., Minn., Gray); Hartville,
June 30, 1901, Nelson 8319 (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.

310 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

Hills, June 18, 1890, 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 4887
(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 (i.e.,
Bradburiana) that are in reality quite different plants. A 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 9 Sheldonii, 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 CRYPTANTHA 311

specific boundaries further and include humilis, sericea, and
aperta. 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 a 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. Sheldonii 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 Sheldonii 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, 0. 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
a familiar one but since Cryptantha glomerata Lehm., which is a
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.

The 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.

[Vol. 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
a 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 shorter 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 }/% 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.

The 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
I Itah and eastern Nevada 32a. var. commixta

Stems 10-20 cm. high; leaves densely tomentose and appressed-setose; styles
scarcely exceeding the mature nutlets; north-central Utah and south-
eastern Idaho 32b. 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 32c. 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 Utah 32d. var. typica

32a. Var. commixta (Macbride) new comb.

Oreocarya commixta Macbride, Contr. Gray Herb. 48: 33. 1916.

19271

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 313

Perennial; stems 15-25 cm. high; leaves spatulate, obtuse, 3-5
cm. 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. Type: 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, type) ; 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

1 605) .

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.

314 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

Specimens examined:

Idaho: Pocatello, June 15, 1893, Palmer 598 (U.S.) ; Pocatello,
spring of 1921, Soth P-9 (R.Mt.); Pocatello, May 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
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, May 5, 1880, Jones 1692 (R.Mt., Calif., U.S.); Grant's
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. 1

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.

1 Cryptantha nana var. ovina Payson var. nov., perennia caespitosa; caulibus 10-
15 cm. altis; foliis spathulatis aut oblanceolatis toraentosis et adpresse setulosis;
sepalia dense tomentosis; nuculis muriculatis; stylo nuculas 0.5-1 mm. superante.

Collected by Georgia 11. Bentley, in vicinity of Currant, Nye County, Nevada,
June, 1916, (R. Mt., type).

1927J

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 315

Distribution: Upper Sonoran Zone, southern Utah, and
southern Nevada. Type: vicinity of Currant, Nye County,
Nevada, June, 1916, Georgia H. 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 Shantzii,

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

[Vol. 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 no 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. They seem to be
geographic units and so are worthy of discrimination in a more

technical treatment.

33. C. cana (A. Nels.) new comb. Plate 29, figs. 92-94.

Oreocarya carta A. 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 cm. 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 x /i °f 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 x /± 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, Bodin (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., type, Gray); Platte Hills, Nutlall (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. (Type: 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 cm. 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 Y^ or % of the stem, densely but rather weakly setose,
foliar bracts not conspicuous; calyx densely and weakly setose

318 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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 }£ 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. Type: 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 (Calif., U.S., Gray, type); Vale, Malheur Co., May
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. carta
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 cm. long, uniformly

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 319

1

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
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 }4 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: 63^ miles north of Jensen

(Utah), Osterhout6414-

Specimens examined:

Utah: 63^ miles north of Jensen, June 19, 1925, Osterhout 6414
(R.Mt., type, Osterh.); Uinta Mts., 1902, Langille 113 (N.Y.,
U.S.); Ft. Duchesne, May 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.
0. nitida Greene, PI. Baker. 3: 21. 1901. (Type: in dry stony
ground at Deer Run, Colorado, 11 June, C. F. Baker 95.)

Perennial and caespitose from a woody root; stems few to

[Vol. 14

320 ANNALS OF 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 cm. long,
uniformly silky strigose, pustulate hairs scarcely differentiated
and difficult of observation in dried material, pustules numerous
on dorsal surface, few and small on ventral surface; inflorescence

rather narrow, nearly continuous, on upper Yi ° r Z A 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 )4 as
long as the tube, lobes united for l /i 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 % 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) ; West water, May 6,
1891, Jones (Pomona); Naturita, May 4, 1914, Pay son 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 OREOCARY> 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., U.S.).

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 % 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 \i~yz their length;

[Vol. 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. Type: Pahria Canyon, Utah, 5300 ft. alt., in
red sand, M. E. Jones 5279p.

Specimens examined:

Utah: Pahria Canyon, May 26, 1894, Jones 5297p (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 is 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 (Eri-
trichium glomeratum var. fulvocanescens Wats. Bot. King's Exp.
243. 1871) as being the plant that must bear the specific name
fulvocanescens. The first legal publication of this name is 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 cm. long, obtuse, blade abruptly or
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 Yi 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 3^ as long as the tube,

united for about 34 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,

324 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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 / 6 of distance
to apex, no elevated margin.

Distribution: Upper Sonoran Zone in south-central Utah.
Type: 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. Fla voculatae . Nutlets usually deeply and con-
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 CRYPT ANTHA 325

usually acute, 1.5-5 cm. long; inflorescence on upper % 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 % their length; fruit ovoid, 3^1 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. Type:
"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 type); Cisco, May 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. Wetherillii 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. The differences between the two species
may be contrasted as follows :

C. 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).

[Vol. 14

326

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fornices long-papillose or " his- Fornices scarcely papillose at

pid."

Corolla tube 6-7 mm. long.
Nutlets tuberculate and more
or less rugose.

all.
Corolla tube 12-14 mm. long.
Nutlets distinctly rugose,

scarcely tuberculate.

Scar open but narrow, slightly Scar very narrow, sides nearly

broader near the middle,
surrounded by a distinct
elevated margin.

parallel, elevated margin
not evident or very slight.

40. C. longiflora (A. Nels.) new comb. Plate 30, figs. 104-106.

Oreocarya longiflora A. Nels. Erythea 7: 67. 1899.

0. horridula Greene, PL Baker. 3: 20. 1901. (Type: Deer
Run, Colorado, 11 June, on a dry bank, C. F. Baker 133.)

A 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 cm. 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 % 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 10 mm. in diameter,

lobes VL to ]4 as long as the tube, united about l /i 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 327

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 4463
(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., type); Westwater,
May 6, 1891, Jones (Pomona); Montrose, May 23, 1912, Pay son
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.
Wetherillii.

0. horridula Greene has not been seen except for a few frag-
ments from the type. A 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 Yi to % 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. ;

[Vol. 14

328 ANNALS OF THE MISSOURI BOTANICAL GARDEN

corolla white, tube 6 mm. long, slightly exceeding the sepals in
an thesis (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-
allv roimnamilate for the eenus, about 5 mm. across, tube 1.5-2

mb, lobes united for about \4 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. No evidence is at hand to show that this species is di-
morphic 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-Oct. 1898, Purpus (Calif.) ; Court
House Wash., s.e. Utah, May 25, 1892, Eastwood (R.Mt., Calif.,

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 Wetherillii (from which it differs
by the narrower leaves, sharply rugose nutlets, and much smaller
flowers) .

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 329

42.

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
cm. 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 cm. 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 % 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 \i
their length; fruit broadly ovoid, in the material examined only
1 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 ^

the length of the nutlet, evident, elevated and tuberculate margin
present.

Distribution: Upper Sonoran Zone in western Colorado and
eastern Utah. Type: Monument Park, near Grand Junction,
Colorado, Geo. E. Osterhout 6188.

Specimens examined:

Colorado: Grand Junction, Monument Park, June 3, 1921,
Osterhout 6188 (R.Mt., type, 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

[Vol. 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 (A. Nels.) new comb. Plate 30, figs. 113-115.
Oreocarya paradoxa A. Nelson, Bot. Gaz. 56: 69. 1913.
0. gypsophila Payson, Bot. Gaz. 60: 380. 1915. (Type: "on
dry gypsum hill in Paradox Valley, Colorado, alt. slightly over

ft., Payson ^58, June

>>

Densely caespitose perennial; caudices much branched and
clothed with the leaf-bases of previous years; stems slender, 5-12
cm. 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 cm. 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

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 2y 2 times as long as the lobes,

lobes united for about M their length; fruit oblate-ovoid

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

the length of the nutlet, margin slightly elevated.
Distribution: Upper Sonoran Zone, western Colorado, and
eastern Utah. Type: on dry "gyp" hills, in Paradox Valley,
Colorado. E. P. Walker 91.

1927]

PAYSON — SECTION OREOCARYA OF CRYPT ANTHA 331

Specimens examined:

Colorado: Paradox, Montrose Co., June 17, 1912, Walker 91
(R.Mt., type, 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: My ton, 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 paradoxa the anthers are described as
being "just below the crests." On 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.
0. eulophus Rydb. Bull. Torr. Bot. Club 31: 637. 1904.
(Type: Dolores, Colorado, 1892, Crandall)

[Vol. 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 cm. 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 4 /b, 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 l / 2
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 curved or twisted, closed, surrounded by an elevated,
tuberculate margin.

Distribution: Transition or higher Upper Sonoran Zone in
southwestern Colorado and southeastern Utah. Type : "Collected
on the Mancos River sage plains in southern Colorado, by Messrs.

827.

& Tracy, 8 July, 1898, and distributed under

>?

Specimens examined:

Colorado: Meeker, June 8, 1902, Osterhout 2610 (Osterh.);
Gunnison, July, 1897, Bethel (Colo.); Gunnison Co., June, 1888,
Eastwood (U.S.) ; 3 miles south of Ridgway, July 16, 1917, Payson
1079 (R.Mt., Mo., Gray); Ridgway, June 18, 1924, Payson &
Payson 8834 (R.Mt., Mo., Gray); Dolores, June 16, 1892,

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA 333

Crandall (N.Y.); Dolores, June 15, 1892, Herb. Colo. Agric. Coll
(N.Y.) ; Mancos, 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. They 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. The very low position of the
anthers makes it seem probable that another form with anthers
near the top of the tube will be found. The position of the
stigma, however, is that which usually occurs in the species with
only one kind of flower.

This species is a 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
*U-*/b of the stem, foliar bracts large and well-developed; calyx
strigose and conspicuously setose, sepals in anthesis linear-
jolate, 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.

334 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

long, equalling or shorter than the sepals in anthesis, crests a
the base of the tube nearly but not quite obsolete, fornices prob

ably

mm. long, slightly papillose, limb 6-8

mm. broad, lobes united for about }/$ 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, May 16, 1894, M. E. Jones 5445p.

Specimens examined :

Utah: Emery, June 16, 1894, Jones 5445 p (Pomona, type,
U.S.); San Rafael Swell, May 15, 1914, Jones (Pomona).

C. mensana is certainly most closely related to C. 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. C. flavoculata (A. Nels.) new comb. Plate 30, figs. 122-124.

Oreocarya flavoculata A. Nels. Erythea 7: 66. 1899.

0. flavoculata spatulata A. Nels. Erythea 7: 67. 1899. (Type:
from gravelly hill tops near Evanston, Wyoming, Nelson 2977,
May 29, 1897.)

0. cristata Eastw. Bull. Torr. Bot. Club 30: 244. 1903. (Type :
Grand Junction, Colo., May 17, 1892, Eastwood.)

0. Shockleyi Eastw. Bull. Torr. Bot. Club 30: 245. 1903.

(Type: Miller Mountain, Esmeralda Co., Nevada, elev. 7500 ft.,
Shockley 244)

0. 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 OF 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 as on the dorsal surface; inflorescence continuous or
somewhat interrupted, rather narrow, usually extending over

upper Yi or % 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 M their length, V> to 2 A 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

336 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

& 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, Tweedy 4259 (U.S.); 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

(R.Mt.); Granger, June 13, 1898, Nelson 4749a (R.Mt.); Kern-
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., type, 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.); McCoys, 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, May 6, 1891, Jones 6715
(Pomona); Grand Junction, May 17, 1892, Eastwood (Calif.);
Naturita, June 20, 1924, Payson & Payson 3865 (R.Mt., Mo.,
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

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,

1927J

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 52041 (Pomona,
U.S., in part) ; San Pitch Mts., June 24, 1908, Tidestrom 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 (U.S.); 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 ft., 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 (Mo.).

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 specific 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 a basis of setose-hispid and silky-strigose

338

[Vol. 14

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 a good match for the type of cristata. 0. Shockleyi 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 0. Shockleyi, nor can the
author see that the nutlets are different.

The difference between the stamen insertion in the two forms

bably never more than 3 mm. and

less than that.

Index to Specimens Cited

1 n the following index the numbers in italics outside the parenthesis are the col-
lector's numbers. Those in black type in the parenthesis are the numbers assigned
to the species in the present monograph.

Aiton, G. B.

31, 64 (28) C. Sheldonii.
Anderson, F. W.

(29) C. Macounii.
Anect, Bro.

164 (l a ) C. Jamesii var. multicaulis.
Austin, Mrs. R. M. & Bruce.

2270 (10) C. nuhigena.
Austin, S. W.

— ■ (5) C. conjerti flora.
Bailey, V.

14 (31) C. Bradhuriana.
Bailey, V., F. V. Coville & F. Funston. Bethel, E.

Barber, M. A.

325 (29) C. Macounii.

Bates, J. M.
5768, 5797 (le) C. Jamesii var.
typica; — (31) C. Bradhuriana; — (33)
C. carta.

Beals, Mrs. W. G.

(4) C. oblata.

Beattie, R. Kent.

3921 (7) C. leucophaea.

Bentley, Georgia H.

— (32c) C. nana var. ovina.

(11) C. Clemensae; 1888 (15) C.

virgmensis.

Baker, C. F.

(6) C. flava; — (44) C. Bakeri.
Bethel, E., F. S. Nielley & I. W. Clokey.
4258 (le) C. Jamesii var. typica.

563 (la) C. Jamesii var. multicaulis; Blankinship, J. W.

455 (lc) 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.

13 (20) C. thyrsi flora; 827 (44) C. Bakeri.
Barber, II. S.

112 (lc) C. Jamesii var. cinerea.

734 (28) C. Sheldonii; 372a, 373,
(31) C. Bradhuriana.

Bodin, J. E.

(le) C. Jamesii var. typica;

(20)

C. thyrsiflora;
Bourgeau, E.

(33) C. cana.

(29) C. Macounii.

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA

339

Bradbury, John.

(31) C Bradburiana.

Brandegee, E. N.

— (28) C. SheUonii.

17

Brandegee, K.

Cotton, J. S.

1027 (7) C. leucophaea; 359 (29) C.
Macounii.

Coville, F. V.

151 1+ (10) C. nubigena.

(5) C. confertiflora; — (14) C. in- Coville, F. V. & F. Funston

solita;

(15) C. virginensis;

(16)

C. tumulosa.

Brandegee, T. S.

#f 51 (46) C. flavoculata.
Coville, F. V. & J. B. Leiberg
91 (18) C. humilis.

(lc) C. Jamesii var. cinerea; 997 Cowen, J. H.

(7) C leucophaea; 898 (13) C. virgata;

(lc) C Jamesii var. cinerea.

(16) C. tumulosa; 4 (20) C. %m- Crandall, C. S.

/tora; 536? (27) C. celosioides .

Brewer, W. H.

/«M (5) C. confertiflora; 1887 (10) C.

Bryan, Lieut. F. T.

(46) C flavoculata.

Buffum, B. C.
640 (31) C. Bradburiana; 642 (46) C.

flavoculata.

Carlton, M. A.

246 (le) C. Jamesii var. typica.

Carlson, John I.

(4) C. oblata.

Cary, Merritt
276 (33) C. cana.

Chestnut, V. K. & E. R. Drew

— (10) C. nubigena.

Clemens, Mrs. Joseph

(11) C. Clemensae; — (32b) C. nana
var. Shantzii.

Clements, F. E. & E. S.

102 (13) C. virgata.

Clifton, R. L.

3039 (la) C. Jamesii var. multicaulis.

Clute, W. N.

37 (Id) C. Jamesii var. disticha.

Cockerell, T. D. A.

(le) C. Jamesii var. typica.
Colo. Agr. College Herb.

(44) C. Bakeri.

Condit, I. J.

(le) C. Jamesii var. typica

Cooper, W. S.

117 (13) C. virgata.
Copeland, E. B.

52 (11) C. Clemensae.

(le) C. Jamesii var. typica;

(6)

C. flava; — (13) C. virgata; — (40) C.

longiflora;
Cusick, Wm.

(44) C. Bakeri.

2028,

(10) C. nubigena.

Davidson, A.

608 (la) C. Jamesii var. multicaulis;

2722 (5) C. confertiflora.

Dawson, G. M.

17088 (29) C. Macounii.

Dodds, G. S.

1832 (13) C. virgata; 2077 (20) C.

thyrsiflora.
Douglas, David.

— (7) C. leucophaea.
Drummond, Thomas.

(29) C. Macounii.
Eastwood, Alice.

8366 (la) C. Jamesii var. multicaulis;
(Id) C. Jamesii var. disticha;

(le) C. Jamesii var. typica;

(6) C.

flava;

(21) C. elata; — (23) C. aperta;

(32d) C. nana var. typica; 5596 }

(36) C. fulvocanescens ;

(39) C.

(40) C. longiflora;
- (44) C. Bakeri;

Wetherillii; —
(41) C. tenuis;
(46) C. flavoculata.

Eggert, H.

— (la) C. Jamesii var. multicaulis;

— (3) C. Palmeri.

Eggleston, W. W.

16184 (1^) C. Jamesii var. multicaulis;
11194a, 15168 (le) C. Jamesii var.
typica; 15379 (20) C. thyrsiflora; 7838,
7959, 12520 (31) C. Bradburiana;
11132 (32b) C. nana var. Shantzii;
11118 (46) C. flavoculata.

340

[Vol. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Ellis, Charlotte C.

866, JfiS (la) C. Jamesii var. multi-

caulis.
Elmer, A. D. E.

1056 (7) C. leucophaea.
Engelmann, G.

(10) C. nubigena; — (20) C. thyrsi- Hanson, H. E.

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.

flora.
Engelmann, H

(20) C. thyrsiflora;
Evermann, B. W.

— (20) C. thyrsiflora.

(22) C.sericea. H avard, V.

A llfi (la) C. Jamesii var. multicaulis;
899 (4) C. oblata.

(la) C. Jamesii var. multicaulis;

Fendler, A.

686 (la) C. Jamesii var. multicaulis;
632 (36) C. fulvocanescens.

Ferris, R. & R. Duthie.

(lb) C. Jamesii var. laxa; —
oblata; — (31) C. Bradburiana.

Hayden, F. V.

(4) C.

34 (10) C. nubigena.
Fin ley, C.

2 (46) C. flavoculata.
Flodman, J. II.

748 (30) C. sobolifera.
Forwood, W. H.

— (19) C. caesjritosa; — (28) C. Shel-
donii.

Heller, A. A.

11016 (If) C. Jamesii var. abortiva;

8211, 9619 (5) C. confertiflora; 13435

(10) C. nubigena;

(18) C. humilis;

272,

(31) C. Bradburiana;

Garrett, A. O.

266, 1737 (32b) C. nana var. Shantzii.
Geyer, C. A.

(31) C. Bradburiana.
Gilbert, H.

(46) C. flavoculata.
Goldman, E. A.

407 (la) C. Jamesii var. multicaulis.
Goodding, L. N.

9185 (25) C. interrupta; 10508 (46) C.
flavoculata.

Heller, A. A. & E. G.

3577 (lc) C. Jamesii var. cinerea;
3731 (20) C. thyrsiflora; 3517 (36) C.
fulvocanescens.

Henderson, L. F.
5212 (27) C. celosioides; 2563 (28) C.
Sheldonii.

Hess, W. L.

106 (le) C. Jamesii var. ty\rica.

1264, ^349 (la) C. Jamesii var. mul- Hick s, Mr. & Mrs. G. H.

ticaidis; 208 (le) C. Jamesii var.

178 (lc) C. Jamesii var. cinerea.

tyj/ica; 814, ^221 (5) C. confertiflora; Hill, G. R., Jr.

53,164 (6) C. flava; 589 (12) C. setosis-

(32b) C. nana var. Shantzii.

sima; 2286 (14) C. insolita; 830 (15) Hindshaw, H. H.

C. virginensis; 996 (17) C modesta;

2 (7) C. leucophaea.

60 (19) C. caespitosa; 2076 (20) C. Hitchcock, A. E.

thyrsiflora; 29, 230, 287 (31) C. firad-

buriana; 1074 (32a) C. nana var.
com mixta; 1282 (46) (7. flavoculata.
Greene, E. L.
82 (la) C. Jamesii var. multicaulis;

1193 (lc) C. Jamesii var. cinerea;
7 (6) C. ^aya; S2P, JW5 (25) C. in-
terrupta; 755 (46) (7. flavoculata.
Hitchcock, A. S.

347 (le) C. Jamesii var. typica.

(lc) C. Jamesii var. cinerea; 300 Holzinger, G. A.

(12)

(le) C. Jamesii var. ty\nca; —
C. setosissima; — (18) C. humilis.
Griffiths, David.

4268, 4912 (la) C. Jamesii var. wwZtt-

cmdis; 5300 (12) C. setosissima.

6 (31) C. Bradburiana.
Hough, W.

S (lc) C. Jamesii var. cinerea; 88 (12)
C. setosissima; 39 (36) C fulvocanes-
cens .

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA

341

Howell, Thos.

481,

(27) C. celosioides.

Jaeger, E. C.

— (la) C. Jamesii var. multicaulis ;
1113 (15) C. virginensis.

James, Edwin.

— (le) C. Jamesii var. typica.

Johnston, E. L.

845 (le) C. Jamesii var. typica.
Johnston, I. M.

Jones, W. W.

339 (la) C. Jamesii var. multicaulis
Kearney, T. H. & H. L. Shantz.

3098 (32b) C. nana var. Shantzii.
Kellerman, W. A.

6 (le) C. Jamesii var. typica.
Kellogg, A.

— (18) C. humilis.
Kelsey, F. D.

102,

Maco

(le) C. Jamesii var. typica; 2818 Kennedy, P. B.

(13) C. virgata;2817 (20) C. thyrsiflora.

Jones, B. J.

Bradburia

Jones, M. E.

4007, 4047, 5297q, 5298b, 5300, 6685,

6687,

1664 (18) C. humilis; 4519 (46) C.
flavoculata.
Kennedy, P. B. & L. N. Goodding.
110 (5) C. confertiflora; 78 (15) C. vir-
ginensis; 146 (46) C '. flavoculata.

(la) C. Jamesii var. multi- Kirkwood, J. E.

caulis; 4042, 5328, 6684, 6688,

(le)

1262 (28) C. Sheldonii.

C. Jamesii var. cinerea; 66, — (le) C. Kirtley

Jamesii var. typica; 6S15f, — (If) C.

(8) C. salmonensis.

Jamesii var. abortiva; 3759 (4) C. ob- Knowlton, F. H.

lata; 5144, 5261 j, 5289t, — (5) C. con-

35 (20) C. thyrsiflora.

fertiflora; 5455c, 5464, 6725, — (6) C. Kunze, R. E.

jlava; 5695al, 5790m, 5812, 601 5al,
6054s, 6686 (12) C. setosissima; 296, —
(13) C. virgata; — (14) C. insolita; 5125,

(12) C. setosissima.

Langille, H. D.

5195a, 6728,

(15) C. virginensis;

117 (6)
flora.

fla

Wm

6692, 6713 (17) C. modesta; 6732, — I

(18) C. humilis; 6726 (19) C. caespi-

tosa; 972 (20) C. thyrsiflora; 5607a Lechiner, H. J.

1030 (27) C. celosioides.

(22a) C. sericea var. typica;
C. sericea var. perennis;
rugulosa; 6673 (26) C. spiculifera;
(28) C. Sheldonii; — (29) C. Macom

(22b)

3 (7) C. leucophaea.

(24) C. Leckenby, A. B.

(30) C. sobolifera; 6698, 6699, 6700,

6705, 6706,

(32a) C. nana var.

commixta; 1692, 6709, 6719,
C. nana var. Shantzii; —

- (32b)
(32c) C.

(7) C. leucophaea.
Leiberg, J. B.

5545 (la) C. Jamesii var. multicaulis;
5748 (12) C. setosissma; 5294 (18)
C. humilis; 125 (27) C. celosioides;
2049, 2223 (34) C. propria.

nana var. ovina; — (32d) C. nana var. Lemmon, J. G.

typica; 6474 (34) C. propria;

(35)

165 (18) C. humilis.

C. breviflora; — (36) C. fulvocanescens; Lemmon, J. G. & wife.

5297p, 531 2ac,

(37) C. echinoides;

(38) C. Jonesiana;

(39) C.

Wetherillii;

— (la) C. Jamesii var. multicaulis;
(12) C. setosissima.

(40) C. longiflora;

W

(42) C. Osterhoutii;

(43) C. para-

doxa; 54-45p,

iflo

52041, 6677, 6678, 6715, 6716, 6717,

(45) C. mensana; Little, E. E. & E. M. Stanton.

6720, 6721, 6726, 2729,
flavoculata.

179 (22a) C. sericea var. typica.

(46) C. Lunell, J.

(31) C. Bradburiana.

[Vol. 14

342

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Lyall, David.

4771, 7067, 10693,

(6) C. flava;

— (7) C. leucophaea.
Macbride, J. F.

93, 875 (26) C. sjnculifera; 2 (32b) C.

nana var. Shantzii.
Macbride, J. F. & E. B. Payson.

950 (lc) C. Jamesii var. cinerea; 705

(6) C. flava; 3848 (8) (\ salmonensis;

3771 (10) C. nubigena; 3039,

(26) C. sjricidifera.

MacDougal, D. T.

49 (la) C. Jamesii var. muHicaulis;

£04 (lc) C. Jamesii var. cinerea; 79

231, 1267, 1937, 7338 (13) C. virgata;
3072, 3120, 4^71, 4749, 4772, 7078,
7255, 10696 (19) C. caespitosa; 3035,
4715 (22b) C. serieea var. perennis;
418, 1362, 1990, 7306, 8263 (20) C.
thyrsijiora; 255, 888, 1966, 4788, 483
6961, 7248, 8319, 9045, 9355, 9411,
9045, 10700, 10741, — (31) C. Brad-

1

j

buriana; 889, 2876, 8309, 9867, —

(33) C. carta; 1511, 4513, 4">7°2, 4616,
4665, 4749a, 4750, 4815, 7090, 9028,
(46) C. flavoculata.

(le) C. Jamesii var. typica; 165 (12) Nelson, A. & J. F. Macbride.

C. setosissima; 169 (28) C. Shcldonii;

60 (31) C. Bradburiana.

Macoun, John.

6801, 11819, 11820, 12800, 17087,

17089, 76741, 785DO, S5007,

(29)

C. Macounii; 5802 (31) C. Bradburi-

ana.

(la) C.

M earns, E. A.

Ill, 359, 360, 2091,

Jamesn var. multicaidis.
Merrill, E. D. & E. N. Wilcox.

486, 497, 726 (6) C. flava; 617 (19) C.

caespitosa; 457 (46) C. flavoculata.

Metcalfe, O. B.

70, 481, 868, 1061 (la) C. Jamesii var.
multicaidis; 1576 (4) C. oblata.
Moodie, M. E.

— (29) C. Macounii.
Moore, E. J.

— (31) C. Bradburiana.

Moyer, L. R.

468,

(31) C. Bradburiana.

Mulford, A. I.

— (32b) C. nana var. Shantzii.
Munz, P. A.

5721 (If) C. Jamesii var. abortiva.
Munz, P. A., I. M. Johnston & R. D.
Harwood.
4209 (16) C. tumulosa.

Nealley, G. C.

(la) C. Jamesii var. rnidticaulis ;
167 (4) C. oblata.
Nelson, A.

1960, 1980,

(18) C. humilis; 1799

(26) C. spiculifcra.
Nelson, A. & E. Nelson.

5429 (28) C. Shddonii.

Nelson, E. W.

6350 (lb) C. Jamesii var. laxa.

Nelson, E.

4906 (le) C. Jamesii var. typica; 4397
(6) C. flam; 4338, 4497 (19) C. caes-
pitosa; 4337 (46) (\ flavoculata.

Nicollet, I. N.

374 (31) C. Bradburiana.

Nuttall, T.

(6) C. flora;

( 19) C. caespitosa;

(20) C. thyrsijiora; — (33) C. cana;

(46) C. flavoculata.
Osterhout, G. E.

628, 9087 (lc) (\ J nun six var. cinerea;
627, 629, 3992, 4979, 5754, — (le) C.
Jamesii var. typica; 6389, 6406 (6) C.
flam; 6195 (9) C. rtricta; 2463, — (13)

C. virgata; 6248 (19) (\ catspitosa; — ,
4979, 5934 (20) C. thyrsijiora; 0996
(21) C. elata; 2122, 3476, 4*940, 5960,

6094,

(22a) C. serieea var. ty\nca;

5927 (31) C. Bro'llniriana; 5902 (33) C.
cana; 5980, 6010, 6200 (32d) C. nana
var. typica; 6416 (35) C. breviflora; 6003

(36) C. fvlvocanescens; 4463, 4497,
5975 (40) C. longiflora; 6138 (42) C.
Osterhoutii; 2610 (44) C. Bakeri; 2750,

4696, 5956, 6006, 6194, 6197, 6252 (46)
C. flavoculata.

477, 2567, 2882, 8805, 9379, — (le) C. Over, W. H.

Jamesii var. typica; 22, 3074, 3098,

5188, 6185,

(31) C. Bradburiana.

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA

343

Overholts, L. 0.

10, 155 (13) C. virgata.

Palmer, Ed.

591 (la) C. Jamesii var. multicaulis;

Pearson, G. A.

210 (la) C. Jamesii var. multicaulis.

Peck, M. E.

10020 (27) C. celosioides .

375 (lc) C Jamesii var. cinerea; 895 Pierson, F. E.

(3) C. Palmeri; — (6) C. jto»a; 357, 59i

SiOS (If) C. Jamesii var. abortiva.

(12) C. setosissima; 189 (26) C. s/ncw- Piper, C. V.

lifera; 598 (32b) C. nana var. Shantzii.

Palmer, Wm.

(29) C. Macounii.

Parish, S. B.

2987,

(7) C. leucophaea; 2294 (28)

C. Sheldonii.
Porter, T. C.

(22b) C. sericea var. perennis.

1480 9 3238, 3694 (If) <?. Jamesii var. p ra tt, A. D.

abortiva; 1316, 1319, 3240, 4887, 19167

(31) C Bradburiana.

(5) C. conjertifiora; 102 J^S (16) C. p r i ng le, C. G.

tumulosa.
Parish, S. B. & W. F. Parish.
1316 (5) C. conjertifiora.

Parry, C. C.

i6£ (5) C. conjertifiora; 173 (15) C.
virginensis; 49 (46) C. flavoculata.

Patterson, H. M.

HO (13) C. virgata;— (22a) C. sericea

var. typica.

Payson, E. B.

(la)

1019 (lc) C. Jamesii var. cinerea; R ama i ey p.

776 (lb) C. Jamesii var. loza; —
C. Jamesii var. multicaulis.

Purpus, C. A.

6065, 7067, 5045a, 5^55 (lc) C. Jamesii
var. cinerea; 5323, 5802 (5) C. con-
fertiflora; 8048 (12) C. setosissima;
(15) C. virginensis; 6070 (32 e) C. nana
var. onm; — (32d) C. nana var.
typica; — (41) C. tenuis; 5869 (46) C.
flavoculata.

1027 (le) C. Jamesii var. typica;
405 (6) C. M' !0i4 (20) C. %m-
,/tora; 27i (36) C. fulvocanescens ; R ^ n " t

A. 107, 1075 (13) C. virgata; 1448 (20)
C. thyrsiflora.

35, 659 (40) C. longiflora; 458 (43) C.
paradoxa; 1079, 1135 (44) C. Bakeri;
246, 337, 670, 997, 1046 (46) C. flavo-
culata.
Payson, E. B. & G. M. Armstrong.
3223 (19) C. caespitosa; 3206, 322,
(22b) C. sericea var. perennis.

Payson, E. B. & E. Bethel.

1608 (le) C. Jamesii var. typica.
Payson, E. B. & L. B.

2533, 2557, 4307 (6) C. flava; 1880 (8)

2120 (3) C. Palmeri.

Rothrock, J. T.

/y

ifi

Rusby, H. H.

/

Jamesii

caulis; 748 (12) C. setosissima.

Rydberg, P. A.

254, 1514 (le) C. Jamesii var. typica;
255 (20) C. thyrsiflora; 893 (31) C.
Bradburiana; 256 (33) C. cana.

C. salmonensis; IfiJfr 4253, 4291 (13) Rydberg, P. A. & E. A. Bessey.

C. virgata; 2578 (19) C. caespitosa;

4882, 4883 (31) C. Bradburiana.

4246, 4685 (20) C. thyrsiflora; 2528 Rydberg, P. A. & E. C. Carlton.

(22a) C. sericea var. typica; 2580 (22b)

70^1 (lc) C. Jamesii var. cinerea.

C. sericea var. perennis; 1768 (26) C. Rydberg, P. A. & A. O. Garrett.

spiculifera; 1746 (29) C. Macounii;
4247 (31) C. Bradburiana; 3876 (36)

9569 (2) C. pustulosa; 8431 (6) C.
flava; 9130 (44) C. Bakeri.

C. fulvocanescens; 4223 (43) C. para- Rydberg, P. A. & F. K. Vreeland.

doxa; 3834 (44) C. Bakeri; 1689, 2583,
3865, 3874, 3875 (46) C. flavoculata.

5702 (lc) C. Jamesii var. cinerea; 5704
(20) C. thyrsiflora.

344

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

Sandberg, J. H.

— (28) C. Shcldonii.
Sandberg, J. H. & J. B. Lei berg

878 (7) C. leucophaea; 164 — (26) C
spicuiifera; 440 (27) C. celosioides.

Saunders, A. A.

— (29) C. Macounii.

Saunders, I). A.

4875 (26) C. sfnculifcra.
Scheuber, K. W.

— (31) C. Bradburiuitii.

Schmoll, H. M.

1281 < 1846 (44) C. Bakeri.
Schoper, M.

28 (32b) C. nana var. Shanteii.
Scribner, (J. L.

174 C31) (\ Bradburiana.
Shantz, II. L.

571 (le) C. Jamesii var. typica.
Sharp, S. S.

132 (31) C. Bradburiana.

Shear, C. L.

4381 (22b) C. scricea var. perennis.

Sheldon, C. S.

527 (lc) C. Jamesii var. cincrea; 216
(13) r. mrja(a; 526 (20) C. thyrsiflora.

Sheldon, E. P.

8815 (28) (7. Shcldonii.

Shockley, W. H.

(5) C. confertiflara; 244 (46) C.
ftavocxdata.

Slaughter, I. W.

7 (32b) C. nana var. Shanteii.
Smith, C. P.

7680 (lc) C. Jamesii var. cinerea;

8912 (le) C. Jamesii var. typica;

8916 (20) (7. thyrsiflora; 1578, 1605

(32b) C. //ana var. Shanteii; 8986 (44)

C. Bakeri.

Sonne, C. F.

— (18) C. hum His.
Soth, Mrs. M. E.

Starz, E.

(29) C. Macounii.
Stevens, G. W.

385 , S024, 3035 (la) C. Jamesii var.
multicaulis.
Stokes, S. G.

— (6) C. flava; — (17) C. humilis; —

P-!* 32b) C. nana var. Shanteii.

Btandley, P. C.

4899, 7126, 7880, 7067 (la) C. Jamesii Visher, S. S.

(32a) C. nana var. commixta; — (32b)
C. nana var. Shantzii; — (39) C
Wetherillii; — (46) C. 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 (le) C. Jamesii var. typica.
Thurber, Geo.

278 f 298 (la) C. Jamesii var. multi-

caulis; 147 (4) C. oblata.
Tidestrom, Ivar.

2031 (6) C. ./fara; i£93, ,9£i0 (46) C.
flavoculata.
Tourney, J. W.

224 (12) C. setosissima.
Tracy, S. M.

7835 (la) C. Jamesii var. multicaulis.
Tracy, S. M. & F. S. Earle.

424 (3) C. Palmeri.
True, G. H,

S65 (18) C. humilis.
Turesson, G. W.

— (28) C. Sheldonii.
Tweedy, Frank.

576 (la) C. Jamesii var. multicaulis;
5219 (le) C. Jamesii var. typica; 4262
(6) C. flava; 5222, 5228, 5676 (13) C.
t^r^a/a; ^jgffl} (19) C. caespitosa; 3570

29) C. Macounii; 816, 3569, 4261,
(31) C. Bradburiana; 4259 (46) C.

flavoculata.

Umbach, L. M.

J# (29) C. Maeranu.

Vasey, G. R.

(4) C. oblata; — (20) C. thyrsiflora.

var. multicdulis; — (4) (7. oHota; 6'03£,

6066 (20) C. thyrsiflora; 15107, 17428, von Schrenk, H.

2S5, 481, 587 (31) C. Bradburiana.

17594 (28) C. Shcldonii.

(31) C. Bradburiana.

1927]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA

345

Voth, H. R.

15, 103 (6) C. flaw.
Waldron, C. B.

— (31) C. Bradburiana.
Walker, E. P.

85, 156 (36) C. fulvocanescens ; 91

(43) C. paradoxa.
Ward, L. F.

893, 557 (Id) C. Jamesii var. disticha;

Wilcox, E. V.

79 (29) C. Macounii.
Wilkes, Charles.

(61) 487 (28) C. Sheldonii.
Williams, R. S.

109 (28) C. Sheldonii.
Williams, T. A.

— (la) C. Jamesii var. typica;

108,

(5) C. confertiflora;

(6) C.

(20)

C. thyrsiflora; — (31) C. Bradburiana;
— (33) C. carta.

flava; 646 (12) C. setosissima; 49 (17) Willits, Vie.

C. modesta; 87 (46) C. flavoculata.
Waring, M. G.

15 (36) C. fulvocanescens.
Watson, S.

92 (29) C. Macounii; 236 (31) C.
Bradburiana.
Wolf, John, & Rothrock, J. T.
700 (20) C. thyrsiflora.

853 (18) C. humilis; 852 (22b) C. Wooton, E. O.

sericea var. perennis; 287 (28) C.

Sheldonii; 852 (46) C. flavoculata.
Webber, H. J.

(31) C. Bradburiana.
White, O. E.

— (31) C. Bradburiana.
Whited, Kirk.

401, 595, 2818, — (la) C. Jamesii var.

multicaulis;

(4) C. oblata;

(12)

C. setosissima; — (29) C. Macounii.
Wright, C.

489 (3) C. Palmeri; 1566 (4) C. oblata.
Zuck, M.

— (36) C. fulvocanescens.

1099,

(27) C. celosioides.

Index to Species and Varieties

Synonyms are printed in italics; species or genera maintained in this paper
are printed in Roman type; new species, varieties, or combinations are indicated
in bold face type.

Page
Cryptantha

affinis 307

aperta 295

Bakeri 331

Bradburiana 307

breviflora 318

caespitosa 281

cana 316

celosioides 299

Clemensae 267

confertiflora 256

echinoides 321

elata 285

flava 259

flavoculata 334

fulvocanescens 319

glomerata 307

Page

holoptera 270

humilis 278

insolita 273

interrupta 296

Jamesii 242

var. abortiva 250

var. cinerea 246

var. disticha 248

var. laxa 246

var. multicaulis 244

var. typica 248

Jonesiana 323

leucophaea 262

longiflora 326

Macounii 303

mensana 333

modesta 277

346

[Vol. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Page

nana 312

var, commixta 312

var. ovina 314

var, Shantzii 313

var. typica 315

nubigena 265

oblata 254

Osterhoutii 329

Palmeri 253

paradoxa 330

propria 317

pterocarya 270

pustulosa 252

rugulosa 295

salmonensis 203

sericea 280

var. perennis 288

var. typica 2S7

setosissima 268

Sheldonii 301

sobolifera 305

spiculif era 298

stricta 264

euffruticosa 242

tenuis 327

thyrsiflora 283

tumulosa 276

utahensis 270

virgata 270

virginensis 274

Wetherillii 324

Cynoglossum

glomeratum 307

Eritrichium

ftdvocanescena 319

glorm ratum 303, 307

var. f ulvocanescens 278

Var. hispidissimum 283

var. humUe 278, 287

var. virgatum 270

Jamesii 248

leucophaeum 262

mitlticaide 244

setos-issimum 268

virgatum 270

Krynitzkia

depressa 277

echinoides 321

Page
f ulvocanescens 319

var. idahoensis 317

glomerata 307

var. acuta 324

var. virginensis

274

Jamesii 248

leucophaea

262

var. alata 256

mensana 333

midticaidis

var. abortiva 250

var. setosa 244

oblata 254

l'almcri 253

pust alata 307

sericea 287, 316

setosissima 268

virgata 270

MyosoHs
glomerata 307

leucophaea 262

suffruticosa 242, 248

Oreocarya

abortiva 250

307

affinis

affi/nis perennis 288

256

295

alata

apt rta

argentea 287

Bah ri 331

br< iriflora 318

caespitosa 281

cana 316

celosioides 299, 301

cilio-hirsuta 298

cinerea 246

commixta 312

co rtfe.rti flora 256

cristata 334

depressa 277

disticha 248

dolosa 313

dura

283

Eastwoodae 334

echinoides 278. 321

elata

285

eulophus 331

flava

259

192 7]

PAYSON — SECTION OREOCARYA OF CRYPTANTHA

347

Page

flavoculata 334

var. spatulata 334

fulvocanescens 319

glomerata 301, 307

gypsophila 330

hispida 278

hispidissima 254, 283

holoptera 270

horridula 326

humilis 278

insolita 273

interrupta 296

Jonesiana 323

Lemmoni 246

leucophaea 262

var. confertiflora 256

longiflora 326

lutea 256

lutescens 259

Macbridii 278

Macounii 303

mensana 333

monosperma 283

multicaulis 244

var. cinerea 246

var. laxa 246

nana 312, 315

nitida 319

nubigena 265

oblata 255

Osterhoutii 329

Page

Pahneri 253

paradoxa 330

Paysonii 255

perennis 288, 307

procera 287

propria 317

pustulosa 252

rugulosa 295

salmonensis 263

sericea 287, 301

setosissima 268

Shantzii 313

Sheldonii 301

Shockleyi 334

spicata 270

spiculifera 298

stricta 264

suffruticosa 248

var. abortiua 250

var. cinerea 246

var. multicaulis 244

tenuis 327

thyrsiflora 283

tumulosa 276

urticacea 283

virgata 270

forma spicata 270

virginensis 274

WetheriMi 324

Rochelia

glomerata 307

[Vol. 14, 1927]

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 pi. 25 (complete fruit), which is times 7 l A.

PLATE 25

Figs. 1-4. C. Jamesii var. multicaulis. Drawn from Wooton Jfil. The nutlets
of all the varieties of C. Jamesii, C. pustulosa, and C. Palmeri are very similar, and
these figures will serve to illustrate them.

Figs. 5-7. C. oblata. Drawn from Jones 3759.

Figs. 8-10. C. confertiflora. Drawn from Goodding 814*

Figs. 11-13. C. flava. Drawn from Goodding 164-

Figs. 14-16. C. leucophaea. Drawn from Sandberg & Leiberg 873.

Figs. 17-19. C. salmonensis. Drawn from Payson & Payson 1880,

Ann. Mo. Bot. Gard., Vol. 14, 1927

Plate 25

y

i

2

3

\
* \

*\

/

7

*1

8

/

/

ii

12

15

■J

4

9

13

5

6

14

V

i\

*-A>. y

16

17

18

19

PAYSON— SECTION OREOCARYA OF CRYPTANTHA

COCKAYNE, BOSTON

IVol. 14, 1927]

350 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 26

Figs. 20-22. C. strict a. Drawn from Osterhout 6195.

Figs. 23-25. C. nubigena. Drawn from specimen collected by Cusick, Pine
Creek, Oregon.

Figs. 26-28. C. Clemensae. Drawn from type, Clemens, Glenn's Pass, California.

Figs. 29 31. C. setosissima. Drawn from specimen collected by M. E. Jones,
Flagstaff, Arizona, August 4, 1884.

Figs. 32-34. C. virgata. Drawn from specimens collected near Laramie, Wyo-
ming, by the author in 1925.

Figs. 35-37. C insolita. Draw T n from specimen collected by K. Brandegee,
Las Vegas, Nevada.

Ann. Mo. Bot. Gard., Vol. 14, 1927

Plate 26

25

21

26

22

27

V

\

29

\

1 <

31

32

34

35

36

PAYSON— SECTION OREOCARYA OF CRYPTANTHA

COCKAYNE, BOSTON

352 ANNALS

[Vol. 14, 1927

Explanation of Plate

plate 27

Figs. 38-40. C. virginensis. Drawn from M. E. Jones 5195 a.

Figs. 41-43. C. tumulosa. Drawn from type, T. S. Brandegee, Providence
Mountains, California.

Figs. 44-46. 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. C. caespitosa. Drawn from Payson & Pay son 2578.
Figs. 53-55. C. thyrsiflora. Drawn from Osterhout 4979.

Ann. Mo. Bot. Gard., Vol. 14, 1927

Plate 27

£»

42

38

39

43

40

44

49

41

46

50

52

53

54

55

PAYSON— SECTION OREOCARYA OF CRYPTANTHA

COCKAYNE, BOSTON

[Vol. 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. C. sericea var. typica. Drawn from Pay son & Pay son 2528.

Figs. 62-64. C. aperta. Drawn from type, Eastwood, Grand Junction, Colorado.

Figs. 65-67. C. rugulosa. Drawn from type, M. E. Jones, Fish Springs, Utah.

Figs. 68-70. C. interrupta. Drawn from Heller 9185.

Figs. 71-73. C. spiculifera. Drawn from type, Sandberg & Leiberg 164.

Figs. 74-76. C. celosioides. Drawn from Suksdorf 888.

Figs. 77-79. C. Sheldonii. Drawn from type, Sheldon 8315.

Ann. Mo. Bot. Gard., Vol. 14, 1927

Plate 28

58

62

67

68

59

73

77

78

76

79

PAYSON— SECTION OREOCARYA OF CRYPTANTHA

COCKAYNK, BOSTON

[Vol. 14, 1927]

35G ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 29

Figs. 80-82. C. Macounii. Drawn from specimen collected by Macoun, at
Moose Jaw, Saskatchewan.

Figs. 83-85. C. sobolifera. Drawn from type, M. E. Jones, Upper Marias
Pass, Montana.

Figs. 86-88. C. Bradburiana.

Figs. 89-91. C. nana var. Shantzii. Drawn from type of Oreocarya dolosa, C. P.
Smith 1605. The nutlets of C. breviflora, C. fulvocanescens, and C. echinoides are
similar to those of C. nana var. Shantzii. The differences are noted in the descrip-
tions.

Figs. 92-94. C. cana. Drawn from type, A. Nelson 8309.

Figs. 95-97. C. projrria. Drawn from Leiberg 2223.

Figs. 98-100. C. Jonesiana. Drawn from type, Jones. San Rafael Swell. Utah.

Ann. Mo. Bot. Gard., Vol. 14, 1927

Plate 29

80

89

i

81

93

97

86

'. IV

» Vi

>y#%

*0 %

*i

90

94

82

83

87

88

92

96

100

I

t

PAYSON -SECTION OREOCARYA OF CRYPTANTHA

COCKAYNE, BOSTON

358

[Vol. 14, 1927]

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 30

Figs. 101-103. C. Wethcrillii. Drawn from type, Eastwood, Court House Wash,

Utah.

Figs. 104-106. C. longijlora. Drawn from Pay son 659.

Figs. 107-109. C. tenuis. Drawn from specimen collected by Pur pus.

Figs. 110-112. C. Ostcrhoutii. Drawn from type, Osterhout 6188.

Figs. 113-115. C. paradoxa. Drawn from Pay son & Payson 4223.

Figs. 110-118. C. Bakeri. Drawn from Payson & Payson 3834.

Figs. 119-121. C. mensana. Drawn from type, Jones 5445 p.

Figs. 122-124. C. flavoculata. Drawn from Payson & Payson 2583.

Ann. Mo. Bot. Gard., Vol. 14, 1927

Plate 30

103

118

109

114

121

122

123

PAYSON— SECTION OREOCARYA OF CRYPTANTHA

COCKAYNE, BOSTON

Annals

of the

Missouri Botanical Garden

Vol. 14 NOVEMBER, 1927 No. 4

COTTON WILT: A PATHOLOGICAL AND
PHYSIOLOGICAL INVESTIGATION 1

DAVID C. NEAL

Plant Pathologist, Mississippi Agricultural Experiment Station
iufus J. Lackland Research Fellow in the Henry Shaw School of Botany of

Washingt

■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. The 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 a year was spent in a laboratory and greenhouse investi-

carried

Washingt

mitted as a thesis in partial fulfillment of the requirements for the degree of doctor

Washingt

Ann. Mo. Bot. Gard., Vol. 14, 1927

(359)

3G0 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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 fundus 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 ('99) 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. The field evidence, how-

19271

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 ('10) 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 : ' ' No 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 ('00) 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. He 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, succumb more readily than those which have grown
on poorer soil." Orton also corroborated the work of Smith ('99)
and Atkinson ('92) 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. He conducted experiments
with a number of fungicides, spraying these on the plants and

IVol. 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. The 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 ('07), 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 ('11) 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, 1 used at the rate of from
300 to 800 pounds per acre, were of no benefit in controlling it.

eplicated, however, and
ivas not indicated.

Lewis C15) originated

from 1905 to 1915, some of which are still widely planted

era

section of the cotton belt. He used both the
method of selection and of hybridization of desirable parent

» 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-anfi, 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 19
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-

6.25 per cent potash. It was applied in the spring of

and

1921 at the rate of 1000 pdunds per acre before planting. His
results are quoted: "The 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, p. 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

[Vol. 14

3C4 ANNALS OF THE MISSOURI BOTANICAL GARDEN

received enabled the plants this year to very largely escape wilt

infection."

Description of Cotton Wilt

A noticeable yellowing of the leaves of the cotton plant accom-
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

infested

the main stem of

plants infected with the disease remains short, while some of the
lower branches grow normally (pi. 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 (pi. 31, fig. 2). The
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 taxonomic study by the
writer. It seems desirable, therefore, to accept the criteria of
taxonomic differentiation of species as established through the
work of Wollenweber and associates ('25). The 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. Micro-conidia are present on the aerial
mycelium and are dominantly non-septate, ovoid to fusoid,
4-6 x 3-4.5 (x. Macro-conidia are usually none to 3-septate,
attenuate at distal end, pedicellate, fusoid to sickle-shaped,
20-25 x 4-6 [i. Chlamydospores are produced in cultures 3
weeks old or less and may be either terminal or intercalary (pi. 32,

1927]

NEAL — COTTON WILT 365

figs. 3-4). The 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. On 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 a 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. On July 15, 1926, the fungus was isolated
from a wilted plant of Trice cotton in the breeding plot at the
Experiment Station, A. & M. College. It was also isolated from
a wilted plant of Delfos 6102 cotton collected at Dunleith, Missis-
sippi, on August 16, 1926, and from Trice 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. The 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.

[Vol. 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:

Oilman ('16) 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° C. 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 ('20) has worked on the temperature relations of
tomato wilt, and he and other investigators state that the tomato
wilt fungus, Fusarium Lycopersici Sacc, 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 ('23) 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 367

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 a 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 ('20), 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

fun

orm Petri dishes

measured amount (10 cc.) 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. The 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

368

ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

loopfuls of the spore suspension were placed at random

surface

As soon as the agar solidified

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. The results are given in
table i.

TABLE I

GROWTH OP FUSARIUM VASINFEOTUM AT VARIOUS TEMI'KllATURES

Temperature
degrees C.

Diameter of colony in cm. at indicated intervals

2 days

Plate No.

6
10
15
L'O
24

28

30
35

40

6 days

10 days

Plate No

Plate No.

15 days

Plate No.

1

1.2
2.5

1.5
2.8
5.4 5.2

9.0*
9.0*
9.0*
9.0*
3.5

9.0*
9.0*9.0*

3

1.2

2.9
5.4

9.0*

0.0*
0.0*

3.0

9.0*
!> . 0*

3.5

1

1.5

3.8

8.2
9.0*

2

9.0*
9.0*

1.6
3.6
8.0
9.0*

3

1.6

3.4
8.1

9.0*
9.0*

t

* Growth over plate,
t Contaminated.

It will be noted from the above results that Fusarium vasin-
fectum grows very slowly at the lower temperatures, and that the

C. The optimum temperature,

m

therefore, of the parasites of the very similar Fusarium diseases of
cotton, tomato, flax, and cabbage is practically the same in pure

gh the host plants have quite different temperature

supplemented by the work of Edgerton
others. The maximum temperature of

relations.

Tisdale C23). and

Fusarium vasinfectum is apparently around 38° C.

The effect of temperature on the growth of Fusarium
fectu?n was also studied by means of liquid cultures. T

dry

of the fungous mat was ascertained at various tempera

i

1927]

NEAL — COTTON "WILT

369

tures ranging from 6 to 40° C. at intervals of 5, 10, and 15 days.
Duggar's 1 solution was used for growing the fungus and the final
concentration of chemicals used in this solution was as follows:
M/4 dextrose, M/5 KN0 3 , M/20 KH 2 P0 4 , M/100 MgS0 4 , and
a trace of FeP0 4 . For obtaining these dilutions the following
stock solutions were prepared: M/2 dextrose, M/l KN0 3 ,
M/4 KH 2 P0 4 , M/10 MgS0 4 , and M/1000 FeP0 4 . For 50 cc. of
medium the following amounts of these solutions were used:
25 cc. dextrose, 10 cc. KN0 3 , 10 cc. KH 2 P0 4 , 5 cc. MgS0 4 , and
6 drops FeP0 4 . Pyrex flasks were used for the cultures, and
these were run in triplicate at each temperature. A 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. The
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 on a filter paper by n-eans of a Buchner filter, dried to
constant weight in a Fries oven, 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 i

milligrams

degrees C.

5 days

10 days

15 days

6

6.20

54.9

81.4

10

28.0

84.5

97.1

15

35.8

98.7

145.2

20

38.2

149.4

251.5

25

49.5

112.3

455.2

28

226.0

338.6

610.0

30

382.3

493.7

725.2

35

286.5

294.2

422.9

38

172.0

247.3

*

Contaminated

A nutrient solution for fungi recommended by Dr. B. M. Duggar

t

370

[Vol. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

noted again that the optimum temperature for Fusarium vasin-
fectum is around 28 and 30° C. 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.

Fig. 1 . Showing the effect of temperature on the growth of Fusarium vasinfedum.

RELATION TO HYDROGEN-ION CONCENTRATION

The effect of the hydrogen- and the hydroxyl-ion
>on the germination of the spores and growth of
& been demonstrated by numerous investigatoi
action of the substratum is a very important fact
g the growth of many disease-producing organis

That the

known fact

subject

and has been reviewed to advantage by Webb

Lehman f 23), and

Webb C21) finds

Fusarium sp., germination is equally as good, if not better, with
an alkaline reaction. However, with the majority of fungi he

1927]

NEAL — COTTON WILT 371

i

employed the maximum for germination was between P H 3.0 and

4.0.

Hopkins ('21), using a synthetic medium in which the hydrogen-
ion concentration was adjusted by means of KH2PO4, KjHP0 4 ,
H3PO4, 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 P H 2.5 toamaximum
of P H 4.0-4.5. It then decreased to a minimum at P H 5.0-5.5 and
rose again to a second maximum, but the highest point was not

determined.

Maclnnes ('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. The organism grew in nutrient media rang-
ing from P H 5.0 to P H 11.7. No 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 P H 5.2-9.2.

The 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
nourish 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 ('22) 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 ('20, '21), Lint ('14,
'15, '16), 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 P H value of 7

bly

with a P
crop.

H

bby, while an acid
produce an almost c

o 70 (VOU 14

6(A 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 inorganic salts and dextrose in the following volume
molecular concentration: M/4 dextrose, M/5 KN0 3 , M/20
KH 2 P0 4 , and M/100 MgSO*. Six drops of M/1000 FeP0 4 were
used for each 50 cc. of solution, and M/5 Ca(N0 3 ) 2 was substi-
tuted for the M/5 KN0 3 . The inorganic salts were Merck's
"blue label" reagent grade, and the dextrose was the grade desig-
nated as "difco standardized." The nutrients were prepared in
500-cc. Pyrex flasks in quantities of 475 cc. for each series, and
this quantity of solution was adjusted to the desired P H value
by the addition of previously determined amounts of 0.1647 N
H3PO4 or 0.1613 N KOH. The range of initial P H values was
from 2.5 to 9.0 After P H adjustments were made, the 400 cc.
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 cc. of solution of each initial P H
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 H and to serve as controls for final P H reading at the end of
designated growth periods. It was found that autoclaving did
not change the P H 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 a temperature
of 29° C. The cultures were also w
exclude the light from the bulbs. H

apped with carbon pap

examined at the end of 8, 12, 16, and

each

dry weight of mat, hydrogen-ion concentration, and titrable
acidity were determined at each examination. The P„ values
were determined colorimetrically by the method of Clark and
Lubs C17).

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 a Fries oven at
100° C, cooled in a calcium chloride desiccator for a uniform
period, and weighed. The filtrate from each mat was diluted
to 100 cc. 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. The 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) was employed. The data obtained are shown
in table in.

TABLE III

GROWTH OF FUSARIUM VASINFECTUM IN NUTRIENT SOLUTION*

Growth

Dry wt.

Cc. N/50 KOH

Wt. of sugar per

Series

Initial

period

mat

Final T H

to neutralize

cc. of solution

No.

Ph

days

mgms.

10 cc. filtrate

mgms.

8

49.0

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

8

397.0

6.2

11.9

. __.

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

5.7

14.2

16

417.1

5.8

13.6

28

761.0

6.9
4.5

1.2

_____

8

116.0

16.8

5

4.0

12

345.2

5.8

13.4

16

419.0

5.8

14.2

28

758.3

7.3

.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

362.3

6.6

9.8

4.8

16

413.0

6.8

7.1

2.88

28

732.6

7.2

2.3

374

[Vol. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

TABLE III (Continued)

Series
No.

Initial

Ph

Growth

period
days

Dry wt.

mat
mgms.

Final P H

Cc. N/50 KOH

to neutralize
10 cc. filtrate

Wt. of sugar per
cc. of solution
mgms.

8

6.0

8

12

16

28

195.0
380.0
441.6

577.8

5.6
6.4
6.4
7.3

14.7
5.9
6.1
3.7

9

6.5

8
12

16

28

199.0
353 .
434.1
452 .

189
318
447
512

6.2
6.6
7.2
7.4

10.6
9.9
3.9
1.3

10

7.0

8
12
16

28

6.4
6.4
6.2

7.2

4.3

4.35
8.6

1.75

11

7.5

8
12

16

28

144
358

455
502

6.4
6.2

6.3
7.4

3.4

9.6

9.4
0.30

12

8.5

8
12
16

28

109
299
421
529

6.9

7.0
7.6
8.4

3.8

.4

.551
1.3J

30.2
22 . 2

9.8

13

8.8

8
12

16

28

87
257
405

487

6.7
6.9

t-

8.5

5.2
2.7

1.21

14

9.0

8
12

16f

28 1

96
251

427

8.4
7.9

7.8

.95|
.70*

.851

* The values given are the averages of 2 cultures,
t Contaminated,
t .2 N HC1.

The relations of time to growth in the series are shown

fig

the end of the 28-day period, the best growth

indicated by the dry weight of mat, occurred in series 3

epresented by

H

and

maximum growth occurred

the cultures started

>. The

1 it O.O.

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 P H exponents. Judging from the behavior of the
cultures in series 1 and 2, the fungus tolerates a culture solution

1927]

NEAL — COTTON WILT

375

Fig. 2. Showing the relation of time to the growth of Fusarium vasinfedum

strongly acid. However, the maximum growth attained at the
close of the experiment was notably less with the more acid
cultures. All series initially acid became less so, while all
initially alkaline became slightly acid excepting a few started on
the extreme alkaline side of P H 7, which, although becoming acid
at first, reversed the reaction later and were distinctly alkaline at
the close of the growth period.

376 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

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 ('99) suggested
this theory in the case of cotton, cowpea, and watermelon plants,
which were infected with closely related fungi of the genus
Fusarium. He 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 ('09),
Stevens ('21), and Orton ('00, '08, '12). 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 a 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

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
obstruction 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 fact that cotton wilt is more severe in a 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 periods of
abnormally dry weather, but may frequently develop suddenly
and in destructive form as soon as such periods are broken by
ample rainfall. Beal ('26) has also noted a greater prevalence
of the disease during periods of dry weather. Therefore, since
a 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 microscopic 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 ('19) found that cultures of Fusarium cubense, grown in
Richards' solution, when filtered free from the fungus, produced
a marked wilting in buckwheat and bean plants, as well as banana
leaves. He obtained the same results using Uschinsky's solution
as a 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 ('19) reports the results of experiments in which leaves

378 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

of different plants were placed in filtrates from Fusarium oxy-
sporum, F. discolor sulphureum, Fusarium from pea, and other
fungi. Wilting occurred within a few hours with some extracts,
a fact which he interprets as not to be explained by changes in
acidity. Excised leaves placed in uninoculated media and in
water did not wilt until considerable time had elapsed. The
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 oxy-
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 ('21) 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 P H 5.0 to P H 7.4 on the
fortieth day. But w r hen the control (uninoculated) solution was
adjusted to P H 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.

In 1921, Ajrekar and Bal ('21), 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
('IS). 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 ('23), Fahmy ('23),
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. The 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 ('26) has shown that nitrates represent-
ing cultures growing on Richards' solution are quite toxic to
cotton plants, while nitrates 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. The 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

[Vol. 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 xyleni in these cases is due to the formation of
toxic substances by the fungus in the soil. This author concludes
that tho 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 mechanical
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-

lenomena

FliNGUS

In order to obtain some information on the probable toxic
properties of substances given off by Fusarium vasinfectum, the
following experiment was performed. The fungus was grown at
room temperature for 18 days in 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 a
mortar, and the mixture extracted with 75 cc. of sterile distilled
water. Another was prepared in the same way and the extract

A final series of filtrates was also prepared

boiled for 10 minutes. A final series of filtrates was also pr
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° C,
and examined at intervals of 3, 6}^, and 36 hours. The data
obtained are presented in table iv.

TABLE IV

EFFECT OF FILTRATES AND MYCELIAL EXTRACTS OF 18-DAY CULTURES

OF FUSARIUM VASINFECTUM ON COTTON SEEDLINGS

Series

Treatment

Wilt symptoms present after

No.

3hr. | 6Hbr. 36 hr.

Filtrate

Extract of mycelial mat

Filtrate t

Control

1

Boiled

None

Positive

Very pronounced

1 a

Unboiled

None

Positive

Very pronounced

lb*

Boiled

Slight

Positive

Very pronounced

1 c*

Unboiled

None

None

None

2

Boiled

None

Positive

Very pronounced

2a

Unboiled

None

None

Positive

2 b*

Boiled

None

Slight

Pronounced

2 c*

Unboiled

None

Slight

Very pronounced

3

Boiled

None

None

Slight

3a

Unboiled

None

Slight

Positive

3 b*

Boiled

None

None

None

3 c*

Unboiled

None

Slight

Positive

4 at
5a§

Boiled
Unboiled
Boiled
Unboiled

None
None
None
None

None
None
None
None

None
None
None
None

* Root removed at base of plants,
t Filtrate diluted 1 to 3.
X Duggar's nutrient solution.
§ 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

382 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

at the end of 3 hours and became quite definite at the end of
63^2 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 (pi. 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 (pi. 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. The toxic substance is approximately thermostable
in both the filtrate and mycelial extract used in these experi-
ments. The P H 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 ATK.

During the last few years different investigators have raised
the question as to the real parasitic nature of the fungus Fusarium
vasinfectum in connection with wilt of cotton. For instance,
Ajrekar and Bal ('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. He 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. He used qualitative microscopical tests for these metals.
Tissues of plants which were infected with Rhizoctonia Solani
Kiihn 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 accum-
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 2 , AK 4 , 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. In
some pots the upper lateral roots were cut before introducing the
fungus, but the tap roots were uninjured. His 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

384

[Vol. 14

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 Fe*Oi FOUND IN TKICE COTTON PLANTS*

Healthy

Total ash in leaves
Iron oxide in leaves
Total ash in stems
Iron oxide in stems
Total ash in roots
Iron oxide in roots

Total ash in plant
Iron oxide in plant

Per cent

14.32
0.028
3.75
0.006
3.45
0.016

21.32
0.050

Diseased

Total ash in leaves
Iron oxide in leaves
Total ash in stems
Iron oxide in stems
Total ash in roots
Iron oxide in roots

Total ash in plant
Iron oxide in plant

Per cent

18.54
0.04
4.11
0.008
3.70
0.014

26 . 35
0.062

* Average of 3 plants

Additional

plants in a 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

The results, howe\

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 anv close

correlation bet

iron accumulations in the tissues of the

plants and wilt infection. The 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 ('24) theory of aluminium and
iron accumulations in the tissues of plants as a primary factor
involved in producing wilt, another experiment bearing upon this

planned

the summer of 1926, a field was visited

South Mississippi Station which was badly infested with

and which had

A

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. The 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

the tissues of the

The plants were allowed

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
The results obtained are shown in table vi.

December 3

TABLE VI

EXPERIMENTS TO DETERMINE THE PATHOGENICITY OF COTTON WILT.

FUSARIUM VA8INFECTUM ATK., IN THE GREENHOUSE.

SEASON 1926— VARIETY DELFOS 6102

Pot No.

Soil treatment

Number infected plants

Apparent wilt Nov. 1

I

1

Unsterile

1A

Sterilized

2

Unsterile

2A

Sterilized

3

Unsterile

3A

Sterilized

4

Unsterile

4A

Sterilized

5

Unsterile

5A

Sterilized

6

Unsterile

6A

Sterilized

7

Unsterile

7A

Sterilized

8

Unsterile

8A

Sterilized

9

Unsterile

9A

Sterilized

Actual wilt Dec. 3

3

1

2

6

5

1

0§
8

8

2

* 2 dead with Rhizoctonia.

t 1 dead with Rhizoctonia.

X 3 killed by Fusarium vasinfeclum.

§ 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

[Vol. 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 (pi. 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. The 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. The recent work of
Butler ('26) 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 ('23) has obtained bearing upon this question as it
relates to corn root-rot. He 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. The 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. A 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. The 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. A
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 ('25) 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 a

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 corn and perhaps sugar-cane to invasion
by soil-inhibiting organisms, it was of interest to study further
the role that such compounds might play in predisposing cotton
plants to infection by Fusarium vasinfeclum. Accordingly, 2
series of water cultures in battery jars containing aluminium
salts in varying concentrations were prepared. Shive's ('16)
R 2 C B 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:

[Vol. 14

388 ANNALS OF THE MISSOURI BOTANICAL GARDEN

SERIES I

p.p.mil. Al.

1. Control, basic nutrient only None

2. Basic nutrient plus aluminium sulphate to make N/2500 ..3.6

3. Basic 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

Al

1. Control, basic nutrient only N

2. Basic nutrient plus aluminium chloride to make N/2500 ..3.6

3. Basic nutrient plus aluminium chloride to make N/1000 . . 9

4. Basic nutrient plus aluminium chloride to make N/500 ... 18

5. Basic nutrient plus aluminium chloride to make N/250 ... 36

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

the

tv

and

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. The effect
of these salts when added to the nutrient solution is best indicated

by the photographs in pi. 33, figs. 1

2. At the end of

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 vn.

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

Culture

Control

N/2500

N/1000

N/500

N/250

Length of tops
inches

Sulphate

11.5
12.0
10
12
8.4

Chloride

Condition of roots

Sulphate

12
10
9.5

10.5

7.8

Good — normal
Good — normal
Good — normal
Good — normal
Fair — few laterals

Chloride

Good — normal
Good — normal
Good — normal
Fair — few laterals
Poor — very few
laterals

After the length of plants in the above experiment had been
determined they were put back into the battery jars and to each

3's RsCr nutrient solution.

was added freshly

pared Shrv

Each culture was then immediately inoculated w r ith 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

which had received the basic

At

the end of one week after inoculation, all cultures were infected
with the fungus (pi. 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 litt
infection by Fusarium
Dastur C24).

ifectum as has been

sed by

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 and Pember ('15)
was used as a basic nutrient. This nutrient had the following
composition :

(Vol. 14

390 ANNALS OF THE MISSOURI BOTANICAL GARDEN

.2N Ca (N0 3 ) 2 .4H 2 15 cc. per litre

AN NH4NO3 10 cc. per litre

.1JVKC1 8 cc. per litre

.2N MgS0 4 .7H 2 8 cc. per litre

8.3 grams per litre CaH 4 (P0 4 ) 2 1 cc. per litre

FeP0 4 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:

Ser
Ser
Ser
Ser
Ser
Ser
Ser
Ser
Ser
Ser

es I — Basic nutrient only.

es II — Same as I but with phosphorus omitted.

es III — Same as I but with phosphorus increased 20 times.

3S IV — Same as I but with potassium omitted.

3S V — Same as I but with potassium increased 10 times.

3s VI — Same as I but with potassium increased 20 times.

3s VII — Same as I but containing 1/10 less nitrogen.

?s VIII — Same as I but with nitrogen increased 2 times.

5fl IX — Same as I but with calcium omitted.

58 X — Same as 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 NaN0 3 , NaJIPC^ 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. The 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 a week, and after 3 such weekly-
applications 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 a
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 October 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. A final inoculation was made on October
20, bits of diseased stems of mature Delfos 6102 cotton plants from
a 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. The numbers
with lateral root discoloration were recorded, and undoubtedly,

had been

ifectum could

probably have been isolated, though time would not permit such
cultural tests for evidence of the fungus. The results obtained
in these nutrition experiments are presented in table viii.

392

ANNALS OF THE MISSOURI BOTANICAL GARDEN

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19271

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 pis. 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 role 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 IX, 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

396 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

was high. The 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. The same may be said of increased 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. The
specific role 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 ('23) 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 ('24) theory of iron accumulation
as being a predisposing factor to invasion by the fungus. How-

this fact is not borne out by quantitat
s of diseased and healthy plants nor by

of

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 zinc 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. The 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. 1 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 as 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 ix.

TABLE IX

EFFECT OF COMMERCIAL FERTILIZER UPON THE DEVELOPMENT

COTTON WILT IN THE GREENHOUSE. RATE OF APPLICATION

800 POUNDS PER ACRE. SEASON 1926

OF

Fertilizer

Relative height of
plants, inches

Vigor

No.
plants

Infected plants after 60 days

formula

Number

Per cent

8-4-0 check

9.5

Good

8

8-4-0

10.0

Good

8

3

37.5

8-4-4 check

11.5

Good

8

8-4-4

10.5

Good

8

8-0-4 check

9

Good

8

8-0-4

8.5

Good

8

5

62.5

0-4-4 check

8.5

Fair

8

0-4-4

8

Fair

8

4

50

1 2-4-4 check

12

Good

8

12-4-4

11

Good

8

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. The results obtained are best indi-
cated in pi. 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

1 From information supplied the writer by Dr. Oswald Schreiner, of the Bureau
of Plant Industry, U. S. Dept. Agr.

398

[Vol. 14

ANNALS OF 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,
A. & M. College, and another at the Poplarville Branch Experi-
ment Station. The 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

Rate

Yield

seed

Total number plants

% infection

Fertil-

*

• -L

1 .

izer

per acre

cotton

in area*

1

in areaj

•

in area

in area

*

formula

lbs.

per acre
lbs.

inocu-
lated

inocu-
lated

inocu-
lated

inocu-
lated

Total

once

twice

once

twice

Check

140

219

42

3.50

5.56

9.06

8-4-8

600

712

288

63

2.20

3.17

5.37

8-4-4

600

572

267

59

3.87

7.35

11.22

8-4-0

600

260

264

61

4.16

5.47

9.63

Chock

128

258

55

3.49

6 . 67

10.16

8-8-4

600

684

268

65

3.01

4.30

7.91

8-6-4

600

512

251

54

4.25

6.18

10.43

8-0-4

600

492

227

52

5.00

7.06

12.06

Check

140

261

51

3.96

1.98

5.94

6-4-4

600

536

279

59

4.66

8.49

13.15

4-4-4

( >00

496

296

64

4.28

7.81

12.09

1 0-5-5

1 600

70S

267

56

3.74

4.80

8.54

Check

20S

257

55

4.41

4.85

9.26

8-4-4 t

600

652

223

42

4.50

6.76

11.26

8-4-4

000

724

273

53

4.03

7.55

11.58

8-4-4

1200

804

254

54

2.37

2.48

4.85

Check

152

226

47

3.70

5.00

8.70

* Soil inoculated May 4 by plowing under re-inoculated sterilized oats,
t Soil inoculated again August 9 by pouring 100 cc. of spore suspension of Fusarium
vasinfectum around roots of plants.

\ Lime applied at rate of 4500 pounds per acre, to approximately neutralize soil.

v

1927]

NEAL — COTTON WILT

399

The plots at the Central Station were seeded to Lone Star 65
cotton 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 in 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 cc. of a spore suspension of the fungus in
August. The 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.
The addition of lime at the rate of 4500 pounds per acre had no
effect in reducing wilt.

The 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

TREATMENTS 5 YEARS

Plot S

Plot 6

Plot 9

Plot 12

400 lbs. acid

400 lbs. acid

400 lbs. acid

phosphate

phosphate

phosphate

150 lbs. ammonium

150 lbs. ammonium

150 lbs. ammonium

300 lbs. ammonium

sulphate

sulphate

sulphate

sulphate

50 lbs. potassium

100 lbs. potassium

100 lbs. potassium

250 lbs. potassium

sulphate

sulphate

sulphate

sulphate

Plot 2

Plot 5

Plot 8

Plot 1 1

Check

Check

Check

Check

Plot 1

■ -— -■—■■-■■ i

Plot 4

Plot 7

Plot 1

400 lbs. acid

400 lbs. acid

200 lbs. acid

400 lbs. acid

phosphate

phosphate

phosphate

phosphate

150 lbs. ammonium

150 lbs. ammonium

150 lbs. ammonium

100 lbs. potassium

sulphate

sulphate

sulphate

sulphate

100 lbs. potassium

200 lbs. potassium

sulphate

sulphate

The 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. The data
obtained are presented in table xn.

400

[Vol. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

TABLE XII

OCCURRENCE OP COTTON WILT IN FERTILIZER PLOTS AT

POPLARVILLE

BRANCH EXPERIMENT STATION. SEASON 1926

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

3

1206

691

102

14.76

4

1278

773

8

1.04

5

648

810

18

2.22

6

1080

762

7

.92

7

1080

660

8

1.21

8

216

612

17

2.78

9

630

665

1

.15

10

486

613

11

72

512

1

.19

12 1

990

590

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,

ulting in a greater prevalence of the disease. At any rat<
iter observed the plants in plot 1 early in July and they
the time quite large, with wilt beeinninsr to anoear ove

the

entire plot.

aphs of the condition of the plants in plots

when they

and 12 were made on September 2, also on October

These

The

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

the disease may be kept in check by such field

treatments. A
graphed by the

field of Delfos 6102 cotton as shown

This

field had been planted to cotton for several years and had devel

oped considerable

At the suggestion of the County

1927]

NEAL — COTTON WILT 401

the grower fertilized the crop with 700 pounds per acre of an
8-4-4 fertilizer and left a small portion of the field untreated as
a 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.

Field Tests of 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. All of the varieties received 600 pounds
per acre of an 8-6-4 fertilizer. The 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

the

season of 1926 are summarized in table xm 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 a good crop of bolls, especially when the plants are
heavily fertilized and the season for growth is favorable.

402

[Vol. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

TABLE XIII

TESTS OP WILT-RESISTANCE AT MISSISSIPPI EXPERIMENT STATION

SEASON 102(5

Variety

Trice Mississippi Sta.
Half & Half, Mahon
Cook, Rhyne
Cook, 307-6
Toole, Mathis
Toole, Perry
Cleveland, Pied.

Cleveland, Humco 20

Cleveland 54

Super Seven

Dixie Triumph

Kelly Big Boll

Sol. & Oates Big Boll

Miller

D. & P. L. No. 4

D. & P. L. No. 5

Watson

Lightning Express

Delf os 6 1 02

Pounds per acre

Lint data

Seed
cotton

891
534
762
654
741
753
903
657
828
888
888
660
531
528
669
600
822
846
771

Lint

272.6
205.6
287.1
228.9
263.1
255.3
293.5
231.9
278.2
291.3
293.9
236.3
188.0
178.5
244.9
192.0
259.8
256.3
241.3

Per
cent

30.6
38.5
36.5
35.0
35.5
33.9
32.5
35.3

33.6
32.8
33.1
35.8
35.4
33.8
36.6
32.0

31.6
30.3
31.3

Length
inches

Hi
%i

%
1

l

i f

Cents
per lb.

Total

dollars
per acre

13.37
11.48

12.03
11.67
12 37
12 03
12.37
12.03
12.70
14 53
12 70
12.95
13.78
13.37
13.37
17.53
16.53
16.53
14.53

43.87

27 . 54
39.27
31.81
38 . 28

36 . 68
43 . 62
33 . 00
41.93
49.49
44.46
35 . 68
30 . 03

28 . 06

37 . 83

38.56
49 . 69
49 . 45
41.42

Hank

in
value

5

19

9

16

11

13

6

15

7

2

4

14

17

18

12

10

1

3

8

Summary

1. The 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

b

Such loss varies from year to year, and

amount of it depends somewhat on environmental factors
3. The disease causes a yellowing of the leaves of the <

usu

accompanied by a stunted appearance of the main stems,
y rather early in the season. Under certain conditions

outbreak

e growing season

The characteristic internal symptom, as in the case of other
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. The fungus is placed in the recognized generic sub-
division "Elegans." This species produces readily (in culture)

1927]

NEAL — COTTON WILT 403

micro- and macro-conidia and chlamydospores. The latter are
formed in corn-meal or potato-dextrose agar cultures in less than
3 weeks. The color of the stroma is marked and the acid effect
well emphasized. The 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. The fungus grows slowly at temper-
atures below 10° C, the optimum temperatures being 28-30° C,
and the maximum temperature about 38° C. It grows well over

rather wide range of hyd

The best

growth occurred in cultures represented by P H values of 3.0,
4.0, 4.5, and 5.5. The maximum growth occurred in cultures
started at P H 3.5. The fungus tolerates a culture solution strongly
acid. The 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 case 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 toxic 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. The
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 Fusarium vasin-
fectum was no more evident in plants subjected to different

AHA lVOL - 14

4U4 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. The 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-

tash, and from 4 to 6

] >hosph

Acknowledgments

In conclusion, the writer wishes to express his grateful appreci-
ion to Dr. 13. M. Duggar, under whose guidance the laboratory
d greenhouse aspects of the problem have proceeded, and to
ank him for the many suggestions and helpful criticisms which

19271

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
Drivileees and facilities of the Missouri Botanical Garden.

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23: 601-823. pi. 1-21. 1923.

, and Trost, J. F. ('23). The 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. 1923.

Hopkins, E. F. ('21). Growth and germination of Gibberella saubinetti at varying
hydrogen-ion concentrations. Phytopath. 11: 36. 1921.

Hutchinson, C. M. ('13). Rangpur tobacco wilt. India Dept. Agr., Mem. Bact.
Ser. 1: 67-83. 1913.

Jones, L. R., and Tisdale, W. B. ('22). The influence of soil temperature upon the
development of flax wilt. Phytopath. 12: 409-413. /. /. 1922.

Kirby, R. S. ('22). The take-all disease of cereals and other grasses. Ibid. 68-88.
pi 2-4, f. 1-3. 1922.

Lehman, S. G. ('23). Pod and stem blight of soy bean. Ann. Mo. Bot. Card. 10:
111-178. pi. 9-13 } f. 1-13. 1923.

Lewis, A. C. ('11). Wilt disease of cotton in Georgia and its control.
Bd. Ent., Bui. 34: 1-31. 1911.

Lint, H. C. ('14). Report of potato scab experiments. N. J. Agr. Exp. Sta.,
Ann. Rept. 35: 477-488. 1914.

Ga. State

, ('15). Ibid. 36: 618-625. 1915.
, ('16). Ibid. 37: 615-625. 1916.

Maclnnes, J. ('22). The growth of the wheat scab organism in relation to hydrogen-
ion concentration. Phytopath. 12: 290-293. /. /. 1922.

Martin, W. H. ('20). The relation of sulphur to soil acidity and to the control of
potato scab. Soil Sci. 9: 393-408. /. 1-5. 1920.

('21). A comparison of inoculated and uninoculated sulphur for the

control of potato scab. Ibid. 11: 75-85. pi. 1, f. 1-3. 1921.
McGeorge, W. T. ('25). The influence of aluminium, manganese and iron salts

upon the growth of sugar cane, and their relations to the infertility of acid island

soils. Hawaiian Sugar Planters' Assoc. Bui. 49: 1-95. /. 1-33. 1925.
Orton, W. A. ('00). The wilt disease of cotton. U. S. Dept. Agr., Div. Veg.

Physiol, and Path. Bui. 27: 1-16. pi 1-4, f. 1-7. 1900.

•, ('08). Cotton wilt. U. S. Dept. Agr., Farmers' Bui. 333: 1-24. /. 1-

11. 1908.

, and Gilbert, W. W. ('12). The control of cotton wilt and root knot.

U. S. Dept. Agr., Bur. PI. Ind. Circ. 92: 3-19. /. 1-12. 1912.
Picado, C. ('23). Sur Taction h distance des champignons phytopathogenes.
Cong. Path. Veg. Strasbourg, pp. 28-34. 1923.

Rast, L. E. ('22). Control of cotton wilt by the use of potash fertilizer. Am. Soc.
Agron., Jour. 14: 222-224. /. 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. /. IS.

1926.
Shaffer, P. A., and Hartman, A. F. ('21). The iodometric determination of copper

and its use in sugar analysis. Jour. Biol. Chem. 45: 349-390, 1921.
Shive, J. W. ('16). A study of physiological balance in nutrient media. Physiol.

Res. 1: 327-397. 1916.
Smith, E. F. ('99). Wilt disease of cotton, watermelon, and cowpea. U. S. Dept.

Agr., Div. Veg. Physiol, and Path. Bui. 17: 1-53. pi 1-10. 1899.

Stevens, F. L. ('21). Diseases of economic plants. 507 pp. Rev. ed. New York,
1921.

Tisdale, W. B. ('23). The influence of soil temperature and soil moisture upon the

Fusarium disease in cabbage seedlings. Jour. Agr. Res. 24: 55-86. pi. 1-2, f.

1-10. 1923.
Webb, R. 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. 1919.

, ('21). Ibid. XV. Ibid. 8: 283-341. /. 1-89. 1921.

Wollenweber, H. W., et al. ('25). Fundamentals for taxonomic studies of Fusarium.

Jour. Agr. Res. 30: 833-843. /. 1. 1925.
Young, H. C., and Bennett, C. W. ('21). Studies in parasitism. 1. — Toxic substances

produced by fungi. Mich. Acad. Sci., Ann. Rept. 22: 205-208. 1921.

[Vol. 14, 1927J

408 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 31

Fig. 1. A cotton plant infected with the wilt fungus, Fusarium vasinfedum.
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 discoloration of the fibro-vascular tissues. At
right, three sections from a healthy plant.

Fig. 3. Photomicrograph of Fusarium vasinfedum taken from a 12-day-old
culture of the fungus growing on potato-dextrose agar showing terminal and inter-
calary formation of chlamydospores. X 100.

Fig. 4. Photomicrograph of Fusarium vasinfedum taken from a 10-day-old
culture of the fungus growing on potato-dextrose agar showing micro- and macro-
conidia. X 100.

•

Ann. Mo. Bot. Gard., Vol, 11, 1927

Plate 31

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410 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 32

Fig. 1. Experiments to determine the pathogenicity of cotton wilt, Fusarium
vasinfcctum Atk., in the greenhouse. Season 1926.

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 Duggars solution and cot-
ton seedlings. At right, cotton seedlings in flasks containing filtrates from 18-day
cultures of Fusarium vamifectum grown on Duggar's solution. Note toxic effect.
All flasks photographed after treatment of 36 hours.

Fig. 3. At left, uninoculated control flasks containing sterile distilled water and
cotton seedlings. At right, cotton seedlings in flasks containing extracts of mycelial
mats from 18-day cultures of Fusarium vasinfcctum grown on Duggar's solution.
Note toxic effect. All flasks photographed after treatment of 36 hours.

Ann. Mo. Bot. Gaud.. Vol. 14, 1927

Plate 32

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NEAL -COTTON WILT

[Vol. 14, 1927]

412 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 33

Fig. 1. Delfos 6102 cotton seedlings grown in Shive's R 2 C 6 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 in Shive's R 2 C 6 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. 3. Delfos 6102 cotton seedlings grown in Shive's R 2 C 5 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 R 2 C 6 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 inoculated jars are infected.

Aw. Mo. Bot. Gard., Vol. I I, 1927

Plate 33

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N E AL— COTTO N W [LT

[Vol. 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 II.

Fig. 3. Series III.

Fig. 4. Series IV.

Fig. 5. Series V.

Fig. 6. Series VI.

Ann. Mo, Hot. Gard., Vol. 14, 1927

Plate 3 I

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NEAL— COTTON WILT

[Vol. 14, 1927]

416 ANNALS OP THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 35

Variety of cotton in each series, Delfos 6102. All plants photographed after
growing in inoculated sand cultures and receiving nutrients indicated for approxi-
mately 7 months.

Fig. 1. Series VIII.

Fig. 2. Series IX.

Fig. 3. Series X.

Fig. 4. Series XI.

Fig. 5. Series XII.

Aw Mo. But. (Iahd., Vol. 11, 1927

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NKAL— COTTON WTLT

]Vol. 14, 1927]

418 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of Plate

PLATE 36

Fig. 1. 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 1926.

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 May hew,
Mississippi. Season 1926.

Ann. J\Io. Hot. Gard., Vol. 14, [927

Plate 36

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X K A L— COTTON WIF.T

[Vol. 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
and 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.

Fig. 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.

t

\\\. Mo. Hot. Gard., Vol. 14, 1927

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[Vol. 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.

Fig. 2. A view of a cotton field near Aberdeen, Mississippi, showing the effect
of a complete fertilizer on wilt control. The rows at left were not fertilized. The
rows at right received 700 pounds per acre of an 8-4-4 fertilizer. Season 1926 s;

Ann. Mo. Bot. Gard., Vol. 14, 1927

Plate Ms

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NEAL— COTTON WILT

[Vol 14, 1927J

424 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Explanation of 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 1926.

Fig. 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. Mil LJot. Gard., Vol. 11, 1927

Platk 3<J

NEAL— COTTON WILT

SPECIES OF CERCOSPORA ON SMILAX IN THE

UNITED STATES

L. O. OVERHOLTS

Prof,

Mycologist

WashingL

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 Sacc;
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 Smilax 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 \x. The spores also are distinctive, being
practically cylindric, 10-celled or more when mature, and measure
45-120 X 5-6 [x. 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. Mo. Bot. Gard., Vol. 14, 1927

(425)

[Vol 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 (pi. 41,
fig. 4), i.e., pale fuscous or pale cinnamon. The 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. As a
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 (pi. 41, fig. 7). Like-
wise the color of the conidiophore becomes darker with age, so
that while in a young condition it may appear rather pale, in an
older stage it assumes a 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 \l long, and essentially the same measure-
ments are obtained from de Thuemen, 'Myc. Univ.' No. 1670.
However, in Nash, 'Plants of Florida,' No. 1893, they measure
45-135 (jl and in a collection by Peck at Manor, L. I. (Herb. N. Y.
State Mus.) they measure 72-225 ji. 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 [jl, while one at Arcadia, Mo. (Overholts Herb. 10426)
has them 75-210 u.: in 'Plants of the Gulf States,' No. 7802. thev

1927)

OVERHOLTS — SPECIES OF CERCOSPORA ON SMILAX 427

are 150-180 p long; and in Nash, 'Plants of Florida/ No. 2125,
they are 105-165 [x. 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 on 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. One of these
is C. Smilacis de Thuemen, in which the conidia are subcylindric,
tapering very gradually to a narrowed apex, and hence in reality
slightly obclavate when seen on the conidiophores. Frequently
they are considerably curved. They measure for the most part
60-135 X 4-5 [a, but some as short as 40-50 \i 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 pi. 40).

The 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. The 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 y., but a few are as short as 30 n

and 4-5 [i in diameter, while occasionally a spore as much as 75 (i
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 pi. 41).

How often it occurs that Cercospora conidia are clavate and not
obclavate on the conidiophores, remains for further study to
determine. As originally founded by Fresenius ('Beitrage zur
Mykologie,' p. 91, 1863) the genus is based on C. Apii, which he
illustrates as with obclavate conidia, and the usual conception

428 ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol. 14

of the genus involves that characteristic. While I have not seen
a large number of clavate spores on conidiophores in C. Smilacina,
yet they have been observed several times, though always ob-
A iously immature, and at a later stage of maturity they may
possibly take a more obclavate form.

I find nothing in the character of the spots produced by these

species to aid in their macroscopic separation,
considerable extent on the different hosts, with

They

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
colled ions the margin is not well marked. The 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.

A few words may be said regarding C. Petersii. The species
was originally described as a H elminthosporium (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

able to arrive at anv decision

s

of

species. In the sense of Atkinson (I.e.) and as determined in
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.

PI. 40.
C. 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 [i long, 3-5 p 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 ix.

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. N. 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 Sacc. Michelia 2: 364. 1881. PI. 41.

t C. Petersii (Berk. & Curt.) Atk. Jcur. Elisha Mitchell Sci-

entif. Soc. 8: 25. 1892.

f 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 ^ 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)

X 6tx.

On living leaves of various species of Smilax.

The species is easily separated from C. Smilacis by the conidia
that are rather abruptly narrowed near the middle, and less
than 75 u long.

430

[Vol. 14, 1927]

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, Economic 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 of Plate

PLATE 40

Cercospora Smilacis

Fig. 1. Conidia from de Thuemen, Myc. Univ. 1768. X 400.

Fig. 2. Fascicle of conidiophores emerging through a stoma. Ellis, N. Am.
Fungi, 1251. X 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, N. 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. X 400.

Ann. Mo. Bot. Gard., Vol. II. 1927

Plate in

OVERHOLTS CERCOSPORA ON SMI LAX

[Vol. 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.
X420.

Fig. 3. Conidia. From Seymour & Earle, Economic Fungi, No. 199. X 420.
Fig. 4. Conidiophores in young condition. From collection at Auburn, Ala.,
1896. X 420.

Fig. 5. Conidia. Same as fig. 4.

Fig. 6. Conidia and two conidiophores. From Bartholomew, Fungi Col. 2808.
X420.

Fig. 7. Fascicle of conidiophores. From collection at Arcadia, Mo., 1926,
Overholts Herb. 10426. X 165.

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.

Fig. 1 1 . Two conidiophores and two conidia. From collection at Wading River,
N .Y., Peck, 1879. X 420.

Aw. Mo. But. Gaud., Vol. 11, 1927

Plate 1 1

OYKRIlol/rS-rKRCosmRA ON SMILAX

GENERAL INDEX TO VOLUME XIV

New scientific names of plants and the final members of new combinations are
printed in bold face type; synonyms and page numbers having reference to figures
and plates, in italics; and previously published scientific names and all other matter,
in ordinary type.

A

A. & M. College, Mississippi, cotton
wilt experiments at, 360, 398

Agaricus Aegerita, 139; aureus, 122;
aurivellus, 152; autumnalis, 134; cape-
rafus, 123; dactyliotus, 174; detersi-
biliSf 161] flammariSf 148; hormophorus,
175; Johnsonianus, 130; luciferus, 162;
marginatum, 132; mutahilis, 138; prae-
cox, 106; radicoms, 175; temnoph yllus ,
112; togularis, 114; unicolor, 134

Aluminium compounds, relation of cot-
ton wilt to, 383, 387, 395

Aplopappus florifer , 19, var. /3, 21

Aster Wrightii, 32; exscapus, 28

B

Bystropogon axillare, 82; minutus, 83;

uniflorus, 82

c

Calcium, effect of, on development of
cotton wilt, 395

Cercospora, species of, on Smilax in the
United States, 425

Cercospora, 425; Apii, 427; mississip-
piensis, 428; nuhilosa, 425; Petersii,
429; Smilacina, 429, 482; Smilacis,
428, 430; subsan guinea, 425

Cotton, wilt-resistant strains of, 401

Cotton wilt, a pathological and physio-
logical investigation, 359; description
of, 364; factors involved in producing,

376
Cryptantha, section Oreocarya, 237

affinis, 307; aperta, 295, 354; Bakeri
331, 358; Bradburiana, 307, 356
breviflora, 318; caespitosa, 281
352; cana, 316, 356; celosioides, 299
354; Clemensae, 267, 350; conferti-
flora, 256, 348; echinoides, 321
elata, 285, 354; flava, 259, 348; flavo-
culata, 334, 358; fulvocanescens

319; glomerata, 307; holoptera, 270
humilis, 278, 352; insolita, 273
350; interrupta, 296, 354; Jamesii
242, var. abortiva, 250, var. cinerea
246, var. disticha, 248, var. laxa, 246

Ann. Mo. Bot. Gabd., Vol. 14, 1927

var. multicaulis, 244, 348, var.
typica, 248; Jonesiana, 323, 356;
leucophaea, 262, 348; longiflora,
326, 358; Macounii, 303, 356;

mensana, 333, 358; modesta, 277,
3»,l; nana, 312, var. commixta, 312,
var. ovina, 314, var. Shantzii, 313,
356, var. typica, 315, 354; nubi-
gena, 265, 350; oblata, 254, 348;
Osterhoutii, 329, 358; Palmeri, 253;
paradoxa, 330, 358; propria, 317,
356; pterocarya, 270; pustulosa, 252;
rugulosa, 295, 354; salmonensis,
263, 348; sericea, 286, var. perennis,
288, var. typica, 287, 354; setosis-
sima, 268, 350; Sheldonii, 301, 354;
sobolifera, 305, 356; spiculifera,
298, 354; stricta, 264, 350; suffruti-
cosa, 242; tenuis, 327, 858; thyrsi-
flora, 283, 852; tumulosa, 276, 352;
utahensis, 270; virgata, 270, 850;
virginensis, 274, 352; Wetherillii,

324, 358
Cynoglossum glomeratum, 307

D

Dioscorea villosa, 425

E

Epling, Carl C. Studies on South Ameri-
can Labiatae, 111, 47

Erigeron florifer, 19

Eritrichium section Pseudomyosotis , 237

Eritrichium fulvocanescens , 319; glomera-
tum, 303, 307, var. fulvocanescens, 278,
var. hispidissimum, 283, var. humile,
278, 287, var. virgatum, 270; Jamesii,
248; leucophaeum, , 262 ; multicaule , 244;
setosissimum, 268; virgatum, 270

F

Fertilizers, commercial, effect of, on
development of cotton wilt, 396

Flammula pulchrifolia, 148; unicolor,
136; viridans, 148

Fungicides used in cotton wilt experi-
ments, 362

(433)

434

[Vol. 14

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fusarium vasinfectum, 360; morpho-
logical and cultural characters of, 364;
pathogenicity of, 382; physiological

studies of, 366

G

Gardoquia, 47; acutifolia, 63; argentea,
71; brevi flora, 59; chili n sis, 56; dis-
color, 62; elliptica, 54; fasciculate,, 72;

foliotoea, 61; Oilliesii, 56; glabrata, 66;

incana, Q4',Jamesonii, 69; nderophylla,
70; multiflora, 53; ohovata, 58; revoluta,

74; rugosa, 76; aahidefolia, 84; sericea,

73; striata, ()8; taxifolia, 67; thymoides,

61; tomentosa, 60

H

Hartvvcll & Pcmber's nutrient solution,

389
Hebeloma hortense, 105, 107

Helmintho8porium Pctersii, 429

llesperothymus of Satureia, 50
Hydrogen-ion concentration, effect of,

on germination of fungi, 370

Hypodendrum oregom use, 140

I

Iron compounds, relation of, to cotton
wilt, 383, 395

K

Krynitzkia depressa, 277 r ; echinoides, 321 ;
fidvocanescens, 319, var. idakoensis,

317; glomerate, 307, var. acuta, 324,
var. virginensis, 274; Jamesii, 248;
leucophaea, 262, var. aZata, 256;
mensana, 333; multicuulis var. ofcor-

/tt>a, 250, var. setosa, 244; oblate, 254;
Palmeri, 253; pustulate, 307; sericea,
287, 316; sttosissima, 268; virgata,
270

Krynitzkia, section Pterygium, 237,
section Pseudo krynitzkia, 237

L

Labiatae, Studies on South American,

III, 47
Larson, lather L. A revision of the

genus Townsendia, 1

M

Maianthcmum canadense, 425
Micromeria, 47; boliviano, , 80; bonarien-

.s/.s", 84; Darwinii, 78; eugenioido , 83;

Gilliesii, 83; nubigena, 76; pasilla, 79;

pulchella, 84

Micromeria section Hesperothymus, 50

Moisture conditions, effect of, on cotton
wilt, 377

Monograph of the genus Pholiota in the

United States, 87
Monograph of the section Oreocarya of

Cryptantha, 211
Myosotis glomerate, 307; leucophaea y 262;

suffruticosa, 242, 248

N

Neal, David C. Cotton wilt: a patho-
logical and physiological investiga-
tion, 359

N cocas mospora vasinfrta, 360

Nitrogen, effect of, on development of
cotton wilt, 393, 395

Nutrition, effect of, upon the develop-
ment of cotton wilt, 389

o

Oreocarya of Cryptantha, a monograph
of the section, 211

Oreocarya, 237; abortiva, 250; uffinis, 307,
var. pi renins, 288; aioto, 256; aperta,
295; argentea, 287; luiheri, 331; 6rm-
Jfora, 318; caespitosa, 281; rexna, 316;
celosioides, 299, 301; ciiio-lirxuta, 298;
enured, 246; commixta, 312; conferti-
flora, 256; cristata, 334; depressa, 277;

disticha, 248; dolnsa, 313; dura 9
Eastwoodae, 334; echinoides, 278, 321 ;

e/a/a, 285; eulophus, 331; jZaaa, 259;
flavoeulata, 334, var. spatulata,
fulvocanesa its, 319 ; glomerate,
307; gypsophila, 330; hispida,
hispidissima. 254, 283; holopiera, 270;
horridula, 326; humilis, 278; insolita,
273; interrupta, 296; Jonesiana, 323;
Lemmoni, 246; leucophaea, 262, var.
confetti flora, 256; longiflora, 326; Zutea,
266; lutescens, 259; Macbridii, 278;
Macounii, 303; mensa.ua, 333; mono-
sperma, 283; multicaulis, 244, var.
cinerea, 246, var. /axa, 246; nana, 312,
315; nitida, 319; nubigena, 265;
oblate, 255; Osterhoutii, 329; Palmeri,
253; puradoxa, 330; Paysonii, 255;
perennis, 288, 307; procera, 287;
propria, 317; pustulosa, 252; rugulosa,
295; salmonensis, 263; sericea, 287,
301; setosissima, 268; Shantzii, 313;
Sheldanii, 301; Shockleyi, 334; spicata,
270; s j) i calif era, 298; striete, 264;
suffruticosa, 248, var. abortiva, 250,
var. m/.< rca, 246, var. multicaulis, 244;
tenuis, 327; thyrsijlora, 283; tumulosa,
276; urticacea, 283; virgata, 270,
forma spicata, 270; virginensis, 274;
WetherillU, 324

283;

334;
301,

278;

1927]

INDEX

435

Overholts, L. 0. A monograph of the
genus Pholiota in the United States,
87 ; Species of Cercospora on Smilax in
the United States, 425

P

Payson, E. B. A monograph of the
section Oreocarya of Crypt antha, 211

Pholiota, a monograph of the genus, in
the United States, 87

Pholiota, 99; Acericola, 108, 182; adi-
posa, 154, 202 ', 204; Aegerita, 139;
Aegerita var. strobiloidca, 140; aerugi-
nosa, 147, 198; aggericola, 119, 121,
174; albivelata, 128; albocrenulata,
150, 200; angustipes, 166, 208; anoma-
la, 127; appendiculaia , 174;aurea, 121,
186; aurivella, 152, 210; aurivel-
loides, 151; autwnnalis, 134, 174;
blattaria, 113; caperata, 123, 188;
cerasina, 141; comosa, 169, 174; con-
fragosa, 160, 206 ', 208; curvipes, 164;
dactyliota, 174; destruens, 168; deter-
sibilis, 161, 174; discolor, 135, 192;
dura, 105, 174, var. xanthophylla , 105;
duroides, 129; erebia, 118, 184) erina-
ceella, 161; filaris, 116; flammans, 148,
200; fulvo-squamosa, 167, 188; fur-
cata, 136; heteroclita, 169, 174; hormo-
phora, 174; Howeana, 110; in decern,
121, 175; Johnsoniana, 130; limonella,
163; lucifera, 162; lutea, 145, 175;
luteofolia, 143; luxurians, 175; margi-
nata, 132, 190; marginella, 137;
McMurphyii, 124; minima, 125; mol-
licula, 175; muricata, 161, 190; muta-
bilis, 138; mycenoides, 115; ombro-
phila, 120; oregonense, 140; ornella,
175; platyphylla, 117; praecox, 106,
180;radicosa, 175;rigidipes, 158;rube-
cula, 142; rugosa, 115, 192; sabulosa,
175; Schraderi, 159; speciosa, 175;
spectabilis, 144, 194, 196; sphalero-

morpha, 109, 111, 175; squarrosa, 157,
£04; squarrosoides, 155, 206; sub-
nigra, 120; subsquarrosa, 151, 156, 176;
temnophylla, 112; terrestris, 126;
terrigena, 127, 176; togularis, 114, 192;
togidaris var. filaris, 116; trachyspora,
125; tuberculosa, 164, 190; unicolor,
134; ventricosa, 146, 176; vermiflua,
104, 178; villosa, 176; washingtonensis,
119, 121, 176

Phosphorus, effect of, on development of
cotton wilt, 392, 395

Poplarville Experiment Station, Missis-
sippi, cotton wilt experiments at, 398

Potash, effect of, on development of
cotton wilt, 397

Potassium, effect of, on development of

cotton wilt, 392, 395

R

Revision of the genus Townsendia,
Rizoa, 47; ovatifolia, 53
Rochelia glomerata, 307

1

s

Satureia, synopsis of the genus, 47
Satureia acutifolia, 63; Andrei, 67;
argentea, 71; axillaris, 82; boliviana,
80, var. tarijense, 81; bonariensis,
84; brevicalyx, 82; breviflora, 59;
Brownei, 51, subsp. eubrownei, 51;
chilensis, 56; connata, 70; Darwinii,
78; discolor, 62; elliptica, 54; ericoides,
70; eugenioides, 83; fascicnlata, 72;
foliolosa, 61; Gilliesii, 56; Gilliesii, 83;
glabrata, 66; guamaniensis, 58; in-
signis, 74; Jamesoni, 69; Kuntzeana,
80; Lindeniana, 75; Loesneriana, 55;
mantaroensis, 63; Matthewsii, 57;
microphylla, 70; multiflora, 53; nubi-
gena, 76, var. glabrescens, 78; obo-
vata, 58; oligantha, 83; pallida, 65;
Pavoniana, 64; plicatula, 69; pusilla,
79; revoluta, 74; rigidula, 72; rugosa,
75; sericea, 73; simulans, 81;
striata, 68; taxifolia, 67; thymoides,
61; tomentosa, 60; vana, 79; Weber-
baueri, 72
Smilax, species of Cercospora on, in the

United States, 425
Stachys speciosa, 54
Stenotics florifer, 19, var. 0, 21
Stropharia Howeana, 110; Schraderi, 159
Studies on South American Labiatae,

III, 47

T

Temperature, influence of, on cotton

wilt, 366
Thymus, 47; Brownei, 51 ; humifusus, 76;

nubigenus, 76; taxifolius, 67
Townsendia, a revision of the genus, 1
Townsendia, 8; alpigena, 17; alpina, 22;
ambigua, 21; arizonica, 27; arizonica
X incana, 27; Bakeri, 16; condensata,
22; dejecta, 17; eximia, 12, 42; ex-
scapa, 28, 46; exscapa, 31, 46, var.
Wilcoxiana, 29; Fendleri, 24; florifer,
19, 42; florifer var. communis, 19;
formosa, 11, 36, 42; Fremontii, 26;
glabella, 16; grandiflora, 13; incana,
25; incana var. ambigua, 26, var.
prolixa, 26; intermedia, 29; lepotes, 28;
leptotes, 27; mensana, 30; mexicana,
23; montana, 17; Parryi, 9; Parryi
var. alpina, 22; pinetorum, 11; Roth-

[Vol. 14, 1927]

436 ANNALS OF THE MISSOURI BOTANICAL GARDEN

wilt

rocku, 16; Bcapigera, 20; scapigera Toxic substances in cotton

var. ambigua, 21, var. caulescens, 21; 380

sericea, 30, 44, 46; sericea, 28, 4! u 46,

var. leptotes, 27, /3 papposa, 29; X

spathulata, 22; strigosa, 24, 40;

strigosa, 18, 19; texensis, 15, 38; Xenopoma, 47, 76; bolivianum, 80, var.

Vreclandii, 12; Watsom, 18; Wil- tarijense, 81; eugmioides, 83

coxiana, 29; Wrightii, 32

Volume XIV

Number 1

Annals

of the

Missouri Botanical

Garde

n

Contents

February, 1927

A Revision of the Genus Townsendia

Esther L Lars en

1-46

Studies on South American Labiatae. III. . . . Carl C. Epling 47-86

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