BULLETIN OF THE UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED STATES NATIONAL HERBARIUM

VoLuME 34.

SYSTEMATIC PLANT STUDIES

Vera Lyora Cotsry, WituiamM L. Stern, Georce K. Brizicxy,
Francisco N. Tamo_ane, Epwarp S. Avyensu, Taomas R. SopErstrRom,
Aaron Gotpserc, Ricnarp W. Pipren, WitL1aM Louis CuLBERSON,
and Carcita F. CULBERSON

SMITHSONIAN INSTITUTION PRESS * WASHINGTON, D.C. © 1968

Publications of the United States National Herbarium

The United States National Herbarium, which was founded by
the Smithsonian Institution, was transferred in the year 1868 to the
Department of Agriculture and continued to be maintained by that
department until July 1, 1896, when it was returned to the official
custody of the Smithsonian Institution. The Department of Agri-
culture, however, continued to publish the series of botanical reports
entitled “Contributions from the United States National Herbarium,”
which it had begun in the year 1890, until, on July 1, 1902, the Na-
tional Museum, in pursuance of an act of Congress, assumed
responsibility for the publication. The first seven volumes of the
series were issued by the Department of Agriculture.

Frank A. Taytor,
Director, United States National Museum.
ii

Parr 1.

Parr 2.

Part 3.

Parr 4.

Parr 5.

Parr 6.

Part 7.

CONTENTS

Diagnostic characteristics of the fruits and florets of
economic species of North American Sporobolus.
By Vera Lyole vary: (Published December 18,
1957) . Ck ee ,

The woods and flora of the Florida Keys: Cappara-
ceae. By William L. Stern, George K. Brizicky,
and Francisco N. Tamolang. (Published Novem-
ber 4, 1963)

Systematic anatomy and ontogeny of the stem in
Passifloraceae. By Edward S. Ayensu and William
L. Stern. (Published August 21,1964)... . .

Taxonomic study of subgenus Podosemum and sec-
tion Hpicampes of Muhlenbergia (Gramineae). By
Thomas R. Soderstrom. (Published July 19, 1967) .

The genus Melochia L. (Sterculiaceae). By Aaron
Goldberg. (Published August 31, 1967)

Mexican ‘‘Cacalioid” genera allied to Senecio (Com-
positae). By Richard W. Pippen. (Published
May 17, 1968) . Lo

The lichen genera Cetrelia and Platismatia (Par-
meliaceae). By William Louis Culberson and
Chicita F. Culberson. (Published July 10, 1968) .

Page

1-24

25-43

45-73

75-189

191-363

. 365-447

449-558

ill

UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED STaTES NATIONAL HERBARIUM

VoLuME 34, Parr 1

DIAGNOSTIC CHARACTERISTICS OF THE FRUITS
AND FLORETS OF ECONOMIC SPECIES
OF NORTH AMERICAN SPOROBOLUS

By Vera Lyota CoLBry

BuLLETIN OF THE UNITED States NatrionaL Museum

SMITHSONIAN INSTITUTION e WASHINGTON, D. C. e 1957

Contents

Introduction .

Historical consideration .

Materials and methods
Acknowledgements ..........
Structural and diagnostic characteristics .
Use of keys and descriptions .
Terminology .

Taxonomic treatment .

Key to fruits .
Key to florets
Descriptions .

Literature cited

Figures
1. Illustrations of terms used to designate fruit and seed morphology .
2. Illustrations of terms used to designate shape .

Plates

1-4. Sporobolus fruits, seeds, and florets.
ats

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HP OoOT ao aAWNnNnNnNe wp B®

hoo
ng

DIAGNOSTIC CHARACTERISTICS OF THE FRUITS
AND FLORETS OF ECONOMIC SPECIES OF NORTH
AMERICAN SPOROBOLUS

By Vera Lyota CoLsry

Seed Technologist, United States Department of Agriculture,
Beltsville, Md.

Introduction

This study is designed primarily for use in laboratories engaged in
the testing of agricultural seed, where the identification of isolated
seeds, fruits and florets is an integral part of the analytical procedure.
Twenty of the twenty-nine species of Sporobolus occurring in North
America are herein described. These were selected on the basis of
their economic value and their possible occurrence in seed samples.

Historical consideration

With the rapid growth of seed testing within the last 30 years the
need for taxonomic descriptions of seeds has been intensified, but
there are relatively few detailed descriptions, keys, or illustrations
available. For the most complete list to such publications the reader
is referred to the annotated bibliography by Murley (1951). U. 8S.
Agriculture Handbook 30 (1950), contains descriptions, keys, and
illustrations of many of the seeds handled as agricultural crops and
occurring as incidentals in them; and the Handbuch der Samenkinde,
by W. Bower and A. Staéhlin (1955), contains descriptions and illus-
trations of approximately 2,500 seeds, but it has no keys and the
illustrations in general do not show much diagnostic detail.

The supplementary information as to the economic value and occur-
rence in seed samples was obtained principally by means of question-
naires sent to the Soil Conservation Service and to seed laboratory
personnel.

Information pertaining to the geographic distribution and habit
was taken from A. S. Hitchcock, ‘Manual of the Grasses of the United
States” (2 ed., revised by Agnes Chase, 1950).

The only reference found to the diagnostic characteristics of the
florets and fruits of Sporobolus appears in U. S. Department of Agri-
culture Handbook 30 (1950), which consists of a short descriptive

key to five species, three of which are illustrated.
1

2 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Materials and methods

In order to arrive at a generalized or typical description for each
species, fruits were examined from all of the mature specimens in the
U. S. National Herbarium of the North American Sporobolus which
were considered in this study. In addition, bulk “‘seed’’ samples
obtained from the Soil Conservation Service, U. S. Department of
Agriculture, and from a few seed laboratories were studied.

The material was examined with the use of a binocular microscope
largely at magnifications of 35, although certain structural details
were frequently confirmed at magnifications of 45 or 60. Measure-
ments, by means of a binocular eyepiece micrometer, were made of
several fruits and florets from each specimen and averages for each
species calculated. The number of individual fruits and florets
measured for the various species ranged from 25 to 496, depending
on the number of samples or specimens examined and the amount of
variation in the material.

Acknowledgements

The writer is indebted to Dr. Lyman B. Smith for his guidance in
the preparation of this paper, and to Dr. Jason R. Swallen for making
available for study the plant collections at the U. S. National Her-
barium, Smithsonian Institution, Washington, D. C.

The manuscript was reviewed by Dr. Oren L. Justice of the Federal
Seed Laboratory, Beltsville, Md., and the fruit key was tested by
seed analyst Elizabeth Fleming. Seed analysts Mildred A. Mauldin
and Lou Coffey submitted supplementary fruit and floret material for
study.

The following members of the staff of the Soil Conservation Service
were particularly helpful in contributing fruit or plant specimens for
study and for submitting information as to the economic uses of
Sporobolus: Dr. A. A. Hanson, Dr. Jack Harlan, M. D. Atkins, Jack
EK. Engleman, and Joe A. Downs.

Dr. D. B. Judd, of the National Bureau of Standards, assisted in the
selection and use of the Munsell Pocket Edition Constant Hue Charts
from which the basic fruit colors were determined.

Structural and diagnostic characteristics

The florets of Sporobolus are from 1-flowered spikelets, so if free
florets are found in seed samples there will be no rachilla present.
The lemma and palea are membranaceous in the majority of species,
although a few are found in which they are chartaceous, coriaceous,
or indurate. The lemma, is 1-nerved, awnless, and the palea is 2-

1 For a complete list of the material examined the reader is referred to the original thesis on file in the
library of the George Washington University.

COLBRY—DIAGNOSTIC CHARACTERISTICS OF SPOROBOLUS 3

nerved, the keels broadly winged. In most species the palea splits
between the two nerves as the fruit ripens. The fruit is free from the
lemma and palea, usually falling readily from the floret at maturity.
It is this characteristic that necessitates fruit descriptions.

The fruit in the grass family as a whole is commonly known as a
caryopsis, since the pericarp is adherent to the seed. However, in
Sporobolus the pericarp is loose. In all but one species herein consid-
ered the pericarp is thin and finely striate or longitudinally reticulate,
slipping off easily when moist. This type of fruit is referred to as an
utricle. Sporobolus heterolepis has a pericarp which is thick, hard
and brittle, not slipping off when moist. It is designated either as
an utricle or as “nutlike.”’

The fruit characteristics found to have diagnostic value were:
Shape, size, and color of the fruit and seed; texture and surface mark-
ings of the pericarp; shape, color, and relative measurements of the
scutellum and the radicle-plumule axis in relation to each other and
to the endosperm; and in one species, the appearance of the area
around the hilum.

The florets did not have distinctive diagnostic characteristics in all
instances. It was only possible to delimit certain species or groups
of species by means of floret differentiations. ‘The diagnostic charac-
teristics found to be of value were: texture, indumentum, length
and width, general shape, shape of the lemma and palea tips, and

color.
Use of keys and descriptions

Two keys were prepared. The key to the fruits is to individual
species, while the key to the florets delimits certain species or groups
of species. In cases where intact florets are found the latter key may
ensure more rapid identification of the material. Owing to the
slight differences in some of the species the fruit key is made very
descriptive.

The descriptions of the species in the keys represent the most
typical material studied. The full taxonomic descriptions should
be used in order to identify correctly highly variable species.

Since color is an important diagnostic characteristic, an attempt
was made to classify the colors of the fruits according to the Munsell
color system. Munsell Pocket Edition Constant Hue Charts were
used. The colors given are essentially the same as noted on the
color chips with which the fruits were compared, but such terms as
strong brown to designate a dark reddish brown were not used, in
order to avoid confusion as to the exact shade meant. The detailed
taxonomic descriptions of the fruits and florets of each species are
preceded by the name and basonym, after which appears the following
information in the designated order: Specimen illustrated; common

4 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

stylar tip
endosperm
embryo area:
scutellum edge
radicle~plumule axis
stylar tip styler tip
endosperm
embryo area:
scutellum edge
radicle-plumule axis

hilun

Ficure 1.—IIlustrations of terms used to designate fruit and seed morphology:
a, Lateral view; 4, dorsal view; ¢, ventral view; d, fruit with pericarp partially

5
J,

Ficure 2.—Illustrations of terms used to designate shape: a, Ovoid; b, obovoid;
ce, ellipsoid; d, oblong; d, oval; 2, lanceolate.

name or names; geographic distribution; habit; economic importance;
and occurrence in seed samples.

The structure of Sporobolus fruits and florets and the terms used
to designate shape in the keys and descriptions are illustrated in
figures 1 and 2. In plates 1-4 are depicted the fruits and florets for
each species.

COLBRY—DIAGNOSTIC CHARACTERISTICS OF SPOROBOLUS 5

The range of dimensions are given in the fruit key and both the
range and median in the descriptions. The fruit measurements do
not include the stylar tip and basal stipe of the pericarp.

Terminology

Certain terms as used in the keys and descriptions are defined
below:

Ca: approximately.

CHARTACEOUS: papery in texture.

Corrackovus: leathery in texture.

Cucu.uaTE: hood-shaped.

Dorsat: the embryo side of the fruit.

Eccenrric: one-sided.

Emsryro Area: the scutellum and the radicle-plumule axis.

INDUMENTUM: any covering, as hairiness or glandularity.

MEMBRANACEOUS: thin, like a membrane.

NaRROWLY: (as applied to shape) at least twice as long as wide.

PaNDURIFORM: fiddle-shaped, drawn in at the middle.

Pittep: marked with indentations or small hollows.

PiumBeEous: lead-colored, or greenish drab.

RETICULATE: netted.

ScaBERULOUS: minutely scabrous (covered with short, stiff hairs).

STRAMINEOUS: Straw colored, or pale yellow.

SrriaTe: marked by a series of fine parallel lines.

TRANSLUCENT: semitransparent; admitting passage of light but diffusing it so
that objecis beyond cannot be clearly distinguished.

TRANSPARENT: clear; having the property of transmitting rays of light so that
objects beyond can be clearly distinguished.

VENTRAL: the hilum side of the fruit.

Taxonomic treatment

Certain species of Sporobolus separated only by slight inflorescence
divergences were found to have equally slight fruit and floret differ-
ences. Perhaps the most difficult in which to find diagnostic charac-
teristics for individual species occurred in the Sporobolus cryptandrus
group. Some specimens of S. cryptandrus with enclosed panicles,
usually occurring late in the season, are very similar to S. contractus.
Fruit differences in color, shape, size, and to a certain extent opaque-
ness and flecking are fairly reliable, although a positive identification
might not be possible in all cases. The only significant floret difference
is size.

Sporobolus contractus and S. giganteus are very similar. There is a
definite overlap between size and thickness of panicles, size of florets
and size of fruits on some specimens. Color and shape of fruits are
similar, size being the only variable. It would be impossible to
identify accurately all fruits of these two species. The type specimen
of S. giganteus (Nash 394) is typical for the description, having dense,

6 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

thick panicles and large fruits and florets. The type specimen of
S. cryptandrus var. robustus (Neally 746), cited as a synonym for S.
giganteus, has panicle, fruit, and floret measurements in the overlapping
size range between S. giganteus and S. contractus.

Sporobolus flecuosus and S. nealleyi have definite individual fruit
characteristics as well as vegetative differences in the plants. The
florets cannot be distinguished in all cases.

The herbarium specimens of Sporobolus cryptandrus were carefully
examined and measured for subspecific variation, as indicated by
Jones and Fassett (1950). The overlap in the specific diagnostic
characteristics is too great for positive allocation of all specimens into
the subspecific categories designated by them, although there are
certain broad regional differences.

Florets and fruits of the following species intergrade to some extent:
Sporobolus vaginiflorus with S. clandestinus, and S. airoides with S.
wrightii; the fruits have more individual diagnostic characteristics
than the florets and can be distinguished in most instances. The
similarity of the fruits and florets of S. vaginiflorus and S. clandestinus
to each other, of interest since the former is an annual and the latter
a perennial, is the only instance in which plants of unlike habit over-
lapped in fruit and floret characteristics. Sporobolus airoides and S.
wright are both perennials, being separated principally by a slight
inflorescence divergence.

While the plants of Sporobolus vaginiflorus and S. neglectus are very
similar, their identification by fruit and floret characteristics was
found to be very dependable. The fruits of the latter are variously
mottled while those of the former are not, and there are also differences
in size and shape. The florets of S. neglectus are chartaceous and
glabrous while those of S. vaginiflorus are coriaceous or indurate
(except for very occasional cleistogamous florets) and usually sparingly
pubescent.

The published descriptions of the florets of S. vaginiflorus do not
cover the range of characteristics for the species. The florets are
described as sparsely pubescent, sometimes mottled with dark spots.
Statements as to the texture of the lemma and palea are not complete.
Examination of the National Herbarium specimens revealed that the
indumentum of the florets varied from sparsely pubescent to shiny,
microscopical, conical glands. This latter type of floret always has
a palea which is split between the two nerves, the split so complete
as to cause the palea to appear double. The florets are indurate and
mottled, with plump, abruptly acute to incurved or cucullate lemma
and palea. Specimens with this kind of floret occur in the limestone
areas of Arkansas and Missouri. There is a gradual intergrading of
this type with the narrower, thinner textured, slightly pubescent floret

COLBRY—DIAGNOSTIC CHARACTERISTICS OF SPOROBOLUS 7

occurring in other regions. It was thought that perhaps the charac-
teristic of the palea splitting between the two nerves might be of
some significance, but it could not be correlated with any other con-
stant variable in the florets or fruits. In general, specimens from the
northeastern and eastern states have paleas that do not split at fruit
maturity. Those from the central, south, and southwestern states
have either split or nonsplit paleas.

The form of Sporobolus neglectus from the Ozark mountains, desig-
nated by Fernald (1933) as S. ozarkanus, has florets and fruits like the
other forms of S. neglectus examined. The type which he described
as S. vaginiflorus var. inaequalis intergrades with florets and fruits of
S. vaginifiorus from other areas.

The varieties of Sporobolus asper were not studied due to lack of
available mature fruit material.

Sporobolus patens was not included, as only one specimen was avail-
able for study. This species has a limited distribution in the United
States, being reported only from southern Arizona. The fruits are so
similar to those of S. pulvinatus that it is doubtful whether these two
species can be distinguished.

Key to fruits

A. Pericarp thick, hard, brittle and opaque; fruits subglobose, ca. 1.0-2.0 mm.
in diameter. ......... +... . + +1. S. heterolepis (pl. 1,a)

A. Pericarp thin, mucilaginous when wet, hyaline; striate or reticulate.
B. Embryo as long as seed to slightly shorter; fruits narrowly eccentric obo-
void, plump, tapering to base; length 2.1-3.0 mm., width 1.0-1.5 mm.
2. S. interruptus (pl. 1,8)

B. Embryo not more than 434 length of seed.
c. Fruits eccentric ovoid, translucent; seeds smooth, not microscopically
pitted; pericarp extremely finely striate, scarcely evident.

p. Fruits bulging dorsally above embryo tip; endosperm light yellowish
brown or dark orange-brown.

gE. Embryo usually less than 14 length of seed; fruits narrowly ovoid;

endosperm usually faintly reddish flecked; scutellum brown or

reddish; length 2.2-3.6 mm., width 0.6-1.3 mm., thickness

0.4-1.0mm........... 3. S. clandestinus (pl. 1,¢)

E. Embryo usually more than ¥% length of seed (often 34); fruits nar-
rowly or broadly ovoid; endosperm rarely reddish flecked;
scutellum brown, reddish, or blackish; length 1.0-3.0 mm.,
width 0.4-1.5 mm., thickness 0.3-1.4 mm.

4. S. vaginiflorus (pl. 1,p)

p. Fruits bulging ventrally toward base; narrowly ovoid; endosperm
light yellowish brown or yellowish orange; embryo area reddish

brown; length 1.0-1.5 mm., width 0.4-0.7 mm., thickness 0.2-0.5
mm................ 5 S, texanus (pl. 1,5)

c. Fruits rarely eccentric ovoid (if so, not translucent); seeds either micro-
scopically pitted or pericarp evidently striate or reticulate.

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

F. Fruits dark shades of red, orange, or brown, or fruits gray, dark reddish
brown flecked or mottled; pericarp frequently dark red or black
striate.

a. Scutellum edge prominent, dark red or black at seed maturity.

H. Fruits tapering to thin edges dorsally and ventrally, strongly
flattened laterally, broad.

1. Fruits ellipsoid; endosperm orange or reddish brown; trans-
lucent at maturity; length 1.7—-2.6 mm., width 0.8-1.2 mm.,
thickness 0.4-0.6mm. .. . 6. S. purpurascens (pl. 1,F)

1. Fruits broadly obovoid or broadly ovoid; endosperm reddish or
reddish brown; opaque or semitranslucent; length 1.5-1.8
mm., width 1.0—-1.2 mm., thickness 0.4—0.7 mm.

7. S. junceus (pl. 2,4)
H. Fruits not tapering to thin edges dorsally and ventrally.

J. Embryo 34 length of seed; seeds reddish brown mottled; endo-
sperm usually grayish and semitranslucent; shape subov al
or suboblong, plump.

k. Fruits in dorsal view sharply keeled above embryo tip;
seutellum edge narrow (ca. 0.1 mm.); radicle-plumule
axis broad, twisted; length 0.8-2.5 mm., width 0.4—1.0
mm., thickness 0.8-0.7 mm. . 8. S. neglectus (pl. 2,B)

kK. Fruits in dorsal view convex the entire length; scutellum edge
broad (ca. 0.2-0.8 mm.); radicle-plumule axis narrow,
almost straight; length 1.1-2.0 mm., width 0.6—-1.3 mm.,
thickness 0.5-1.l mm... .. . 9. S. asper (pl. 2,c)

J. Embryo less than 34 length of seed; seeds not mottled; endo-
sperm reddish or yellowish orange; shape suboblong,
usually strongly compressed laterally; pericarp dark reddish
striate, highly mucilaginous at seed maturity, and then
frequently forming a dark rim on the edges of the fruits;
length 0.8-1.4 mm., width 0.5-0.8 mm., thickness 0.2-0.4
mm.... ~ 2... 10 S. poiretii (pl. 2,p)

a. Scutellum edge not prominent, about same color as endosperm;
pericarp strongly dark red or black striate; fruits plump, length
0.9-1.7 mm., width 0.4-0.9 mm.

L. Stylar tip rounded; fruits suboblong or suboval, dark yellowish
brown or light brown; seeds rarely reddish flecked; thickness
0.4-0.7 mm.... toe ee _ ILS. airoides (pl. 2,8)
L. Stylar tip usually pointed and strongly laterally compressed;
fruits predominantly ovoid, dark reddish brown (occasionally
light brown); seeds usually reddish flecked; thickness 0.3—0.7
mm... ... . 12. S. wrightii (pl. 2,Fr)

F. Fruits light shades of red, orange, brown, or gray, never dark reddish
brown mottled; pericarp never dark red or black striate, but
frequently light red, rusty, or purplish striate.

M. Fruits flattish at base, embryo without a prominent overhang; hilum
within a basally prominent black half-rim; fruits suboval or
broadly ovoid; uniformly light reddish orange; endosperm
faintly light reddish flecked; length 0.7-1.0 mm., width 0.3-
0.7mm... .... 13. S. buckley (pl. 3,4)

mM. Fruits pointed at base, embryo with a prominent overhang; hilum
not within a basally prominent black half-rim.

COLBRY—DIAGNOSTIC CHARACTERISTICS OF SPOROBOLUS 4)

n. Scutellum in dorsal view pandurate; in lateral view of fruits, inner
scutellum reflected in a broad, dark semicircle through the
endosperm; radicle-plumule axis broad.

o. Embryo area greenish black; fruits light yellowish gray or yel-
lowish orange; obovoid; length 0.8-1.0 mm., width 0.4—-0.7
mm., thickness 0.2-0.5 mm . . 14. S. pulvinatus (pl. 3,8)
o. Embryo area dark reddish brown; fruits light orange reddish;
obovoid, suboval or subelliptic; length 0.7-1.0 mm., width
0.4-0.6 mm., thickness 0.2-0.4 mm.
15. S. pyramidatus (pl. 3,c)
nN. Scutellum in dorsal view oval; in lateral view of fruit, inner
scutellum either not reflected or narrowly and unevenly re-
flected through the endosperm; radicle-plumule axis narrow.
p. Fruits extremely light yellowish brown or pale orange-yellow,
sometimes almost white; endosperm not rusty or light red-
dish flecked but usually internally “starchy” or granularly
mottled.
qa. Fruits not strongly flattened laterally; average length less
than 1.0 mm.; pericarp frequently purplish striate, some-
times rusty striate.
r. Fruits broadly obovoid, frequently ‘‘pinched’’ laterally
toward the center; embryo area rusty or light reddish
brown; length 0.7-0.9 mm., width 0.5-0.8 mm.
16. S. flexuosus (pl. 3,D)
r. Fruits narrowly ovoid, not “pinched” laterally toward the
center; embryo area rusty or purplish brown; length
0.7-1.0 mm., width 0.3-0.5 mm.
17. S. nealleyi (pl. 3,£)
qa. Fruits strongly laterally flattened; average length over 1.0
mm.; pericarp not purple striate, but sometimes rusty
striate; fruits broadly elliptic or broadly ovoid.
s. Fruit length 0.8-1.2 mm., width 0.4-0.8 mm.
18. S. contractus (pl. 3,F)
s. Fruit length 1.0-1.9 mm., width 0.6-0.9 mm.
19. S. giganteus (pl. 4,a)
p, Fruits darker: moderate orange, brownish orange or light reddish
brown; endosperm faintly rusty or light reddish flecked,
rarely internally “starchy” or granularly mottled; fruits
laterally flattened at stylar tip; ovoid or obovoid; pericarp
sometimes rusty or light reddish striate; length 0.6-1.3
mm., width 0.3-0.8 mm .. 20. S. eryptandrus (pl. 4,B)

Key to florets

A. Florets coriaceous, indurate, or chartaceous.

B. Florets coriaceous or indurate; pubescent or microscopically striate with
shiny conical glands; opaque; color gray, tan, or purplish, unevenly
splotched with black, purple, or green.

c. Florets 4.5-13.0 mm. long; palea 0.5-6.0 mm. longer than the lemma;
palea acuminate (rarely acute), not split between the two nerves;
lemma acute or acuminate ... . . 3. S. clandestinus (pl. 1,c)
c. Florets 1.6-7.0 mm. long; palea subequal to 2.0 mm. longer than lemma;

430126—5T——2

10 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

lemma and palea acuminate, acute or incurved or cucullate; palea
sometimes split between the two nerves . . 4. S. vaginiflorus (pl. 1,p)
zs. Florets chartaceous; glabrous; smooth; opaque or semiopaque; color white,
greenish or purplish tinged, or purplish; length 1,1-3.0 mm,
8. S. neglectus (pl. 2,B)
a. Florets membranaceous; essentially glabrous (some species sparingly micro-
scopically scaberulous) ; opaque, semiopaque, or transparent.
p. Length of florets 3.0 mm. or over.
gE. Florets 1.0 mm. thick or over.
y. Lemma and palea acute; florets ca. 1.1 mm. thick, 1.1 mm. wide, and
6.0mm.long. ... . ... . .2 S.interruptus (pl. 1,8)
ry, Lemma and palea subacute; florets distended by the globose fruits; ca.
2.0 mm. thick, 2.0 mm. wide, and 4.0 mm. long.
1. S. heterolepis (pl. 1,A)
v. Florets less than 1.0 mm. thick (S. asper occasionally 1.1 mm. thick).
a. Florets bronze; 3.0-4.0 mm. long and ca. 1.0 mm. wide; lemma acute;
fruits often sticking to the edges of the palea.
H. Palea acute... ....... . .6. S. purpurascens (pl. 1,F)
H. Palea obtuse or subacute... .. . . . .7. S.junceus (pl. 2,4)
a. Florets not bronze.
1. Floret length 4.0-7.0 mm., two to three times as long as the fruit;
lemma and palea subacute; color pale yellowish gray, light green
or purplish tinged... . . ... . . . .9. S. asper (pl. 2,c)
1. Floret length 2.2-3.5 mm., not more than twice as long as the fruit;
lemma and palea acute or acuminate; color stramineous or

plumbeous. . . . .. es. . 19. S. giganteus (pl. 4,4)
p. Length of florets under 3.0 mm.
J. Florets bronze tinged... ........ . .11. S. airoides (pl. 2,5)

12. S. wrightii (pl. 2,F)
J. Florets not bronze tinged.
K. Palea obtuse, usually erose and dentate; lemma acute (sometimes
subobtuse).
L. Florets frequently smutted; fruits usually sticking to the outside
or tips of the florets; length 1.5-2.0 mm.
10. S. poiretii (pl. 2,p)
u Florets not smutted; length ca. 1.0-2.0 mm.
13. S. buckleyi (pl. 3,4)
14. S. pulvinatus (pl. 3,8)
15. S. pyramidatus (pl. 3,c)
Kk. Palea acute; lemma acute.
M. Florets usually opaque or semiopaque.
nN. Floret length 2.1-2.7 mm.; fruits frequently sticking to the outside
or tips of the florets... . . . . . .5. S. texanus (pl. 1,5)
nN. Floret length 1.5—2.0 mm.; florets usually dark purplish tinged.
17. S. nealleyi (pl. 3,£)
mM. Florets usually transparent or semiopaque (rarely opaque).
o. Floret length (1.4)-1.8-(2.0) mm. . . 20. S. eryptandrus (pl. 4,B)
o. Floret length (1.5)-2.1-(2.7) mm. . . 16. S. flexuosus (pl. 3,D)
o. Floret length (1.5)-2.4-(2.9) mm. . . 18. S. contractus (pl. 3,F)
o. Floret length (2,2)-2.9-(3.5) mm. . .19. S. giganteus (pl. 4,A)

‘
CONTR. U. S. NAT. HERBARIUM, VOL. 34 COLBRY, PLATE 1

A, S. heterolepis, X 6.3 B, S. interruptus, X 6.3

C, S. clandestinus, N 6.3 D, S. vaginiflorus, X 6.3

=

I, S. texanus, X% 11.3 F, S. purpurascens, X 9.4

SPOROBOLUS FRUITS, SEEDS, AND FLORETS.

CONTR. U.S. NAT. HERBARIUM, VOL. 34 COLBRY. PLATE 2

A, S. junceus, X 9.4 B, S. neglectus, X 10.7

C. S. asper, X 10.7 D, S. potretti, N 11.35

ES. atrotdes, NX V1.3 By S. cortghtit, NX 113

SPOROBOLUS FRUITS, SEEDS, AND FLORETS.

CONTR. U.S. NAT. HERBARIUM, VOL. 34

A, S. buckley, X 11.3

2

A

‘on VA
we
te ——
i 4 aN f

C, S. pyramidatus, X 11.3

Ih, S. nealleyi, X 11.3

COLBRY, PLATE 3

D, S. flexwosus, X 11.3

I, S. contractus, X 11.3

SPOROBOLUS FRUITS, SEEDS, AND FLORETS.

CONTR. U.S. NAT. HERBARIUM, VOL. 34

COLBRY, PLATE 4

\, S. eiganteus, X% 11.3

B, S. eryptandrus, X 11.3

SPOROBOLUS FRUITS SEEDS, AND FLORETS.

COLBRY—DIAGNOSTIC CHARACTERISTICS OF SPOROBOLUS 11

Descriptions
1. Sporobolus heterolepis (A. Gray) A. Gray, Man. 567. 1848. Based on
Vilfa heterolepis A. Gray. PLATE 1,4
Vilfa heterolepis A. Gray, Ann. Lyc. N.Y. 3:233. 1835. Watertown, New
York, Crawe.

Fruits: Length, width and thickness ca. the same: (1.0)-1.5-(2.0)
mm.; opaque; shape subglobose, eccentric in latera) view; color
grayish yellow, frequently greenish gray or purplish splotched;
pericarp thick, brittle, indehiscent and minutely longitudinally
reticulate.

Seeds: These were removed from the pericarp with extreme diffi-
culty. Although reported as being free from the pericarp, they are
partially adherent at the stylar tip.

Length, width, and thickness ca. the same as for the fruits; opaque;
longitudinally blackish or reddish striate or reticulate, and glandular
shiny; shape irregularly obovoid or skull-like, flattened dorsally;
color of endosperm variable (yellow, yellowish gray, light brown,
grayish greenish yellow, or grayish reddish brown) ; embryo strikingly
darker, sometimes almost black; embryo large, prominent, almost
completely covering the dorsal side of the seed; scutellum very broad;
radicle-plumule axis short and narrow.

Florets: Length (3.0)—4.0-(4.9) mm.; width and thickness ca. the
-same as for the fruits; membranaceous; glabrous; color plumbeous or
stramineous, frequently purplish or greenish tinged; palea slightly
longer and wider than the lemma; nerves of the lemma and palea
glabrous or slightly scaberulous toward the tips; palea at fruit ma-
turity splits completely between the 2 nerves, the 2 apices subacute;
lemma subacute, at maturity distended at the base by the subglobose
fruit.

Specimen illustrated: C. O. Johnston, December 1920, in meadow,
Manhattan, Kansas.

Common name: Prairie dropseed.

Geographic distribution: Quebec to Saskatchewan, south to Con-
necticut, eastern Texas, and Colorado; prairies.

Habit: Perennial, in dense tufts.

Economic importance: A true prairie species, disappearing under
moderate to heavy grazing; common in hay meadows; common in
eastern Kansas, but seed has never been produced or collected in
quantity for seeding.

Occurrence in seed samples: Not reported.

12 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

2. Sporobolus interruptus Vasey, Bull. Torrey Bot. Club 15:8, 1888. Arizona,
Coues and Palmer 66 in 1886; San Francisco Forest, Rusby 15 in 1883
(the Rusby specimen, distributed as No. 885, the type). PLATE 1,8

Fruits: Length (2.1)-2.6-(8.0) mm.; width (1.0)-1.1-(1.5) mm.;
thickness (1.0)-1.1-(1.3) mm.; opaque or semitranslucent; shape
narrowly eccentric obovoid; convex ventrally, plump, tapering to
the base; embryo large, prominent, almost completely covering the
dorsal side of the seed; color of endosperm yellowish reddish brown,
grayish tinged; embryo darker; pericarp brownish black striate or
reticulate, the ridges and interspaces grayish mucilaginous.

Florets: Length ca. 6.0 mm.; width and thickness ca. the same
as for the fruits; opaque; membranaceous; glabrous; color plumbeous
or stramineous; lemma and palea about the same length, the tips
acute; nerves of palea glabrous; nerve of lemma glabrous to minutely
scaberulous toward the tip; palea splits at fruit maturity.

Specimen illustrated: Joe A. Downs 951, Soil Conservation Service
“‘seed’’ sample, Arizona.

Common name: Black dropseed.

Geographic distribution: Arizona, grassy plains and hills.

Habit: Perennial, densely tufted.

Economic importance: Good forage grass, but doubtful if
it will ever be used in revegetation.

Occurrence in seed samples: Not reported.

3. Sporobolus clandestinus (Biehler) Hitche., Contr. U. 8. Nat. Herb. 12:150.

1908. Based on Agrostis clandestina Biehler. PLATE I,c

Agrostis clandestina Biehler, Pl. Nov. Herb. Spreng. Cont. 8. 1807. Penn-
sylvania, Muhlenberg.

Fruits: Length (2.2)-2.9-(3.6) mm.; width (0.6)-0.9-(1.38) mm.;
thickness (0.4)—0.6-(1.0) mm.; translucent; shape narrowly eccentric
ovoid, bulging dorsally above the embryo tip; embryo usually less
than % the length of the seed; color of endosperm and 1adicle-plumule
axis light yellowish brown or dark orange brown; endosperm usually
faintly reddish flecked; scutellum edge broad, flat, bandlike, darker
than the endosperm and the radicle-plumule axis; pericarp almost
smooth, sometimes faintly reddish striate.

Florets: Length (4.5)~-7.2—(13.0) mm.; width and thickness ca. the
same as for the fruits; opaque; coriaceous; sparingly long-pubescent;
microscopically striate with shiny conical glands; color gray, tan, or
purplish, unevenly splotched with black, purple, or green; palea
(0.5)-1.6—-(6.0) mm. longer than the lemma; lemma acute or acumi-
nate; palea usually acuminate, rarely acute; palea does not split
at fruit maturity.

Specimen illustrated: Jason R. Swallen 5625; Nov. 16, 1938; dry
sandy ground east of Gainesville, Florida.

COLBRY-—DIAGNOSTIC CHARACTERISTICS OF SPOROBOLUS 18

Common name: Hidden dropseed.

Geographic distribution: Connecticut to Wisconsin and Kansas,
south to Florida and Texas; sandy fields, pine barrens, hills and
prairies.

Habit: Perennial.

Economic importance: Palatable; especially valuable in winter
when stock seem to prefer it to other warm-season grasses.

Occurrence in seed samples: Reported in native grass and legume
seed samples, in samples of S. eryptandrus, and in lespedeza (Lespe-
deza spp.) samples.

4, Sporobolus vaginiflorus (Torr.) Wood, Class-book ed. 1861. 775. 1861.

Based on Vilfa vaginiflora Torr. PuaTE 1,p

Vilfa vaginiflora Torr. ex. Gray, N. Amer. Gram. and Cyp. 1: No.3. 1834;
Trin., Mém. Acad. St. Pétersb. VI. Sci. Nat. 41:56. 1840. New Jersey.

Fruits: Length (1.0)-2.0-(3.0) mm.; width (0.4)-0.7-(1.5) mm.:
thickness (0.3)—-0.5-(1.4) mm.; translucent; shape narrowly or broadly
eccentric ovoid, bulging dorsally above the embryo tip; embryo
usually more than half length of seed, frequently three-quarters; color
of endosperm and radicle-plumule axis light yellowish brown or dark
orange brown; endosperm rarely reddish flecked; scutellum edge
broad, flat, bandlike, at full maturity strikingly darker than the
endosperm and the radicle-plumule axis (sometimes almost black) ;
pericarp almost smooth, rarely faintly reddish striate.

Florets: Length (1.6)-3.3-(7.0) mm.; width and thickness ca. the
same as for the fruits; opaque; coriaceous or indurate; usually spar-
ingly pubescent; microscopically striate with shiny, conical glands (6
specimens observed in which the indumentum consisted of these shiny,
conica] glands alone); color gray, tan or purplish, unevenly splotched
with black, purple, or green; length of palea in relation to the lemma
variable, but never more than 2.0 mm. longer than the lemma; lemma
and palea tips acuminate, acute or incurved to cucullate; palea
usually does not split at fruit maturity (only 10 specimens were ob-
served with split paleas) ; cleistogamous florets like the others or rarely
membranaceous, whitish.

Specimen illustrated: Vera L. Colbry 17; Oct. 24, 1954; low clay
ground, near stream; Prince Georges County, Maryland.

Common name: Poverty dropseed.

Geographic distribution: Maine and Ontario to Minnesota and
Nebraska, south to Georgia, Texas, and Arizona; sandy soil or open
waste ground.

Habit: Annual.

Economic importance: Considered a weed; no value except for
erosion control when it invades abandoned fields.

14 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Occurrence in seed samples: Has been found in native grass and
legume seed samples, in S. cryptandrus samples, and in lespedeza
(Lespedeza spp.) samples from the southeastern states.

5. Sporobolus texanus Vasey, Contr. U. 8S. Nat. Herb. 1:57. 1890. Screw
Bean, Presidio County, Texas, Nealley (755). PLATE 1,5

Fruits: Length (1.0)-1.3-(1.5) mm.; width (0.4)-0.5-(0.7) mm.;
thickness (0.2)—0.4-(0.5) mm.; translucent; shape narrowly eccentric
ovoid, bulging ventrally toward the base; color of endosperm light
yellowish brown or yellowish orange; embryo area reddish brown;
scutellum edge narrow, the inner part slightly sunken around the
elevated, twisted radicle-plumule axis; pericarp very finely striate or
reticulate, frequently partially slipped off mature fruits, causing them
to adhere to the florets.

Florets: Length (2.1)-2.3-(2.7) mm.; width and thickness ca. the
same as for the fruits; opaque or semiopaque; membranaceous;
essentially glabrous, sometimes sparingly scaberulous toward the tips;
color plumbeous or stramineous, frequently greenish or purplish
tinged; palea slightly shorter than the lemma; lemma and palea tips
acute, the nerves minutely scaberulous toward the tips; palea splits
at fruit maturity; fruits with the pericarp partially slipped off fre-
quently adhering to the tips or sides of the florets.

Specimen illustrated: S. M. Tracy 8197; Oct. 6, 1902; Carlsbad,
New Mexico.

Common name: Texas dropseed.

Geographic distribution: Kansas and Colorado to Texas and
Arizona; mesas, valleys, and salt marshes.

Habit: Perennial, in close, hemispherical tufts.

Economic importance: Palatable, but no place in a seeding pro-
gram; one of the first grasses to come into alkaline flats in river bottom
succession; eventually crowded out by climax forms.

Occurrence in seed samples: Not reported.

6. Sporobolus purpurascens (Swartz) Hamilt., Prodr. Pl. Ind. Oce. 5. 1825.

Based on Agrostis purpurascens Swartz. PLATE 1,F
Agrostis purpurascens Swartz, Prodr. Veg. Ind. Oee. 25. 1788. Jamaica,
Swartz.

Fruits: Length (1.7)—2.1-(2.6) mm.; width (0.8)-1.0-(1.2) mm.;
thickness (0.4)-0.5-(0.6) mm.; translucent at maturity; shape ellips-
oid; strongly laterally flattened; convex laterally, and tapering to
thin, sharp edges dorsally and ventrally; slightly bulging ventrally
and Jaterally toward the base; color of endosperm orange or reddish
brown; scutellum edge broad (ca. 0.2 mm. wide toward the lower edge
and 0.1 mm. wide at the upper edge), the outer rim elevated; scutellum
dark, black at full seed maturity; radicle-plumule axis elevated and
strongly keeled, the same color as the endosperm; pericarp dull,

COLBRY—DIAGNOSTIC CHARACTERISTICS OF SPOROBOLUS 15

minutely striate, the striations often reddish; pericarp readily slipping
off mature fruits.

Florets: Length 3.0 to 4.0 mm.; width and thickness ca. the same
as the fruits; palea transparent; lemma semiopaque, membranaceous,
glabrous; color bronze, sometimes purplish tinged; lemma and palea
tips acute, usually spread apart at the tips, exposing the fruits; palea
splits at fruit maturity; fruits with the pericarp partially slipped off
often sticking to the lemma and palea.

Specimen illustrated: C. L. Lundell & Amelia A. Lundell 8846;
May 10, 1940; in sand along railroad; live-oak belt; north of Encino,
Brooks County, Texas.

Common name: Purple dropseed.

Geographic distribution: Southern Texas and eastern Mexico; West
Indies to Brazil; sandy prairies.

Habit: Perennial, in dense bunches.

Economic importance: Not reported.

Occurrence in seed samples: Not reported.

7. Sporobolus junceus (Michx.) Kunth, Rév. Gram. 1:68. 1829. Based on

Agrostis juncea Michx. PLATE 2,4

Agrostis juncea Michx. Fl. Bor. Amer. 1:52. 1803, not A. juncea Lam.,
1783. Carolina, Michaux.

Fruits: Length (1.5)-1.7—(1.8) mm.; width (1.0)-1.0-(1.2) mm.;
thickness (0.4)-1.6—-(0.7) mm.; opaque or semitranslucent; shape
broadly obovoid or broadly ovoid; strongly laterally flattened;
convex laterally and tapering to fairly thin edges dorsally and ven-
trally; slightly bulging ventrally and laterally toward the base;
color of endosperm reddish or reddish brown; scutellum, radicle-
plumule axis and pericarp the same as for S. purpurascens.

Florets: Palea tips obtuse or subacute; otherwise the same as for S.
purpurascens.

Specimen illustrated: Albert Ruth 91; July 1898; dry sandy soil,
Lookout Mountain; DeSoto Falls, Alabama.

Common name: Pineywoods dropseed.

Habit: Perennial, in dense bunches.

Economic importance: Not reported.

Occurrence in seed samples: Not reported.

8. Sporobolus neglectus Nash, Bull. Torrey Bot. Club 22:464. 1895. Mas-
sachusetts to Kentucky, Tennessee, and Kansas (type, Woodruff Gap,
New Jersey, Britton in 1887). PLATE 2,B

Fruits: Length (0.8)—1.4-(2.5) mm.; width (0.4)-0.7-(1.0) mm.;
thickness (0.3)-0.5-(0.7) mm.; semitranslucent; shape suboval or
suboblong; plump; keeled dorsally above the embryo tip; com-
pressed laterally toward the stylar tip; in ventral and dorsal views
lanceolate; embryo three-quarters length of seed; color of endosperm

16 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

grayish yellowish brown or brownish orange; endosperm strongly
reddish brown mottled and usually pitted; scutellum edge narrow
(ca, 0.1 mm. broad), bandlike, at full maturity almost black; radicle-
plumule axis broad, elevated, twisted, and lighter in color than
the scutellum edge; pericarp finely striate.

Florets: Length (1.1)—2.2-(3.0) mm.; width and thickness slightly
more than for the fruits; opaque or semiopaque; chartaceous; gla-
brous; color white, variously greenish or purplish streaked, or purplish;
florets wide toward the center; lemma and palea tips acute (palea
frequently abruptly acute); palea glabrous on the nerves; lemma
scaberulous on the nerve near the tip; palea splits at fruit maturity.

Specimen illustrated: H. L. Bolley 867; Aug. 27, 1891; Fargo,
North Dakota.

Common name: Puffsheath dropseed.

Geographic distribution: Quebec and Maine to Montana, south to
Virginia, Tennessee, and Texas; also Washington and Arizona;
dry open ground and sandy fields.

Habit: Annual.

Economic importance: No value except for erosion control when it
invades abandoned fields or other bare areas.

Occurrence in seed samples: Reported as occurring in native grass
and lezume samples and in S. cryptandrus samples.

9. Sporobolus asper (Michx.) Kunth, Rév. Gram. 1:68. 1829. Based on
Agrostis aspera Michx. PLATE 2,c
Agrostis aspera Michx., F\. Bor. Amer. 1:52. 1803. Illinois, Michaux.

Fruits: Length (1.1)-1.6-(2.0) mm.; width (0.6)-1.0-(1.3) mm;
thickness (0.5)-0.8-(1.1) mm.; semitranslucent; shape suboval or
suboblong; plump; in dorsal view convex the entire length, in ventral
and dorsal views suboblong; embryo three-quarters length of seed;
color of endosperm grayish reddish orange or brownish orange; endo-
sperm reddish brown mottled and usually pitted; scutellum edge broad
(ca. 0.2-0.3 mm.), bandlike, at full maturity almost black; radicle-
plumule axis narrow, elevated, almost straight, and lighter in color
than the scutellum edge; pericarp coarsely striate or reticulate.

Florets: Length (4.0)-6.0-(7.0) mm.; width 1.25 to 2.0 mm.;
thickness ca. the same as the fruits; opaque or semiopaque; mem-
branaceous; glabrous; color pale yellowish gray, light green or purplish
tinged; florets 2 to 3 times as long as the fruits, completely enclosing
them; lemma and palea tips subacute; nerves of the palea glabrous;
the nerve of the lemma scabrous toward the tip; palea splits at fruit
maturity.

Specimen illustrated: H. W. Clark 22; Nov. 22, 1904; east of depot
grounds; Lake Maxinkuckee, Indiana.

Common name: Tall dropseed.

COLBRY-——DIAGNOSTIC CHARACTERISTICS OF SPOROBOLUS 17

Geographic distribution: Vermont to Montana, south to Louisiana,
Arizona and Texas; eastern Washington; prairies and sandy meadows.

Habit: Perennial.

Economic importance: Valuable for forage and erosion control;
fairly palatable and a rapid grower; seldom planted intentionally
but is usually present in the commercial mixed bluestem (Andropogon
spp.) used for seeding purposes.

Occurrence in seed samples: Reported in bluestem (Andropogon
spp.) and other native grasses and legumes, and in lespedeza (Les-
pedeza spp.) from Kentucky.

10. Sporecbolus poiretii (Roem. and Schult.) Hitche., Bartonia 14:32. 1932-
Based on Axonopus potretii Roem. and Schult. PLATE 2,D

Azonopus poiretii Roem. and Schult., Syst. Veg. 2:318. 1817. Based on
Agrostis compressa Poir., “‘n. 78”, not A. compressa Willd., 1790, nor
Poir. (op. cit.) No. 82, on the following page.

Agrostis compressa Poir. in Lam., Encycl. Suppl. 1:258. 1810, not A. com-
pressa Willd., 1790, nor Poir. (op. cit.) 1:259. 1810, nor Sporobolus com-
pressus Kunth, 1933. Carolina, Bose.

Fruits: Length (0.8)-1.1-(1.4) mm.; width (0.5)-0.6-(0.8) mm.;
thickness (0.2)—0.3-(0.4) mm.; translucent or opaque; shape oblong-
ish, tapering to the base; usually strongly laterally compressed, the
edges often reddish black rimmed from the mucilaginous pericarp;
color of the endosperm and radicle-plumule axis reddish or yellowish
orange; embryo less than half the length of the seed; scutellum edge
elevated, about 0.1 mm. wide, and reddish black at maturity; pericarp
minutely striate, the striations often dark reddish; pericarp becoming
mucilaginous at seed maturity, causing the fruits to adhere to the
florets.

Florets: Length 1.5-2.0 mm.; width and thickness ca. the same
as for the fruits; semiopaque; membranaceous; essentially glabrous,
but microscopically sparingly scaberulous; color stramineous or pale
plumbeous; lemma acute; palea obtuse, erose; palea splits at fruit
maturity; fruits with the pericarp partially slipped off often sticking
to the lemma and palea; florets often affected with a black fungus.

Specimen illustrated: A. S. Hitchcock 523; March 1903; Miami,
Florida.

Common names: Smutgrass; rattail smutgrass.

Geographic distribution: Virginia to Tennessee and Oklahoma,
south to Florida, Texas, and the warmer parts of America to Argentina;
on ballast in Oregon and New Jersey; tropical Asia, apparently intro-
duced in America; open ground and waste places.

Habit: Perennial.

Economic importance: None reported in the United States; utilized
for forage in arid regions in southern Europe, according to Fiori (1923).

18 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Occurrence in seed samples: Reported as occurring in Dallis grass
(Paspalum dilatatum) and in bluestem (Andropogon spp.) seed samples.
11. Sporobolus airoides (Torr.) U.S. Rep. Expl. Miss. Pacif. 7:21. 1856. Based

on Agrostis airoides Torr. PLATE 2,E
Agrostis atroides Torr., Ann. Lye. N. Y. 1: 151. 1824. Branches of the
Arkansas River near the Rocky Mountains, James.

Fruits: Length (0.9)—1.1-(1.7) mm. ; width (0.5)—-0.7—(0.9) mm. ; thick-
ness (0.4)-0.5-(0.7) mm.; opaque; shape irregularly suboblong or sub-
oval; plump; the stylar tip rounded and not much laterally compressed ;
color of endosperm and embryo area about the same: a dull, dark,
yellowish brown or light brown; endosperm rarely reddish flecked;
embryo frequently over half the length of the seed; pericarp strongly
dark reddish or blackish striate or reticulate, rarely slipped off
mature fruits.

Florets: Length (1.5)-1.9-(2.5) mm.; width and thickness ca. the
same as for the fruits; opaque or semitransparent; membranaceous;
essentially glabrous but at high magnifications minutely scaberulous;
color plumbeous, stramineous or purplish, often bronze tinged; lemma
and palea acute or subacute, the nerves scaberulous toward the tips;
florets spread apart at maturity, exposing the tips of the fruits; palea
splits at fruit maturity.

Specimen illustrated: Hdgar A. Mearns 2303; Sept. 11, 1893; near
White Water, on Mexican boundary line; New Mexico.

Common name: Alkali sacaton; bunchgrass.

Geographic distribution: South Dakota and Missouri to eastern
Washington, south to Texas and southern California; Mexico;meadows
and valleys, especially in moderately alkaline soil.

Habit: Perennial, in large, tough bunches.

Economic importance: Excellent results from seeding in alkaline
lowlands; valuable forage and erosion control plant.

Occurrence in seed samples: Occasionally part of native grass or
legume seed samples or received as pure seed samples.

12. Sporobolus wrightii Munro ex. Scribn., Bull. Torrey Bot. Club 9:103. 1882.
Pantano, Arizona, Pringle. PLATE 2,F

Fruits: Length (0.9)-1.3-(1.7) mm.; width (0.4)-0.7—(0.9) mm.;
thickness (0.3)—0.5-(0.7) mm.; opaque (occasionally semitranslucent) ;
plump; shape predominantly ovoid, plumper toward the base (rarely
suboval or obovoid); stylar tip usually pointed and strongly laterally
compressed; color of endosperm and embryo area about the same: a
dull, dark reddish brown or occasionally light brown; endosperm
usually faintly reddish flecked; embryo frequently less than half the
length of the seed; pericarp strongly dark reddish or blackish striate
or reticulate, frequently slipped off mature fruits.

Florets: Essentially the same as for S. atroides.

COLBRY-—DIAGNOSTIC CHARACTERISTICS OF SPOROBOLUS 19

Specimen illustrated: Edgar A. Mearns 2345; Sept. 16, 1893; Dog
Spring, Dog Mountains, New Mexico.

Common name: Sacaton.

Geographic distribution: Southern and western Texas and Okla-
homa to southern California and central Mexico; mesas and valleys.

Habit: Perennial, in large dense tufts.

Economic importance: Excellent results from seeding in southern
New Mexico; useful for grazing when young; also furnishes hay and
and makes good winter range.

Occurrence in seed samples: Occasionally part of native grass seed
mixtures; received as pure seed samples in U. 8. Department of
Agriculture Soil Conservation Service seed laboratories.

13. Sporobolus buckleyi Vasey, Bull. Torrey Bot. Club 10:128. 1883. Texas,
Buckley. PLATE 3,4

Fruits: Length (0.7)-0.8-(1.0) mm.; width (0.3)-0.5-(0.7) mm.;
thickness (0.2)-0.3-(0.4) mm.; translucent or opaque; shape suboval
or broadly ovoid; plump and flattish at the base, the embryo without
a prominent overhang; laterally compressed at stylar tip; color of
endosperm and embryo light orange reddish, the embryo sometimes
slightly darker; endosperm faintly light reddish flecked and pitted;
embryo area inconspicuous, usually less than half the length of the seed;
hilum within a basally prominent, black half-rim; pericarp usually
minutely reddish striate.

Florets: Length 1.2 to 1.7 mm. long; width and thickness ca. the
same as for the fruits; opaque; membranaccous; essentially glabrous,
but microscopically slightly scaberulous, especially on the nerves;
color plumbeous, purplish or stramineous; lemma acute, longer than
the palea, which is subacute or obtuse; palea splits at fruit maturity;
fruits with the pericarp partially slipped off often sticking to the
florets.

Specimen illustrated: Jason R. Swallen 1492; Apr. 10, 1931; deep
shade, in Rabb Palm Grove; Brownsville, Texas.

Common name: Buckley dropseed.

Geographic distribution: Texas and eastern Mexico.

Habit: Perennial.

Economic importance: Not reported.

Occurrence in seed samples: Not reported.

14. Sporobolus pulvinatus Swallen, Journ. Washington Acad. Sei. 31:351. f. 4.
1941. Adamana, Arizona, Griffiths 5107. PLATE 3,B

Fruits: Length (0.8)-0.9-(1.0) mm.; (0.4)-0.6-(0.7) mm.; thick-
ness (0.2)-0.4-(0.5) mm.; semitranslucent; shape obovoid; obtuse
at stylar tip and pointed at the base from the overhanging embryo;
ventral surface frequently longitudinally grooved; color of the endo-

20 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

sperm yellowish gray or yellowish orange; endosperm often micro-
scopically pitted; embryo area conspicuous, greenish black at matur-
ity; scutellum in dorsal view panduriform; in lateral view of the fruit
the inner scutellum reflected in a broad, dark semicircle through the
semitranslucent endosperm; radicle-plumule axis broad in relation to
the scutellum edge; pericarp finely striate.

Florets: Length (1.4)-1.5-(1.7) mm.; width and thickness ca. the
same as for the fruits; semiopaque or opaque; membranaceous; essen-
tially glabrous, but microscopically sparingly scaberulous; color
stramineous or plumbeous, sometimes purplish tinged; lemma acute
or subobtuse; palea the same length or slightly shorter than the lemma,
obtuse, slightly dentate; palea splits at fruit maturity; fruits with
the pericarp partially slipped off often sticking to the florets.

Specimen illustrated: A. S. Hitchcock 7820; Oct. 15, 1910; along
street; El Paso, Texas.

Common name: None given.

Geographic distribution: Texas, New Mexico and Arizona; northern
Mexico; sandy land.

Habit: Annual.

Economic importance: Not reported.

Occurrence in seed samples: Not reported.

15. Sporobolus pyramidatus (Lam.) Hitche., U. 8. Dep. Agr. Mise. Pub.
243:84. 1936. Based on Agrostis pyramidata Lam. PLATE 3,¢
Agrostis pyramidata Lam., Tab]. Encyel. 1:161. 1791. South America.

Fruits: Length (0.7)-0.8-(1.0) mm.; width (0.4)-0.5-(0.6) mm.;
thickness (0.2)—0.3-(0.4) mm.; usually translucent; shape obovoid,
suboval, or subelliptic; obtuse at stylar tip and pointed at the base
from the overhanging embryo; ventral surface occasionally longitudi-
nally grooved; color of the endosperm light orange reddish; endo-
sperm often microscopically pitted; embryo area conspicuous, dark
reddish brown at maturity; scutellum in dorsal view panduriform; in
lateral view of the fruit the inner scutellum reflected in a broad, dark
semicircle through the translucent endosperm; pericarp finely striate.

Florets: Length (1.2)-1.5-(1.9) mm.; width and thickness ca. the
same as for the fruits; semiopaque or opaque; membranaceous; essen-
tially glabrous, but microscopically sparingly scaberulous; color
stramineous or plumbeous; lemma acute, usually slightly longer than
the palea; palea subacute or obtuse; palea splits at fruit maturity;
fruits with the pericarp partially slipped off often sticking to the
florets.

Specimen illustrated: A. S. Hitchcock 5343; June 27, 1910; sand,
along railway near bay; Corpus Christi, Texas.

Common name: Whorled dropseed.

Geographic distribution: Kansas and Colorado to Louisiana and

COLBRY—DIAGNOSTIC CHARACTERISTICS OF SPOROBOLUS 21

Texas; southern Florida; tropical America; sandy or gravelly soil,
especially along streets and along the seashore and in the interior in
alkaline soil.

Habit: Perennial, in spreading or prostrate tufts.

Economic importance: Too short to be a productive grass; reported
by Parodi (1928) as good in saline soils of the pampas, Argentina,
where other types of pasture grasses are scarce.

Occurrence in seed samples: Not reported.

16. Sporobolus flexuosus (Thurb.) Rydb., Bull. Torrey Bot. Club 32:601.
1905. Based on Sporobolus cryptandrus var. flecuosus Thurb.
PLATE 3,D

Sporobolus cryptandrus var. flecuosus Thurb. in 8. Wats., Bot. Calif. 2:269
1880. Based on Vilfa cryptandra var. flecuosa Thurb.

Vilfa eryptandra var. flecuosa Thurb. ex. Vasey, in Rothr., in Wheeler U. 8.
Survey W. 100th Merid. Rept. 6: 282. 1878. Nevada and Arizona,
Wheeler Exped.

Fruits: Length (0.7)-0.8-(0.9) mm.; width (0.5)-0.6-(0.8) mm.;
thickness (0.2)—0.4—(0.5) mm. ; opaque, semitranslucent or translucent;
shape asymmetrically broadly ovoid; pointed at the base from the
overhanging embryo; obtuse and scarcely laterally flattened at stylar
tip; in ventral and dorsal views predominantly oblongish, frequently
“pinched” laterally toward the center; color of endosperm light
yellowish brown or pale orange yellow, sometimes almost white;
endosperm usually microscopically pitted and internally strongly
“starchy” or granularly mottled; embryo area conspicuous, rusty or
light reddish brown; radicle-plumule axis narrow, elevated, almost
straight; in lateral view the scutellum edges usually unevenly reflected
through the endosperm; pericarp frequently purplish or rusty striate.

Florets: Length (1.5)—2.1—(2.7) mm.; otherwise like S. cryptandrus.

Specimen illustrated: Starr 171A; Oct. 4, 19385; New Mexico.

Common name: Mesa dropseed.

Geographic distribution: Western Texas to southern Utah, Nevada,
southern California, and northern Mexico; mesas.

Habit: Perennial.

Economic importance: Fair palatability; invader; occurs in lower
Sonoran Zone on medium or light textured soils; at the present
time the only species available for seeding around Las Cruces, Deming,
and Lardsburg, New Mexico, on light-textured soils.

Occurrence in seed samples: Occasional pure seed samples in U. S.
Department of Agriculture Soil Conservation Service seed laboratories.
17. Sporobolus nealleyi Vasey, Bull. Torrey Bot. Club 15:49. 1888, name

only; Contr. U. 8. Nat. Herb. 1:57. 1890. Brazos Santiago, Tex.,
Nealley. PLATE 3,5

Fruits: Length (0.7)-0.8-(1.0) mm.; width (0.3)-0.4-(0.5) mm.;

thickness (0.2)-0.2-(0.3) mm.; opaque or semitranslucent; shape

22 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

narrowly ovoid or narrowly obtuse at stylar tip; pointed at the base
from the overhanging embryo; color of endosperm light yellowish
brown or pale orange yellow, frequently almost white; endosperm
sometimes microscopically pitted and usually internally ‘‘starchy”
or granularly mottled; embryo area conspicuous, rusty or purplish
brown; radicle-plumule axis narrow, elevated, almost straight; in
lateral view the scutellum edges usually unevenly reflected through
the endosperm; pericarp very finely striate, frequently purplish striate,
sometimes rusty striate.

Florets: Length (1.5)—1.7—(2.0) mm. ; florets usually opaque and very
dark green or purplish tinged; otherwise like S. eryptandrus.

Specimen illustrated: J. D. Schoeller & R. S. Campbell 462; Forest
Service 49290; Nov. 13, 1925; three-quarters of a mile east of Middle
Well, in Caliche, Jornada Range Reserve, New Mexico.

Common name: Nealley dropseed.

Geographic distribution: Western Texas, Nevada, New Mexico,
and Arizona; gypsum sands.

Habit: Perennial, with rhizomes.

Economic importance: Quite common around Roswell, Carlsbad,
and Partalu, New Mexico; thought to have forage value.

Occurrence in seed samples: Not reported.

18. Sporobolus contractus Hitchc., Amer. Journ. Bot. 2:303. 1915. Based
on Sporobolus strictus Merr. PLATE 3,F
Sporobolus strictus Merr., U. 8. Dep. Agr., Div. Agrost. Cir. 32:6. 1901,
not S. strictus Franch., 1893. Based on Sporobolus eryptandrus var.
strictus Seribn.
Sporobolus cryptandrus var. strictus Scribn., Bull. Torrey Bot. Club 9:108.
1882. Camp Lowell, Arizona, Pringle.

Fruits: Length (0.8)-1.0-(1.2) mm.; width (0.4)-0.6-(0.8) mm.;
thickness (0.2)—0.3-(0.5) mm.; opaque, semitranslucent or translu-
cent; shape broadly ovoid or broadly elliptic; pointed at the base from
the overhanging embryo; strongly laterally flattened; obtuse or pointed
at stylar tip; color of endosperm light yellowish brown or pale orange
yellow, sometimes almost white; endosperm usually microscopically
pitted and internally strongly “starchy” or granularly mottled;
embryo area conspicuous, rusty or light reddish brown; embryo area
usually less than half the length of seed; radicle-plumule axis narrow,
elevated, almost straight; in lateral view the scutellum edges usually
unevenly reflected through the endosperm; pericarp frequently rusty
striate, sometimes greenish striate.

Florets: Length (1.5)-2.4-(2.9) mm.; specimens with stramineous
or pale plumbeous florets exceeding those with darker florets; other-
wise like S. cryptandrus.

Specimen illustrated: O. B. Metcalfe 780; Sept. 22, 1903; Gila
River bottom near cliff; Grant County, New Mexico.

COLBRY—DIAGNOSTIC CHARACTERISTICS OF SPOROBOLUS 23

Common name: Spike dropseed.

Geographic distribution: Arkansas, Colorado to Nevada, south to
western Texas, southeastern California, and Sonora; adventive in
Maine; mesas, dry bluffs, and sandy fields,

Habit: Perennial.

Economic importance: Not used much in reseeding; fits in where S.
cryptandrus and S. fexuosus overlap.

Occurrence in seed samples: Occasionally in native grass seed
mixtures or in U. 8. Department of Agriculture Soil Conservation
Service seed laboratories, received as pure seed samples.

19. Sporobolus giganteus Nash, Bull. Torrey Bot. Club 25:88. 1898. Dona
Ana County, N. Mex., Wooton 394. PLATE 4,4

Fruits: Length (1.0)-1.3-(1.9) mm.; width (0.6)-0.8-(0.9) mm.;
thickness (0.3)-0.4-(0.5) mm.; otherwise essentially the same as for
S. contractus.

Florets: Length (2.2)-2.9-(3.5) mm.; otherwise essentially the same
as for S. contractus.

Specimen illustrated: Paul C. Standley; Oct. 13, 1906; sand hills,
southwest of Mesilla, New Mexico.

Common name: Giant dropseed.

Geographic distribution: Oklahoma and western Texas to Colorado
and Arizona; mesas and sandhills.

Habit: Perennial.

Economic importance: Minor forage and erosion control value;
used on sand dunes in New Mexico with fair results; palatable plant
on dunes and blow-out areas in Oklahoma.

Occurrence in seed samples: Occasionally in native grass seed
mixtures or in U. S. Department of Agriculture Soil Conservation
Service seed laboratories as pure seed samples.

20. Sporobolus cryptandrus (Torr.) A. Gray, Man. 576. 1848. Based on
Vilfa cryptandra Torr. PLATE 4,B
Vilfa cryptandra Torr. ex. Trin., Mém. Acad. St. Pétersb. VI. Sci. Nat.
41:69. 1840. Based on Agrostis cryptandra Torr.
Agrostis cryptandra Torr., Ann. Lye. N. Y. 1: 151. 1824. Canadian River
(Texas or Oklahoma), James.

Fruits: Length (0.6)-0.8-(1.3) mm.; width (0.3)-0.5-(0.8) mm.;
thickness (0.2)-0.3-(0.5) mm.; translucent, semitranslucent, or
opaque; shape ovoid or obovoid (rarely suboval); pointed at the base
from the overhanging embryo; laterally compressed toward the stylar
tip; bulging toward the base; in ventral and dorsal views predominantly
lance-shaped; color of endosperm moderate orange, brownish orange
or light reddish brown; endosperm faintly light reddish or rusty
flecked and usually pitted; endosperm internally clear or “‘starchy”
or granularly mottled; embryo area usually inconspicuous, the same

24 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

color or a slighly deeper shade than the endosperm (rarely very much
darker); radicle-plumule axis narrow, elevated, almost straight; in
lateral view the scutellum edges evenly or unevenly reflected through
the endosperm; pericarp very finely striate, sometimes faintly rusty
or reddish striate (rarely with purplish striations).

Florets: Length (1.4)-1.8-(2.0) mm.; width and thickness ca. the
same as for the fruits; opaque, transparent or semitransparent;
membranaceous; glabrous to minutely microscopically scaberulous,
especially toward the tips of the florets and the nerves; color variable,
from plumbeous, frequently purple tinged, to stramineous; lemma
and palea acute or acuminate; palea splits at fruit maturity.

Specimen illustrated: Hula Whitehouse 17190; Sept. 25, 1946; sandy
prairie; 4 miles southeast of Farwell on highway 84; Parmer County,
Texas.

Common name: Sand dropseed.

Geographic distribution: Maine and Ontario to Alberta and Wash-
ington, south to North Carolina, Indiana, Louisiana, southern Cali-
fornia, and northern Mexico; sandy, open ground.

Habit: Perennial, usually in rather small tufts.

Economic importance: An invading species in many types of soils;
more valuable for erosion control and seeding on problem spots than
for forage; not as palatable as most other range grasses.

Occurrence in seed samples: Reported in native grass and legume
seed samples or received as pure seed samples.

Literature cited

Bews, J. W. The world’s grasses. 66. 1929,
Brower, W. & Stahlin, A. Handbuch der Samenkinde. 1955.
Fernald, M. L. Two segregates in Sporobolus. Rhodora 35: 108-110. 1933.
Gray, A. Ann. Lye. N. Y. 3:233. 1835.
Hitchcock, A. 8. Manual of the grasses of the United States, ed. 2, revised by
Agnes Chase. U.S. Dep. Agr. Misc. Pub. 200, pp. 413-432. 1950.
Jones, E. K., & Fassett, N. C. Subspecific variation in Sporobolus cryptandrus,
Rhodora 52:125-126. 1950.
Lawrence, G. H. M. Taxonomy of vascular plants. 389. 1951.
Munsell Pocket Edition Constant Hue Charts. 1929.
Murley, M. L. Seeds of the Cruciferae of northeastern North America. Amer.
Midl. Nat. 46:1-81. 1951.
Parodi, L. R. Revision de las gramineas argentinas del genero Sporobolus.
Rev. Fac. Agron. y Vet. Univ. Buenos Aires, 6:122. 1928.
Fiori Véase A. Nuova flora analitica d’Italia. 1:95. 1923.
U.S. Department of Agriculture. Testing agricultural and vegetable seeds. U.S.
Dep. Agr. Handbook No. 30, pls. 12, figs. 196-198, and 34, pp. 194-261.
1950.

UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED STATES NATIONAL HERBARIUM

VoLumE 34, Part 2

THE WOODS AND FLORA OF THE FLORIDA KEYS:
CAPPARACEAE

By Wixi L. Stern, Georce K. Brizicky,

AND Francisco N. TAMOLANG

BULLETIN OF THE UNITED STATES NATIONAL MuSEUM

SMITHSONIAN INSTITUTION e WASHINGTON, D.C. ¢ 1963

THE WOODS AND FLORA OF THE FLORIDA KEYS:
CAPPARACEAE

By Wittram L. Stern,’ Georce K. Brizicky, AND
Francisco N. TAMOLANG 2

Introduction

Capparaceae * A. L. de Jussieu, are primarily a tropical and sub-
tropical family of xerophilous and heliophilous trees, shrubs, vines,
and herbs characterized by acrid juice. The family comprises about
45 genera with approximately 700 species which are widely distributed
in the New and Old Worlds. Almost half the genera are monotypic.
Representatives of two pantropical genera occur on the Florida Keys:
the native Capparis L., and Cleome L., which is probably naturalized.
The present study deals only with the woody Capparis. (An excellent
discussion of the taxonomic status of Capparaceae, ecology, mor-
phology, pollination mechanisms, and distribution, is given in Jacobs,
1960.)

Capparis is a pantropical genus of about 250 species, best repre-
sented in America and Africa. Pax and Hoffmann (1936) subdivided
Capparis into 14 sections. These appear to be fairly natural, although
they are based almost exclusively on exomorphic, and perhaps not
always unquestionable, characters. Chromosome numbers are known
for a few species (2n=18, 30, 38, 40, 84; Darlington and Wylie, 1956)
and can hardly be of any assistance at the present time. The genus
as a whole appears to be insufficiently known; more comprehensive
investigations based on adequate material, and a revision would be
desirable. The two species of Capparis which occur on the Florida
Keys probably represent two sections of the genus, as noted below.

Capparaceae, usually placed in the order Rhoeadales near Papa-
veraceae and Fumariaceae, are presently assumed to be most closely
“1 Previous numbers in this series appeared in the journal Tropical Woods (Yale University, School of
Forestry) under the running title, ‘“The Woods and Flora of the Florida Keys”; “Introduction” by W. L.
Stern and G. K. Brizicky, vol. 107, pp. 36-65, 1957; ‘‘Compositae” by 8, Carlquist, vol. 109, pp. 1-37, 1958;
“Goodeniaceac” by W. L. Stern and G. K. Brizicky, vol. 109, pp. 38-44, 1958; ‘“‘Passifloraceae’’ by W. L.

Stern and G. K. Brizicky, vol. 109, pp. 45-53, 1958; and “‘Wood Anatomy and Phylogeny of Batidaceae” by
J. McLaughlin, vol. 110, pp. 1-15, 1959.

2 Department of Botany, Smithsonian Institution, Washington, D.C.; Gray Herbarium, Harvard Uni-
versity, Cambridge, Massachusetts; and Forest Products Research Institute, University of the Philippines,
College, Laguna, Philippines, respectively.

3 This taxon is more widely known as Capparidaceae, However, the authors have chosen to follow the
recommendations in Appendix 1, ‘Nomina familiarum conservanda,”’ of the 1961 edition of the International
code of botanical nomenclature, where the approved name is Capparaceae.

25

26 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

related to Cruciferae, with affinities to Moringaceae, Resedaceae, and
Tovariaceae. Hutchinson’s (1926) order Capparales (“Cappari-
dales”) has been considered by Takhtajan (1959) to comprise all the
above-mentioned families and Koeberliniaceae. Papaveraceae and
Fumariaceae are excluded. Data from the biochemistry of these
families (Hegnauer, 1961) seem to be in favor of the Takhtajan
delimitation of Capparales. However, conclusions based on these
data should probably be considered preliminary, since the material
investigated is inadequate.

This study is based in large part on the wood specimens listed in
table 1; these are on deposit for reference in several institutions
including the division of plant anatomy of the U.S. National Museum.
Associated herbarium vouchers from the U.S, National Herbarium,
and other institutions, have also been examined where necessary.
Microscope slides of wood prepared expressly for this research are on
deposit in the division of plant anatomy.

Capparis cynophallophora L. Sp. Pl. 504. 1753.
Capparis jamaicensis Jacq. Enum. Pl. Carib. 28. 1760.

The Jamaica caper, Capparis cynophallophora, is a shrub or small
tree with lepidote branchlets. Leaves are elliptical, oblong-elliptical
or narrowly elliptical, rounded or notched at apex, rounded at base,
not conspicuously reticulate-veined, glabrous on the upper surface,
rusty or silvery lepidote with peltate trichomes on the lower surface,
and exstipulate. Flower buds are 4-angled. The opened flowers
are about 3 cm. in diameter. Sepals are more or less equal (appearing
I-seriate), free, lanceolate to ovate-lanceolate, acute, and valvate in
aestivation. The receptacular nectariferous glands are liguliform.
Staminal filaments are purplish. The one-locular ovary develops a
torulose fruit-body which is lepidote, and 20-30 em. long. The
seeds are more or less reniform, and the embryo is coiled (cochleate).

Disrrisutrion.—C. cynophallophora occurs in exposed areas in the
coastal hammocks on most of the Florida Keys. It can be found in
similar habitats north to Pine Key, Boca Ciega Bay (Pinellas Co.)
and Cape Canaveral (Brevard Co.) on peninsular Florida. The
Jamaica caper probably occurs on all the West Indian Islands,
southern Mexico, Central America, and northern South America.
In Panama, specimens from both coasts have been seen.

Capparis flexuosa (L.) L. Sp. Pl. Ed. 2. 722. 1762.
Capparis cynophallophora L. Syst. Nat. Ed. 10. 2:1071. 1759.
Morisonia fleruosa L. Pl. Jamaic. Pug. 14. 1759; Amoen. Acad. 5:398. 1760.
The limber caper, Capparis flecuosa, is a glabrous shrub or small
tree with leaves of similar shape to C. cynophallophora (fig. 1). How-
ever, in this species, they are conspicuously reticulate-veined and

WOODS OF FLORIDA KEYS—STERN AND OTHERS 27

glabrous, minutely stipulate, and each bears an oblong, axillary gland.
Subglobular flower buds open into blooms about 5-7 cm. in diameter.
The sepals are unequal (distinctly 2-seriate), the exterior ones smaller
than the interior, somewhat connate at the base, subcircular, and
distinctly imbricate in aestivation. Receptacular nectariferous glands
are broad and low appearing as glandular spots or pits. The staminal
filaments are white. The ovary is 2-locular in the material investi-
gated by W. R. Ernst.t Fruit-body (fig. 1) is continuous or some-
what torulose, glabrous, 10-20 cm. long; seeds are irregularly shaped,
obliquely ellipsoidal, ovoid or obovoid, and slightly angular; the
embryo is straight, and the cotyledons infold one another and cover
the radicle with the lobes of their obliquely cordate bases.

DistripuTion.—The range and habitat of C. flexuosa resemble
those of the preceding, but this species appears to be more
widespread. Specimens have been seen from as far north as Merritt
Island (Brevard Co.) in Florida, from Mexican states on both coasts
(north to Sinaloa on the west, and Tamaulipas on the east), from the
West Indies, and from both coasts of Central America. The limber
caper has been collected in Colombia, Ecuador, Venezuela, Paraguay,
Trinidad and Tobago, and Brazil (Santa Catarina) in South America.
Macbride (1938) thinks it probable that this species also occurs in
Peru.

The nomenclatural confusion in which Capparis cynophallophora
L. was usually known as C. jamaicensis Jacq., and C. flecuosa L. was
known as (C. cynophallophora 1.., was cleared up by Fawcett (1914).
Although (C. eynophallophora L. was placed into the section Cyno-
phalla by Pax and Hoffmann (1936), the morphological characters °
of this species clearly show that it belongs to the section Quadrella
where De Candolle (1824) had already placed it. Radlkofer (1884,
1887) confirmed the place of C. cynophallophora (“C. jamaicensis’’)
in this section, the original description of which he emended by adding
some anatomical characters of the leaves.

Xylem Anatomy

The transverse section of Capparis cynophallophora L. exhibits
alternating concentric bands of cells which make up the groundmass
of the wood. Cells in these bands are nonseptate fiber-tracheids.
Bands with cells having conspicuously wider lumina and somewhat
thinner walls alternate with bands comprising narrow-lumened fiber-
tracheids with thicker walls. These bands vary considerably in
width, and adjacent bands commonly merge to form irregular arcuate

4Dr. Ernst has examined the Florida Keys capers in preparation for publishing “The genera of
Capparaceae and Moringacceae in the southeastern United States” (1963).

§Four-angled flower buds, lepidote pubescence, equal and valvate sepals, and the presence of scler-
enchymatous idioblasts (‘‘spiculares’’) in the mesophyll of leaves.

28 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

segments. That these are bands associated with growth is evident
from ther formation and arrangement, but whether they are re-
flective of climatic conditions is questionable.

Fiber-tracheids have obscurely bordered pits which average 1.6
in diameter. The thickness of radial cell walls varies from thin to
thick, and ranges from 0.62—4.20 » with an average of 2.07 u. The
range in length of fiber-tracheids is from 292-661 » with an average of
487 uw. Most cells fall between 415 and 523 yw in length.

The wood is diffuse-porous and pores are moderately small ranging
from 8-107 » in tangential diameter, averaging 62 4 wide. Most are
between 33 and 83 u in diameter. In transverse outline pores are
circular to oval. There are normally 20 pores per square millimeter
of which 24 percent are solitary. Mostly, however, pores are clustered
and in radial multiples of 2-4 (5). Aggregations of pores present a
complex picture appearing usually as somewhat irregular clusters of
cells with one or two large central pores surrounded by several smaller,
more or less wide-lumened cells. These aggregations, which shall be
designated ‘‘pore complexes,’ consist of several cell types and modi-
fications: ‘‘normal’’ vessels with regular elements, extremely narrow
vessels, and axial parenchyma cells. Vessel element walls vary from
thin to thick, from 1.7-5.0 », and average 3.8 u. Mostly they range
between 3.3 and 8.3 win thickness. Perforation plates are simple in
even the very narrowest vessel elements, and are circular as viewed
in the radial section. Jind walls of vessel elements (fig. 6) are mostly
oblique and range from 0°-55° from the horizontal. Intervascular
pitting is alternate, and the pits are crowded which sometimes
produces angular pit borders. Pits are tiny, ranging from 2-3 uw in
diameter, and appear to be obscurely vestured (cf. Bailey, 1933).
However, this could not be determined with assurance on the sections
available. Vessel elements average 192 » in length and range from
94-296 u. Most vessel elements are between 139 and 235 u in length.
Vessel—axial parenchyma pitting resembles the intervascular pitting
in that it is alternate, but pits may be oblong in shape as well as
circular.

Vascular rays (figs. 6, 7) as viewed on the tangential section are
numerous and average 12 per mm. Most rays are uniseriate, but it
is also common to see essentially uniseriate rays with short, bi- and
triseriate central segments. Other rays are mostly bi- and triseriate
throughout. Uniseriate rays are low and average 20 cells high
(0.34 mm.). They vary from 4-30 cells (0.29-0.46 mm.) in height.
The multiseriate rays are correspondingly low being roughly of the
same dimensions as the uniseriate rays. Rays are essentially homo-
cellular; that is, composed completely of procumbent cells (Kribs’
homogeneous type I). Pitting between vascular ray cells and cells

WOODS OF FLORIDA KEYS—STERN AND OTHERS 29

of the axial parenchyma is alternate. Crystals occur more or less
commonly in ray cells and fall within Chattaway’s (1955) category 6,
‘“thomboidal, square or diamond-shaped.”

Axial parenchyma is vasicentric consisting of a uniseriate sheath
around pores or pore complexes, and sometimes it is also mixed among
the cells in the complex. Crystals were not noted in axial parenchyma
cells.

In Capparis flecuosa (L.) L., the banded nature of the imperforate
tracheary elements of the groundmass (figs. 2, 3) as seen on the
transverse section is not as sharply defined here as in the C. cynophal-
lophora specimen described in the foregoing. The areas of narrow
and wide-lumened cells are highly irregular, and the distinction
between the areas is not sharp.

Imperforate tracheary elements of the groundmass are nonseptate
fiber-tracheids characterized by small, minutely bordered pits (average
1.8 » in diameter). Radial thickness of cell walls varies from very
thin to thick, and ranges from 0.62-6.14 » with an average of 2.25 uy.
Fiber-tracheids range from 292-584 yw in length, averaging 418 yg.
The most frequent range is between 385 and 492 u.

The wood is diffuse-porous, the pores are circular to oval, and are
moderately small varying from 21-96 yw with an average of 56 y» in
tangential diameter (figs. 2, 3). Most are between 24 and 90 » in
width. There are about 35 pores per square millimeter of which 6
percent are solitary and the remainder in radial groups of 2-10
(mostly 2-5) with many uregular clusters. Pore complexes (figs. 2, 3)
consist of a few large pores surrounded by other cells as noted above
under the description of C. cynophallophora. Walls of vessel elements
range from thin to thick, from 1.5-7.5 uw, and average 5.1 4. Perfora-
tion plates are simple in all vessel elements and circular to oval.
Vessel elements have oblique end walls which vary from 0°-70° from
the horizontal. Intervascular pitting is alternate and crowded; the
pits seem to be vestured and are very small (2-4 yu, average 3.2 yu)
with circular to somewhat angular borders. Vessel elements range
in length from 63-261 w and average 175 uw. The most frequent range
is between 156 and 210 uw. Vessel—axial parenchyma pitting is
alternate and pits usually have circular borders.

Vascular rays as seen on the tangential section are numerous and
average 10 per mm. Most rays are bi- and triseriate, uniseriate rays
being of less frequent occurrence. Both uniseriate and multiseriate
rays are low, the former ranging from 3-26 cells high (0.06—-0.26 mm.),
the latter ranging from 5-60 cells high (0.09-1.1 mm.). Rays are
homocellular composed essentially of procumbent cells (Kribs’
homogeneous type I). Multiseriate rays commonly have several-
celled uniseriate wings. Pitting between ray and axial parenchyma

30 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

cells is alternate. Ray cells often contain crystals designated by
Chattaway (1955) as “rhomboidal, square or diamond-shaped.”

Axial parenchyma (figs. 2, 3) consists of a vasicentric sheath sur-
rounding pores or pore groups. Strands may also occur among the
vessels in the group. No crystals were seen in the cells of axial
parenchyma.

Anatomical distinctions between the two species of Capparis
collected from the Florida Keys, as described here, are slight. Minor
variations in cell size occur, especially in the extreme ranges, but these
are of little or no consequence considering the great coincidence of
average figures and those for most frequent range. Most noticeable
of the differences between the two species is the width of vascular
rays. In C. cynophallophora, rays are largely uniseriate (figs. 6, 7),
whereas in C. flecuosa bi- and triseriate rays predominate. Regard-
less, rays with bi- and triseriate segments occur in the former, and
uniseriate rays appear in the latter. We may expect that this dis-
tinction then, is one of degree rather than the two conditions being
mutually exclusive. A statistical analysis of many specimens of each
species would doubtlessly determine the taxonomic validity of this
potential anatomical distinction,

Analyses of Cells and Tissues

The descriptions above would seem to belie the complex nature of
the wood, and especially the difficulties in interpretation of certain of
the cellular components. In order best to interpret and evaluate the
constituent elements of the wood in the Florida Keys capers, and for
purposes of comparison, prepared slides of all species of Capparis were
borrowed from the Samuel James Record Memorial Collection at
Yale University; wood specimens of Capparis from the National Col-
lections of wood were examined; and fluid-preserved specimens of the
Keys species of Capparis were specially gathered for this study. All
wood samples employed in these studies were associated with cor-
responding herbarium vouchers except as noted in table 1.6 Further-
more, wherever possible vouchers were examined to substantiate the
authenticity of identification; in some doubtful cases twigs from
vouchers were examined microscopically and compared with the
wood samples to assure concurrence.

6 The authors are deeply indebted to Dr. Graeme Berlyn and Mr. Richard A. Mark of Yale University
for the privilege of examining Capparis slides and some herbarium vouchers from the Record Memorial
Collection, and to Mr. John R. Millar of the Chicago Natural History Museum for allowing us to see and
compare herbarium vouchers of Capparis with wood specimens used for comparative studies. Dr. James
A. Duke of the Missouri Botanica] Garden aided in verifying the identification of some herbarium vouchers
on deposit in St. Louis and for sending us bits of twigs for comparison. For his help in preparing slides, and
for other kindnesses, the authors are grateful to Mr. Edward S, Ayensu, research assistant at the Smith-
sonian Institution. Particular gratitude is due Mr. George Stevenson of Plantation Key, Florida, and Dr.

P, B. Tomlinson of the Fairchild Tropical Garden in Miami, Florida, for their most kind assistance in col-
lecting fluid-preserved stems of C. flexuosa and C. cynophallophora together with herbarium vouchers.

WOODS OF FLORIDA KEYS—STERN AND OTHERS 31

GrounpMass.—After a preliminary study of the material on hand,
we found that a convincing interpretation of the cells and tissues of
the wood of Capparis would require fluid-preserved specimens. But
nowhere was this need more strongly felt than in the correct identifica-
tion of the cells comprising the tissues surrounding the pore complexes
to be described below. Bands are more or less obvious on the trans-
verse section of all woods examined (figs. 2, 3), except for C. nitida
(fig. 4) and C. schunkei. At first it seems obvious that these bands are
merely rings of growth demarcated by alternating layers of thick- and
thin-walled imperforate tracheary elements. But, in C. pulcherrima
and C. angulata (figs. 10, 11), it seems equally apparent that at least
some of the bands are truly and purely parenchymatous. In certain
species, e.g., C. decidua, a confounding of bands occurs; some seem to
be totally prosenchymatous and others at least partly parenchymatous.
One is tempted here to call this arrangement boundary parenchyma
after the definition of Jane (1956). In C. nitida (fig. 4) and a few
other species pore complexes are much simplified and seem to consist
of a solitary pore, a short radial multiple, or a small cluster surrounded
by a several- to many-seriate sheath of vasicentric parenchyma. In
the slides prepared from previously dried wood, certain groundmass
cells which look like imperforate tracheary elements (fibers) take on
some of the characteristics of parenchyma (compare fig. 2 with fig. 3),
i.e., cytoplasmic remains are present within the lumina. This was
indicated by the strong affinity for haematoxylin shown by the cell
contents. Walls are thinner in these cells than in adjacent prosenchym-
atous elements, and the compound middle lamella seems thicker and
possesses a great affinity for haematoxylin and little for safranin.
The foregoing observations raised questions in the minds of the authors
which seemed solvable only after the examination of fluid-preserved
rather than dried tissues.

For purposes of cytological examination, the fluid-preserved
collections of Stevenson (Newcomer’s fixative) and Tomlinson
(F. A. A.) were utilized. In all, five specimens were available: two
of C. flexuosa, two of C. cynophallophora, and a single, unnamed speci-
men from the Fairchild Tropical Garden. Our observations indicate
that in the specimens of Capparis investigated a well-formed fusiform
nucleus is present in every imperforate tracheary element, from bark
to the inner edge of the radial section (i.e., for a distance of 6-10 mm.),
in all but the Stevenson collection of C. cynophallophora (fig. 13).
Here, the innermost elements lack a nucleus. Furthermore, starch
erains occlude the cell cavity completely, except for the narrowed tips,
of practically all imperforate tracheary elements in the specimens of
C. flexuosa (fig. 13). In C. cynophallophora starch occurs mostly in the
thick-walled imperforate tracheary elements at the margins of “growth

687-311—63——2

32 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

rings,” but not in every cell. As in C. flecuosa, starch grains totally
occlude the cell cavity. In the unnamed Tomlinson specimen starch
occurs in bands corresponding to the alternating layers of thick- and
thin-walled imperforate tracheary elements—it is present only in the
thick-walled cells.

We may assume from the above that the imperforate tracheary
elements of the groundmass in the five Capparis specimens under
analysis, partake of the nature of parenchymatous cells. It is probable
that they continue to perform vital as well as mechanical functions
long after formation by the vascular cambium. In this connection it
is interesting to consider Carlquist’s (1961) remarks, which point out
the possibility of axial parenchyma cells arising from libriform wood
fibers through the mechanism of “fiber dimorphism.”

These findings contrast stronely with the widely held characteriza-
tion of imperforate tracheary elements as being dead at ontogenetic
maturity. ‘The living nature of these cells is indicated by the presence
of a presumably functioning protoplast which, among other activities,
stores starch. The cell wall pitting in these elements, however,
consists of tiny vestigially bordered or simple pit-pairs, whereas in
vasicentric parenchyma cells in the same specimen the pit-pairs are
as large as intervascular pits, simple, half-bordered, or even fully
bordered.

Fahn and Leshem (1963), in a study of xylem anatomy of woody
desert plants from Israel and other woody plants native to Switzer-
land, reported living fibers in 70 percent of 60 species in 24 plant
families. Furthermore, they state that fibers over the entire sap-
wood contained living protoplasts. Included in their study was
Capparis spinosa Li. which showed living fibers in the xylem. Esau
(1953) mentions that “In contrast to other wood fibers, the septate
forms long retain their protoplasts and serve for storage of starch,
oils... .’’ Cumbie (1960) also mentions the presence of nucleated
fibers in certain transitional herbaceous—woody legumes, and Carlquist
has remarked on the presence of these in Compositae in correspondence
with the authors. Harrar (1946) examined the wood of over two
thousand species of plants from the Yale collections and recorded
the presence of starch in the septate fiber-tracheids of many hundreds
of these. It is reasonable to assume that these starch-containing
cells, especially in sapwood, are living, and Harrar also believes
they may function in a manner similar to axial parenchyma.

It is likely that further cytological studies based on fluid-preserved
material of the stem will force us to revise, or at least to broaden,
our currently held notions as to the physiological condition of the
imperforate tracheary element. Our observations here, and those of
Fahn, indicate relatively long-lived protoplasts in imperforate trache-

WOODS OF FLORIDA KEYS—STERN AND OTHERS 33

ary elements; furthermore, it is not only the septate fiber-tracheids
which retain their vital functions for more or less long periods after
formation; the phenomenon may also occur in nonseptate elements.
The universality of this phenomenon still remains to be tested.

Interesting physiological questions concern the ultimate fate of
this stored starch. If it is dissolved and subsequently transported,
must a still-living cell be postulated? How does the dissolved starch
pass from the storage cells to where it is ‘‘needed”’ elsewhere in the
plant? The tiny pits in libriform fibers and fiber-tracheids would
certainly present a significant barrier to free passage. Even then,
through what cells would the dissolved starch be conducted?

These observations on fluid-preserved wood, coupled with those on
previously dried wood, helped us considerably to interpret with a
degree of assurance, the nature of the groundmass cells in Capparis.
By and large, the cells of the groundmass are imperforate tracheary
elements, either libriform wood fibers or fiber-tracheids.

AXIAL PARENCHYMA.—The growth rings are not to any extent
dependent for distinction upon the occurrence of boundary paren-
chyma, and are visible because of alternating layers of thick- and
thin-walled fibers (figs. 2, 3). The situation in C. decidua, where
axial parenchyma cells are involved in growth-ring margins, is of
sporadic occurrence on the transverse section, and is dependent on
the presence of closely adjacent, tangential rows of pore complexes
next to the growth-ring margin. The seemingly apotracheal paren-
chyma cells are in reality vasicentric parenchyma cells mixed among
the cells of the groundmass at the growth-ring boundary.

All species have narrow sheaths of vasicentric parenchyma sur-
rounding the pore complexes (figs. 2-4, 10, 11). Cells adjacent to
these narrow sheaths which appear parenchymatous, as in C. nitida
(fig. 4), are merely thin-walled, wide-lumened, imperforate tracheary
elements as described above. In two species, C. pulcherrima and
C. angulata (figs. 10, 11), true banded axial parenchyma occurs, as
well as the vasicentric, and sometimes aliform and confluent configura-
tions. These bands are composed of several uniform cell layers which
contrast strikingly with the parenchymatous-appearing, thin-walled,
prosenchymatous cells that mark the boundaries of growth rings in
other species.

PorE COMPLEXES.—AIl species of Capparis investigated were char-
acterized by the presence of more or less complicated groupings of
pores and associated cells as observed on the transverse section (figs.
2, 3, 10, 11). These pore complexes consist of a few large vessels,
a variable number of small vessels, sometimes imperforate tracheary
elements, and always associated axial parenchyma. As indicated
previously the latter is always paratracheal, forming a complete

34 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

sheath around the pore complex, and sometimes strands are inter-
mixed among other elements within the complex. Parenchyma cells
are relatively thick-walled and lignified. The tiny vessel elements
which occur in association with the large vessel elements are inter-
esting in that they always possess simple perforations (figs. 2, 3, 11,
12). They may be narrower than adjacent imperforate tracheary
elements; sometimes they are imperfect and contain only a single
perforation; and occasionally perforations appear on the axial walls.

To interpret these complexes solely on the transverse section is
impossible. It is hardly possible to distinguish tiny pores from trans-
verse sections of associated imperforate tracheary elements (figs. 2, 3,
10, 11). Without doubt then, our analyses of pore groupings are in
error—an error difficult to overcome, for if there is a single feature
of the wood which might be said to characterize Capparis, it is the
pore complex described above.

VASCULAR RAYS.—Vascular rays in Capparis are essentially homo-
cellular and are composed of procumbent cells (figs. 5-9). However,
in a number of species—C. decidua, crotonantha, linearis, oxysepala,
quind, schunket, tenuisiliqua, verrucosa, and angulata—rows of squar-
ish marginal cells are of sporadic occurrence, and it is possible that
this phenomenon occurs in all species of Capparis to a certain extent.
In some of the species examined other peculiarities are present to a
more limited extent. Thus, inflated ray cells which resemble secre-
tory cells of some lauraceous woods occur in C. ferruginea, nobilis,
quina, and tenuistiiqua; and sheath cells are of infrequent occurrence
in the rays of C. nobilis, schunket, verrucosa, and angulata. Seriation
of rays varies from 1-7 cells wide. Height of uniseriate rays is from
1-22 cells, of the multiseriate rays, from 3-55 cells (figs. 6-9).

OTHER FEATURES.—Storied structure occurs in only one of the
species studied, C. angulata (fig. 9). The cells which comprise bands
of parenchyma, and the vessel elements, are in marked seriation as
viewed on the tangential section. Cozzo (1944, 1946) and Cozzo and
Cristiani (1950) record partial storying in C. salicifolia Gris. and C.
tweediana Eichl. Rhomboidal crystals were observed in most of the
species under investigation, and it is probable that all species of
Capparis possess the genetic potential to form crystals under con-
ducive environmental circumstances. Most commonly, crystals were
found in the ray cells, but sometimes they were also observed in the
cells of the axial parenchyma.

Phylogenetic Specialization in the Wood

The structure of the wood of Capparis is indicative of a high level
of evolutionary specialization, as shown by the presence of simple
perforation plates in vessels and alternate intervascular pitting (figs.

CONTR. U.S. NAT. HERBARIUM, VOL. 34 PART 2, PLATE 1

f f
@

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= -

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hic. lL. —-Capparts flexuosa (L.) L.. Crawl Key, Florida; upper portion of shrub showing
mature fruits, < l/4 (FL. Stern 1579, US).

CONTR. U.S. NAT. HERBARIUM, VOL. 34 PART ?, PLATE 2

4 )
d . {
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Lunisertate sheath of vasicentric xia) parenchyma whichis discernible tn this material

nal
prepiired from flutd- preserved tissues, 160) (G. Stevenson oon LSw 30300). Pro. 3
Co fexwood CL) Lo. transverse seetion for comparison with feure 2 Notice dhatoon thas

seclion, prepared from dried tisstles. itis not possible to distineuish the uniseriate shes

axial parenelyina from the surrounding tnperforatre cells, IO Cf. Ge fark good,

CONTR. U.S. NAT. HERBARIUM, VOL. 34 PART 2. PLATE 3

A
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sheath of uniseriate

pores, = 160 (LL Wi

vasicentric axial parenchyma, and uncomplicated
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CONTR. U.S. NAT. HERBARIUM, VOL. 34 PART 2, PLATE 4

Mig. 6. Capparts eyvnophallophora 1... tangential seetion of wood, 200 (6. Sterensan

ay
USw 30299). Bie. 7. Co evnophallophora L., tancential section of Wood. 200 (PB.
Pomlinson POEL O24). Vascular rays in these sections are illustrative of Group Trvpe:
mostly unisertare with occasional biseriates beariny uniseriate extensions

PART 2, PLATE 5

34

NAT. HERBARIUM, VOL

CONTR US

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HERBARIUM, VOL. 34

PART 2. PLATE 7

iG. 12. Capparty tenwniliqgua Jacg.. radial section of wood io
tiny vessel clement Ceenter) associ

ISO0 (S.J. Record SU

Wor id Lo den myst re

show Silay)
é ated in complex with vessel of nornial dias
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from vascular cambiumy: starch vrains arc alse vistble in the dumina otf the the
L379 (6. Steceiven vn L Sw 30300)

WOODS OF FLORIDA KEYS—STERN AND OTHERS 35

5, 9, 12), the highly peculiar pore complexes (figs. 2, 3, 10, 11), para-
tracheal parenchyma and other parenchyma modifications (figs. 2-4,
10, 11), homocellular vascular rays (figs. 5-9), and imperforate trache-
ary elements with simple or vestigially bordered pits. This fact causes
us to wonder at the relatively low taxonomic placement of Capparaceae
and the comparative primitiveness of the families with which it is
ordinarily associated in some schemes of plant classification.

The authors appreciate that the phylogenetic position of a family
containing 45 genera cannot be established through a brief study of
one genus using a single criterion, i.e., xylem anatomy. It is sug-
gested, however, that Capparaceae, with almost 50 percent mono-
typic genera, may be a highly derived taxon in which the evolution-
ary links among the genera have passed out of existence over the
ages. What remains today are the end-products of evolution; the
connecting stages, in the form of transitional taxa, are extinct.

Bessey (1914) derives the Rhoeadales (containing Capparaceae)
directly from Ranales; Engler and Diels (1936) also place Rhoeadales
in close association with Ranales; and in the Bentham and Hooker
(1862) system, Capparaceae are placed in the Parietales next to
Ranales. Wettstein (1935) treats Capparaceae in Rhoeadales and
mentions that they are related to both Cruciferae and Papaveraceae.
Furthermore, he states that Capparaceae contain the types from
which the Cruciferae could have arisen. Pax and Hoffman (1936)
say that without doubt Capparaceae belong in Rhoeadales and are
most closely allied to Cruciferae.

In light of the above, it is rather interesting to examine the position
of Capparaceae in Hutchinson’s (1959) treatment. Here they occur
quite high in the Lignosae as the type of Capparales. He derives
this order from the Dilleniales via Bixales and Pittosporales. Hutch-
inson regards the similarity in flower and fruit structure between
Cruciferae and Capparales as parallel evolution. Futhermore, in
addition to Capparaceae, Hutchinson includes Moringaceae and
Tovariaceae (Tovaria is included in Capparaceae by Bentham and
Hooker) in Capparales. Takhtajan (1954), like Hutchinson, gives
Capparales a more derived placement in his scheme. He shows the
order emanating from Cistales, along with Cucurbitales, Salicales, and
Tamaricales. However, Takhtajan (1959) retains Cruciferae in
Capparales, whereas Hutchinson includes it in Cruciales, a separate
order.

Capparis wood differs markedly from that in some families ordi-
narily associated with Capparaceae (1.e., Papaveraceae, Cruciferae,
and Resedaceae), in having homocellular rays consisting solely of
procumbent cells. The pore complexes described above for Capparis
have not been recorded in the woods of the Papaveraceae, Cruciferae,

36 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

or Resedaceae which are anatomically specialized along other lines
(cf. Metcalfe and Chalk, 1950). In certain respects, the wood of
Moringa resembles Capparis in that rays are homocellular and storied
axial parenchyma is present. Metcalfe and Chalk (1950) place some
emphasis on the presence of myrosin cells in parietalean taxa as a
significant indicator of relationship. All the taxa mentioned above
possess myrosin cells except Papaveraceae (Hegnauer, 1961). It is
noteworthy in this regard that Moringaceae, which Hutchinson con-
siders in his taxonomically elevated Capparales, is placed by Bentham
and Hooker in Sapindales. Without the benefit of detailed anatomi-
cal and associated studies in the other genera of Capparaceae, and com-
parative investigations into the anatomy of Papaveraceae, Cruciferae,
Resedaceae, and Moringaceue, a brief survey such as this of the anatomy
of families reputedly related to Capparaceae can only lead to wild
speculation. It is sufficient to say that our study lends some ana-
tomical support to Takhtajan’s and Hutchinson’s placement. of
Capparaceae in a separate order, more highly placed than the pari-
etalean families with which it is usually associated.

Anatomically, the most highly advanced species among those
investigated appears to be C. angulata. That this is so is indicated
by the combination of pronounced storying among the cells of the
axial parenchyma and vessels, coupled with the highly modified
banded, vasicentric, aliform, and confluent parenchyma distribution
(figs. 9, 11). C. pulcherrima is also more highly specialized than the
other species; it lacks the storied structure of C. angulata, but exhibits
the same kind of parenchyma specialization (fig. 10).

Intrageneric Alignment

The species of Capparis can be divided into two groups based
upon the seriation of rays. At first it seemed desirable to make a
separation into three groups: (1) species with uniseriate rays, (2) species
with rays having short multiseriate segments supporting several-
to many-celled uniseriate wings, and (3) species with multiseriate
rays and only a single apical cell, i.e., fusiform as viewed in the tan-
gential section. However, closer scrutiny caused us to alter our
course and to unite the first two groups. We found that woods with
predominantly uniseriate rays always had some rays which fell into
the second group, and woods in the second group often had many uni-
seriate rays (figs. 6, 7), so that apparently no consistent separation
between groups 1 and 2 could be made (ef. remarks on anatomical
distinction between C. flexwosa and C. cynophallophora). Therefore,
only two groupings, based on ray width, were devised: (I) species
with predominantly uniseriate rays, or with rays composed of short
bi- (or tri-) seriate segments having conspicuous several- to many-

WOODS OF FLORIDA KEYS—STERN AND OTHERS 37

celled, uniseriate extensions (figs. 6, 7), and (II) species with pre-
dominantly multiseriate rays which appear fusiform on the tangential
section and possess only one or two cells at their apices (figs. 8, 9).
Further studies involving more species might show transitions be-
tween these two groups, but on the basis of this investigation they
appear sufficiently exclusive of each other.

The species studied in our investigation were dispersed into the
sections of Capparis recognized in the treatment of Pax and Hoffmann
(1936). The appropriate ray classification was placed next to each
binomial and the results are outlined in table 2. Although representa-
tion of species in the various sections is in some cases meager, it is
noteworthy, excepting section Cynophalla, that the sections are
homogeneous with respect to the ray type shown. Whether this
taxonomic-anatomic device would stand up if further species were
introduced, remains to be shown. Or, as may be the case in Cyno-
phalla, the anatomical evidence may indicate a lack of taxonomic
homogeneity within the section. Substantiation for the latter was
mentioned previously. Recent work of W. R. Ernst has shown
that C. cynophallophora belongs in section Quadrella, rather than
Cynophalla.

Summary

Although Capparis flecuosa and C. cynophallophora are easily
separated, even by their gross foliar morphology, the anatomical
distinctions are more subtle, and possibly less reliable. In C. ecyno-
phallophora, vascular rays are largely uniseriate, whereas in C.
flexuosa, bi- and triseriate rays predominate.

Observations made on sections cut from fluid-preserved material,
showed that the banding in the wood of the two capers from the
Florida Keys, and in other species, is caused solely by alternating
layers of thick- and thin-walled prosenchymatous cells. Axial
parenchyma is strictly paratracheal and is not normally a part of
these bands. The groundmass is purely prosenchymatous. Fiber-
tracheids in O. flecuosa and C. cynophallophora may remain living for
longer or shorter periods after formation by the vascular cambium
and ontogenetic maturity takes place in these cells. This phenomenon
is probably more widespread than heretofore realized.

The wood anatomy of Capparis can be characterized by the presence
of pore complexes, alternate intervascular pitting and simple per-
foration plates, paratracheal parenchyma, homocellular vascular rays,
and imperforate tracheary elements with simple or vestigially bordered
pits. Also, certain species exhibit storied structure. These features
are all associated with phylogenetic advancement and indicate that

35 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Capparis, and possibly Capparaceae, occupies a more derived position
than it is usually afforded.

Using the seriation of vascular rays, the species of Capparis can be
separated into two groups. These groups follow the Pax and Hoff-
mann sectional delimitations of the genus except for the section
Cynophalla, which appears anatomically heterogeneous. Floral
studies of the two Keys species of Capparis, both placed in Cynophalla
by Pax and Hoffmann, give taxonomic support to this suggestion.
More intensive studies on a broader sampling of species will probably
produce at least two more anatomical features useful in separating
the species of Capparis: configuration of the axial parenchyma, and
presence or absence of storying. Application of these features will
serve to test the proposed separation of Capparis by ray seriation,
and may enable us further to refine our anatomical system.

39

OF FLORIDA KEYS-—-STERN AND OTHERS

WOODS

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40

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WOODS OF FLORIDA KEYS—STERN AND OTHERS 4l

TABLE 2.—Anatomical subdivision of Capparis based on ray structure

Par & Hoffmann

Section Species Ray Type
Capparis (‘‘Eucapparis’’) galeata II
Cynophalla eynophallophora I

flexuosa I
tarapotensis II
nitida II
verrucosa I
portoricensis II
linearis I
Mesocapparis crotonantha II
Homback decidua II
Capparidastrum tenuisiliqua I
oxysepala I
baducca I
quina I
sola I
Quadrella odoratissima I
Colicodendron schunkei II
angulata II
Calanthea ferruginea I
pulcherrima I
Busbeckia nobilis II

Literature Cited

Baruey, I. W.

1933. The cambium and its derivative tissues. No. VIII. Structure,
distribution and diagnostic significance of vestured pits in
dicotyledons. Journ. Arnold Arb., vol. 14, pp. 259-273.

BENTHAM, G., and Hooker, J. D.
1862. Capparideae. Genera plantarum, vol. 1, part 1, pp. 103-110.
BgEssEy, C. E.
1915. The phylogenetic taxonomy of flowering plants. Ann. Missouri Bot.
Garden, vol. 2, pp. 109-164.
CARLQUIST, S.
1961. Comparative plant anatomy. Pp. i-ix+146.
Cuattraway, M. M.

1955. Crystals in woody tissues; Part I. Tropical Woods, vol. 102,

pp. 55-74.
Cozzo, D.

1944. La estructura estratificada del lefio de Capparis salicifolia Gris.
Revista Ingen. Agron., vol. 6, pp. 151-153.

1946. Relaciédn anatémica entre la estructura del lefio de las especies
argentinas de ‘‘Capparis’” y ‘‘Atamisquea.”’ Lilloa, vol. 12,
pp. 29-37.

Cozzo, D., and Crist1an1, L. Q.

1950. Los generos de fanerogamas argentinas con estructura lefiosa
estratificada. Revista Inst. Invest. Cienc. Nat. Buenos Aires
Bot., vol. 1, pp. 363-405.

42 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Cumetrg, B. G.
1960. Anatomical studies in the Leguminosae. Tropical Woods, vol. 113,
pp. 1-47.
DarRuinaTon, C. D., and Wrutn, A. P.
1956. Chromosome atlas of flowering plants. Pp. i-xix+519.
Dr Canpo.ie, A. P.
1824. Capparideae. Prodromus systematis naturalis regni vegetabilis, vol.
1, pp. 237-254.
Enauer, A., and Diets, L.
1936. Syllabus der Pflanzenfamilien. Ed. 11, pp. i-xlii+419.
Ernst, W. R.
1963. The genera of Capparaceae and Moringaceae in the southeastern
United States. Journ. Arnold Arb., vol. 44, pp. 81-95.
Esau, K.
1953. Plant anatomy. Pp. i-xii+735.
Fann, A., and Lesnem, B.
1963. Wood fibers with living protoplasts. New Phytol., vol. 62, pp. 91-98.
Fawcett, W.
1914. Notes on Jamaican species of Capparis. Journ. Bot., vol. 52, pp.
142-144.
Harrar, E. 8.
1946. Note on starch grains in septate fiber-tracheids. Tropical Woods,
vol. 85, pp. 1-9.
HeGNAuER, R.
1961. Die Gliederung der Rhoeadales sensu Wettstein im Lichte der In-
haltstoffe. Planta Medica, vol. 9, pp. 37-46.
HvutcuHinson, J.
1926. The families of flowering plants. Vol. 1. Dicotyledons. Pp. i-
xii+ 328.
1959. The families of flowering plants. Vol. 1. Dicotyledons. Ed. 2,
pp. i-xi-+510.
Jacors, M.
1960. Capparidaceae. Flora Malesiana, ser. 1, vol. 6, part 1, pp. 61-105.
JANE, F. W.
1956. The structure of wood. Pp. i-vii+427.
Macsripe, J. F.
1938. Capparidaceae. Flora of Peru. Publ. Field Mus. Bot., vol. 13,
part 2, no. 3, pp. 984-1006.
Merca.re, C. R., and Cuauk, Lh.
1950. Anatomy of the dicotyledons. Vol. 1, pp. i-lxiv-+ 1-724; vol. 2, pp.
725-1500.
Pax, F., and HorrMann, K.
1936. Capparidaceae. Jn Eneler and Prantl, Die natiirlichen Pflanzen-
familien, ed. 2, vol. 17b, pp. 146-233.
RapDLKOFER, L.
1884. Ueber einige Capparis-Arten. Sitz-ber. Akad. Miinchen II, vol. 14,
pp. 101-182.
1887. Ueber einige Capparis-Arten. Sitz-ber. Akad. Miinchen II, vol.
17, pp. 365-422,
STERN, W. L., and CHamsers, K. L.
1960. The citation of wood specimens and herbarium vouchers in anatomical
research. Taxon, vol. 9, pp. 7-13.

WOODS OF FLORIDA KEYS—STERN AND OTHERS 43

TakarTas(1)An, A. L.
1954. Origins of angiospermous plants. Pp. 68 [Translated from the Rus-
sian by Olga H. Gankin, 1958].
1959. Die Evolution der Angiospermen. Pp. i-viii+344.
WETTSTEIN, R.
1935. Handbuch der systematischen Botanik. Pp. i-x+1152.

.S. GOVERNMENT PRINTING OFFICE;:1963

UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED StaTES NATIONAL HERBARIUM

VOLUME 34, Part 3

SYSTEMATIC ANATOMY AND ONTOGENY OF
THE STEM IN PASSIFLORACEAE

By Epwarp S. AYENsu and

Witiiam L. STERN

SMITHSONIAN INSTITUTION ¢ WASHINGTON, D.C. © 1964

SYSTEMATIC ANATOMY AND ONTOGENY OF
THE STEM IN PASSIFLORACEAE

By Epwarp S. AyeEnsu and Wiiutam L. STERN!

Although the taxonomy of the passionflower family, Passifloraceae,
has been studied extensively, especially that of the genus Passiflora
(Killip 1938), the anatomy and ontogeny of the woody stem have
been little explored. This was clearly pointed out by the studies of
Stern and Brizicky (1958), who recorded for the first time in the family
the presence of anomalous stem growth and storied structure in the
xylem of Passiflora multiflora L. Interestingly, the occurrence of
these two phenomena seemed to accord with Killip’s suggestion that
P. multiflora, the single member of his subgenus Apodogyne, ought to
be segregated as a separate genus on the basis of its sessile ovary.
Obaton (1960) encountered anomalous stems in Adenia cissampeloides
Harms (included phloem; i.e., phloem strands surrounded by xylem),
and Crossostemma laurifolium Planch. (intruded phloem; i.e., ribs or
flanges of phloem extending into the xylem cylinder from the bark side
of the stem) of the Passifloraceae. Her observations indicate that
certain other species of Adenia (e.g., Adenia lobata Engl.) have normal
stem structure, even in the thickest portions. She felt’ compelled,
therefore, to segregate as a distinct species a specimen of Adenia lobata,
obtained from Bingerville, Ivory Coast, which showed abnormal stem
structure. Furthermore, she asserts that Adenia kontiensis A. Chev.,
which has long been confused with Adenia lobata, is distinguished from
it by the less numerous, thicker, and more rounded tubercles on the
stem. Because Obaton’s study of stem structure in Passifloraceae
lacks detailed anatomical descriptions and specimens are not docu-
mented by herbarium voucher citation, it is of little use taxonomically.

Except for the works of Stern and Brizicky, and Obaton, there is no
record in the anatomical literature that anomalous stem growth occurs
in the Passifloraceae. Schenck (1893) states that growth in the stem
of Passifloraceae is normal; Solereder (1908) emphasizes that even the
climbing forms of Passifloraceae have normal structure. Solereder,
furthermore, repudiates Schwendener’s (1874) assertion that Passiflora

1 We extend our thanks and appreciation to all who have assisted us in this study, particularly Prof. I. W.
Bailey of Harvard University; Dr. Alberta M. W. Mennega of the State University of Utrecht, Netherlands;
Dr. Richard A. Howard of the Arnold Arboretum; Mr. Richard A. Mark of the 8. J. Record Memorial
Wood Collection at Yale University; Senhor J, do Espirito-Santo, Bafaté, Portuguese Guinea, and Dr. E.
Laing, Ghana University, Legon.

45

46 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

trifasciata has 6 medullary vascular bundles which are provided on the
inside and the outside with small bast-coverings (bicollateral) for
the protection of the conducting cells. Dr. L. Chalk of Oxford Univer-
sity, in an informal communication, suggested that Schwendener’s
remarks at least imply that he suspected the occurrence of anomalous
structure in this particular species of Passiflora. Neither Metcalfe
and Chalk (1950) nor Boureau (1957) make mention of this feature in
Passifloraceae.

In the taxonomic studies of Passifloraceae by Harms (1925), Masters
(1871), and Killip (1938), no mention of atypical stem development in
this family has been noted. Ordinarily, development of the anomalous
configuration commences in the thicker portions of the stem, not reg-
ularly collected for herbarium specimens. Because of this condition,
taxonomists have perhaps overlooked these strange stems.

Owing to certain similarities in the anatomy and morphology of
Passifloraceae and Flacourtiaceae, taxonomists have found it difficult
to decide to which the genus Paropsia should be assigned. Bentham
and Hooker (1867) included under Passifloreae, the genera Passiflora,
Tacsonia, Deidamia, Basanthe, Tryphostemma, Paropsia, Smeathman-
nia, Barteria, and Crossostemma. Harms (1893) later transferred
Paropsia to Flacourtiaceae, and Gilg (1925) included the genera
Paropsia, Soyauxia, Hounea, Smeathmannia, Paropsiopsis, and Bar-
teria under the tribe Paropsieae of this same family. Fernandes and
Fernandes (1958a, 1958b) disagree with Harms and Gilg and assert
that Paropsia should be placed in Passifloraceae. Brizicky (1961)
pointed out that the present delimitations of the family seem to be
fairly natural. However, he remarked that a taxonomic re-evalua-
tion of Passifloraceae and their closest allies, such as Flacourtiaceae—
Paropsicae, based on comparative anatomy and other botanical dis-
ciplines, is needed.

In a preliminary report on the woods of Flacourtiaceae, Tupper
(1934) indicated that axial parenchyma is generally lacking. How-
ever, he encountered abundant axial wood parenchyma in the genus
Paropsia in support of den Berger’s (1928) observation that this genus
has abundant, diffuse wood parenchyma. Metcalfe and Chalk (1950)
mention that although there are certain anatomical differences be-
tween Paropsia and the genera of Passifloraceae, there are also an-
atomical similarities, and, for example, the occurrence of fibriform
vessel members in some species may support its affinity with
Passifloraceae.

Palynological studies by Erdtman (1952) have shown that pollen
erains of Paropsia are more similar to those of Passifloraceae than to
those of Flacourtiaceae. The pollen grains of Paropsia brazzeana, for
instance, are usually 6-colporate and subprolate; the sexine is as thick

AYENSU AND STERN-—STEM IN PASSIFLORACEAE 47

as the nexine, and reticulate. Erdtman suggests that the tribe
Paropsieae (Flacourtiaceae) should, at least in part, be referred to
Passifloraceae.

In general, our investigations uphold the anatomical findings of
Tupper and den Berger, and support Bentham and Hooker (1867),
and Fernandes and Fernandes (1958a, 1958b) in their assertions that
the genus Paropsia belongs with Passifloraceae.

Passifloraceae consists of 12 genera with over 500 species distrib-
uted throughout the tropics. Eight of the genera are confined to
the Old World, three are exclusively American and one, Passiflora, is
predominantly New World. Most of the Old World genera have few
species; more than two-thirds of all species in the family belong to
Passiflora.

Members of the Passifloraceae (figs. 1, 2) are herbaceous or woody,
tendril bearing lianas, or erect shrubs and trees. Leaves are alternate,
simple or compound, and have entire, toothed, or deeply cleft margins,
with predominantly 3-5 nerved blades. Petioles are often glandulif-
erous; stipules are setaceous or broadly ovate, but sometimes
foliaceous.

Flowers may be brilliantly colored or greenish, many centimeters in
diameter or less than one. They may be axillary or in cymes, but are
usually reduced to solitary or paired peduncles. Bracts are ordinarily
present. Five (4), rarely 3, basally connate sepals are present. The
petals 5(4), or wanting, are distinct or briefly basally connate, often
smaller than the sepals. Sepals and petals are imbricate. The floral
tube is usually associated with a corona. The androecium consists
of 5 stamens, rarely 10, usually opposite the petals; anthers are
versatile, 2-locular at anthesis, and dehisce longitudinally. The
gynoecium consists of 3-5 carpels; stigmatic branches are capitate,
clavate, or discoid; the styles number as many as the carpels; the
ovary is stipitate, rarely sessile, 1-locular with numerous anatropous
ovules on 3-5 parietal placentae. The fruit is a berry or a loculicidal
capsule; the seed has a straight embryo and fleshy endosperm. Both
Killip (1938) and Masters (1871) have excellent descriptions of these
highly modified flowers, accompanied by drawings and explanations,

Materials and Methods

Table 1 lists the 60 stem and wood specimens (44 species of 9
genera) examined. The citation of specimens follows the recom-
mendations of Stern and Chambers (1960), and herbarium vouchers
are listed as well as the names and numbers of the collectors. Names
of species are in accordance with those stated in Killip (1938) for
American Passifloraceae.

48 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Microtechnical procedures followed were modified from those
outlined by Wetmore (1932) and are standard for the preparation of
sections of stem and wood. Microscope slides of wood specimens
prepared for this investigation have been deposited in the Division
of Plant Anatomy of the Smithsonian Institution (USw).

Diagnostic characters for descriptions of passifloraceous woods
were selected from Tippo’s (1941) list. For the most part, terms
used in the anatomical descriptions comply with those approved by
the Committee on Nomenclature of the International Association of
Wood Anatomists (1957). In a few instances, however, certain
liberties were taken with the definitions.

Lack of sufficient wood material in a number of cases made the
preparation of macerations impracticable, hence, length measurements
of fibers and vessel elements were taken from sections. Therefore,
our method of making measurements cannot conform exactly to the
recommendations of Chalk and Chattaway (1934, 1935) that total
lengths of cells be used. Tangential vessel diameters and lengths of
fibers and vessel elements were recorded from 30 cells in each specimen.

Because of the conflicting and ambiguous results brought on by
employing only names to categorize vascular rays (Kribs 1935), it
was thought best to employ the suggestions of the Committee on
Nomenclature (1957). These direct the anatomist to describe rays
using brief sentences or phrases to point out the width, height,
frequency of occurrence, and cellular composition. Classification of
axial wood parenchyma used here is in most part that presented by
Metcalfe and Chalk (1950). The various types of parenchyma
distribution fall into two main categories: apotracheal, in which
parenchyma strands show no regular relation to vessels; and para-
tracheal, in which parenchyma strands show a close and regular
association with vessels.

Four descriptive terms have been chosen to characterize the
appearance on the transverse surface of the mature vascular cylinder
of stems in Passifloraceae: (1) NoRMAL TypE—a stem in which
xylem and phloem cylinders are roughly circular and concentric with
little or no distortion of pith and cortex; (2) INCLUDED PHLOEM
TYPE—similar to (1), but with patches of phloem embedded in the
xylem cylinder; (3) INTERRUPTED TyPE—wedge-shaped sections of
xylem, alternating around the stem with similarly shaped sections
of phloem, characterized by marginal series of steps; (4) DISPERSED
typE—iregularly shaped, disoriented strands of xylem and phloem
associated with fragments of vascular cambium spread throughout a
parenchymatous matrix.

AYENSU AND STERN—STEM IN PASSIFLORACEAE

49

TABLE 1.—Specimens of Passifloraceae stems examined

Citation of specimens follows the recommendations of Stern and Chambers (1960). WOOD COLLECTIONS:
(Yw) The Samuel James Record Memorial Collection, Yale University, New Haven; (USw) Division
of Plant Anatomy, Smithsonian Institution, Washington; (Aw) The Arnold Arboretum, Cambridge;
(Uw) Botanical Museum and Herbarium, Utrecht. Hernaria: (TEF) Inspection Générale des
Eaux et Forts, Tananarive; (FHO) The Imperial Forestry Institute, Oxford; (Y) Yale School of
Forestry, New Haven; (A) The Arnold Arboretum, Cambridge; (US) U.S. National Herbarium,
Smithsonian Institution, Washington; (NY) New York Botanical Garden, New York; (U) Botanical
Museum and Herbarium, Utrecht; (K) Royal Botanic Gardens, Kew.]

Species
Adenia lobata (Jacq.)
Engl.
Androsiphonia adeno-
stegia Stapf
Barteria nigritiana
Hook. f.

Crossostemma laurifolium

Planch.

Dilkea johannesii Rodr.

Mitostemma glaziovii
Mast.

Paropsia adenostegia
(Stapf) Engl.

Paropsia madagascariensis

(Baill.) H. Perrier
Paropsia schliebenii
Sleumer

Paropsia vareciformis
Mast.

Passiflora arborea
Spreng.

Passiflora auriculata
H.B.K.

Passiflora auriculata
H.B.K.

Passiflora coccinea Aubl.

Passiflora caerulea L.

Passiflora coriacea A.
Rich.

Passiflora edulis Sims

Passifiora edulis Sims

Passiflora edulis Sims

Collector Herbarium Location and
and Number Geographical Origin Voucher Catalog Number
E.S. Ayensu. Ghana US
8.n.
G. P. Cooper Liberia Y Yw 15191
276
E. 8S. Ayensu. Ghana US
$.n.
C. Vigne 3503 Ghana FHO Yw 29724
A. Ducke 329 Brazil Y Yw 34093
A. C. Smith British Y Yw 35812
3117 Guiana
G. P. Cooper Liberia Yw 15140
211
— Madagascar TEF USw 27402
H. J. Schliee Tanganyika Y Yw 30003
ben 429
(5442)
H. E. Desch Malaya — Yw 29187
2003
(32695)
J. Cuatreeasas Colombia Y Yw 44493
22014
R. H. Wood- Panama A Aw 362
worth & Canal Zone
P. A. Vestal
362
W. L. Stern Trinidad US USw 30687
1585A
B. Maguire Suriname NY Yw 44096
24179
— — — Aw 5323
R. H. Wood- Panama A Aw 501
worth & Canal Zone
P.A. Vestal
501
— Cuba (cult. — Aw 5283
Soledad)
E. S. Ayensu. Ghana US
8.n.
W. L. Stern Kauai, US USw 25985
& S. Carl- Hawaii

quist 1287

50 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

TABLE 1.—Specimens of Passifloraceae stems excamined—Continued

Species
Passiflora elliptica
Gardn.
Passiflora fuchsiiflora
Hemsl.

Passiflora fuchsiiflora
Hemsl.

Passiflora fuchsiiflora
Hemsl.

Passiflora fuchsiiflora
Hemsl.

Passiflora gigantifolia
Harms (?)

Passiflora glandulosa
Cav.

Passiflora glandulosa
Cav.

Passiflora glandulosa
Cav.

Passiflora glandulosa
Cav.

Passiflora glandulosa
Cav.

Passiflora glandulosa
Cav.

Passiflora glandulosa
Cav.

Passiflora glandulosa
Cav.

Passiflora laurifolia L.

Passiflora maguirei
Killip

Passiflora maliformis
Vell.

Passiflora menispermi-
folia H. B. K.

Collector
and Number

H. 8. Irwin
2222

F. P. Jonker
& Daniels
817

F. P. Jonker
& Daniels
1098

J. Lanjouw
& J.C.
Lindeman
2639

B. Maguire
& D, B.
Fanshawe
22807

A. Rimbach
50

A. C. Smith
2822

BAFOG 4033

F. P. Jonker
«& Daniels
788

F. P. Jonker
& Daniels
830

J. Lanjouw
& C.J.
Lindeman
1820

J. Lanjouw
& J.C.
Lindeman
2765

J. C. Lindman

4530

A. A. Pulle
353

B. Maguire
& D. B.
Fanshawe
23285

R. H. Wood-
worth & P. A. Canal Zone

Vestal 637

Geographical Origin

Brazil

Suriname

Suriname

Suriname

British
Guiana

Ecuador

British
Guiana

French

Guiana
Suriname

Suriname

Suriname

Suriname

Suriname
Suriname

Cuba
British
Guiana

Cuba (cult.
Soledad)
Panama

Voucher

Y

U

qa Ga kK

ar GQ 4

TH

A

Herbarium Location and
Catalog Number

Yw 53094

USw 30677

USw 30678

Uw 1859,
USw
30686

Yw 43907

Yw 20743
Yw 35711
USw 30679

USw 30680

USw 30681

Uw 1559,
USw
30685

USw 30682

Uw 3142
USw 30683

Aw 25273
Yw 44006

Aw 5284

Aw 637

AYENSU AND STERN—STEM IN PASSIFLORACEAE 51

TaBLE 1.—Specimens of Passifloraceae stems examined—Continued

Species
Pagsiflora misera H.B.K.

Passiflora mollissima
(H.B.K.) Bailey = Tac-
sonia mollissima
H.B.K.

Passiflora multiflora L.

Passiflora nitida H.B.K.

Passiflora punctata L.

Passiflora quadrangularis
L.

Passiflora quadriglandu-
losa Radscheid

Passiflora riparia Mart.
ex Mast.

Passiflora rubra L.

Passiflora securiclata
Mast.

Passiflora seemannii
Griseb.

Passiflora spinosa (Poepp.

& Endl.) Mast.

Passiflora variolata
Poepp. & Endl.

Passiflora vitifolia
H.B.K.

Passifiora vitifolia
H.B.K.

Passiflora vitifolia
H.B.K.

Collector
and Number

R. H. Wood-
worth &
P. A. Ves-
tal 456

A. Rimbach
19

W. L. Stern
& G. K.
Brizicky
522

J. J. Wurdack
&L.S8.
Adderley
43025

R. H. Wet-
more & E.
Abbe 26

M. Grant
2617

G. H. M.
Lawrence
607

A. C, Smith
3157

A. C. Smith
3116

R. H. Wet-
more & E.
Abbe 147

J. J. Wurdack
& L. 8.
Adderley
43009

J. J. Wurdack
&L.S.
Adderley
43230

R. H. Wet-
more & R.
H. Wood-
worth s.n.

J. J. Wurdack

& L.S.
Adderley
43026

F. Woyt-
kowski 509

Geographical Origin
Panama
Canal Zone

Ecuador

Plantation
Key,
Florida

Venezuela

Panama
Canal Zone

Tahiti
Colombia
British

Guiana

British
Guiana

Panama
Canal Zone

Venezuela

Venezuela

Panama

Canal Zone

Venezuela

Peru

A

NY

NY

Herbarium Location and
Voucher

Catalog Number
Aw 456

Yw 19485

Yw 51303

USw 17808

Aw 26

Yw 26455

Aw 21303

Yw 35837

Aw 5286
Yw 35811

Aw 147

Usw 17798

USw 17923

Aw 854A

USw 17809

Yw 52812

52 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

TaBLE 1.—Specimens ef Passifloraceae stems examined—Continued

Collector Herbarium Location and
Species and Number Geographical Origin Voucher Catalog Number
Passiflora sp. nov. J.J. Wurdack Venezuela NY USw 17874
& L.8.
Adderley
43154
Passiflora sp. B. Maguire Venezuela NY USw 24782
28327
Passiflora sp. J.J. Wurdack Venezuela NY USw 17834
& LS.
Adderley
43055
Passiflora sp. F. P. Jonker Suriname U USw 30684
& Daniels
1227
Smeathmannia J. Espirito- Portuguese US USw 30675
laevigata Sol. ex R. Santo s.n, Guinea
Br.
Smeathmannia pubescens G.P. Cooper Liberia Y USw 5792
Sol. ex R. Br. 442
Smeathmannia pubescens G. P. Cooper Liberia K Yw 13701
Sol. ex R. Br. 51

Anatomy of the Family

Growth rings are absent in most of the species investigated except
for Paropsia schliebenti and Smeathmannia pubescens. All of the
woods are diffuse-porous, the randomly distributed pores ranging in
size from very small to moderately large. The occurrence of solitary
pores (figs. 3-6) characterizes the wood of Passifloraceae, although
radial (figs. 7-9) and tangential multiples (fig. 4), as well as pore
clusters, are found in some species. Pores in all species are rounded;
however, ‘flattened’? pores (fig. 3) sometimes appear. Pores are
variable in size even in different species of the same genus. In some
species of Passiflora, for example, vessel diameter is less than 100 gz,
while in other species the vessel diameter is over 3004. Vessel
element lengths are extremely short in Adenia and Passiflora to
moderately short in Crossostemma and Smeathmannia.

Slope of the end walls in the vessel elements varies from oblique
(fig. 14) to transverse. Oblique end walls are regularly associated with
wood from arboreal species. Perforation plates are almost exclu-
sively simple (fig. 11); in Paropsia adenostegia, both simple and sca-
lariform perforation plates (fig. 12) may occur in the same section. The
scalariform plates here have few bars, and at times show a tendency
toward the reticulate configuration. Metcalfe and Chalk (1950)
observed a few scalariform plates in Androsiphonia and Mitostemma
glaziovii, but none was observed in M. glaziovii during this
investigation.

AYENSU AND STERN—STEM IN PASSIFLORACEAE 53

The imperforate tracheary elements generally consist of fiber-
tracheids (fig. 13) bearing pits with very distinct borders. However,
tracheids (fig. 13) occur in some species, and very thick-walled libri-
form wood fibers (fig. 10) are commonly found in the arboreal mem-
bers of the family. The fiber lengths range from moderately short
to medium-sized.

Intervascular pitting is mostly alternate (fig. 21), and the pit
apertures are generally elliptical or flattened, and tend to be limited
by the border. In Passiflora fuchsiiflora coalesced apertures were
observed.

Fibriform vessel members (fig. 13) of Woodworth (1935) were
observed in most species of Passiflora and in Paropsia schliebenii.
These cells are longer than vessel elements in the same species, and
bear simple perforation plates that occur about one-fourth the distance
from each end of the element. Fibriform vessel members are fusi-
form cells more or less randomly distributed within the groundmass,
and show fewer axial connections among conspecific elements than
occur among elements making up typical vessels. Lateral wall
pitting of these cells is similar to that of the fiber-tracheids.

Vascular ray tissue (figs. 10, 17) is composed of both uniseriate and
multiseriate rays. Some species have typically high multiseriate rays;
in others, only biseriate rays with long uniseriate wings occur. The
tendency for rays to widen centrifugally, as seen in transverse sections,
is common in Passiflora. In most species of Passiflora storied uni-
seriate rays (fig. 17) are found among higher, nonstoried multiseriate
rays. Vascular rays may be heterocellular or homocellular. In
Paropsia vareciformis rays consist of procumbent cells associated with
uniseriate margins of square cells. Rays consist exclusively of upright
cells in Passtflora maguireit and Passiflora mollissima. Ontogenetic
splitting of multiseriate rays into uniseriate and biseriate rays was
observed in many species. Chalk and Chattaway (1933) reported on
the presence of perforated ray cells in Androsiphonia adenostegia.
Similarly perforated ray cells were observed in Passiflora vitifolia and
Passiflora elliptica (fig. 15).

Axial wood parenchyma is generally moderately abundant to very
abundant. The usual arrangement is apotracheal (figs. 7, 8, 20);
however, paratracheal parenchyma (figs. 3-6, 9) does occur, and in
some species it is vasicentric and aliform. Combinations of diffuse
and diffuse-in-aggregates, with a tendency toward banded, and even
vasicentric arrangements are found in the same species (fig. 9);
narrow bands occur in others. In Paropsia, the bands are wavy;
banded and aliform arrangements are both observed in Dilkea
johannesit; in Crossostemma laurifolium, only vasicentric parenchyma
arrangement occurs.

54. CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Pith flecks (fig. 16) occur in the wood of about 70 percent of the
Passiflora species studied. Three types of anomalous stem develop-
ment (see pp. 48, 60, 61) were observed in the species investigated: mcluded
phloem type, interrupted type (figs. 24, 25), and dispersed type
(fig. 23). Tyloses and gummy deposits were present in ray cells and
vessels (fig. 8) of a few species. Rhomboidal crystals, and occasion-
ally druses, were observed in most species investigated, particularly
in ray cells.

Anatomy of the Species

Crossostemma laurifolium: Growth rings absent; pores mainly soli-
tary, rounded, average diameter 198 », range 140 u-210 uw; average
vessel element length 424 y, range 311 u—-442 yu; vessel wall thickness 2 p-
5 w; perforation plates simple; vessel element end wall inclination
almost transverse; intervascular pitting opposite to alternate; imper-
forate tracheary elements fiber-tracheids, average length 614 yu, range
497 u-756 u; fibriform vessel members absent; vascular rays mostly
multiseriate without any uniseriate extensions, 2-5 cells wide, 8-12
cells high, few uniseriate, 5-14 cells high, mainly homocellular consist-
ing of procumbent cells, uniseriates sometimes showing upright cells;
axial parenchyma vasicentric; crystals present in ray cells; uniseriate
rays vaguely storied; stem anomalous (interrupted) ; pith flecks absent.

Dilkea johannesti: Growth rings absent; pores mostly solitary,
rounded, average diameter 205 y, range 104 y-211 u; average vessel
element length 378 u, range 335 u-401 yu; vessel wall thickness 4 p-Sy;
perforation plates simple; vessel element end wall inclination trans-
verse; intervascular pitting alternate; imperforate tracheary elements
fiber-tracheids, average length 560 4, range 433 -741 u; fibriform
vessel members absent; vascular rays mostly multiseriate without
any uniseriate extensions, 2-6 cells wide, 5-13 cells high, some uni-
seriate, 4-15 cells high, mainly heterocellular; axial parenchyma
diffuse, abundant; crystals present in ray cells; uniseriate rays vaguely
storied; stem normal; pith flecks absent.

Mitostemma glaziovi: Growth rings absent; pores mostly solitary,
rounded, average diameter 112 yu, range 92 u-130 yu; average vessel
element length 416 uw, range 392 y-478 yw; vessel wall thickness 3 p-5 yp;
perforation plates simple; vessel element end wall inclination trans-
verse; intervascular pitting alternate; imperforate tracheary elements
fiber-tracheids, average length 550 yu, range 520 u-719 yu; fibriform
vessel members absent; vascular rays mostly multiseriate without
any uniseriate extensions, 2—4 cells wide, 7-12 cells high, few uniseriate,
8-13 cells high, heterocellular; axial parenchyma diffuse; crystals
present in ray cells; storying absent; stem normal; pith flecks absent.

AYENSU AND STERN—STEM IN PASSIFLORACEAE 55

Paropsia adenostegia: Growth rings inconspicuous; pores mainly
solitary, few radial multiples, angular to rounded, average diameter
50 wu, range 35 »-56 4; average vessel element length 450 yu, range
420 p-773 gw; vessel wall thickness 2 u-3 »; perforation plates scalari-
form and simple; vessel element end wall inclination oblique; inter-
vascular pitting alternate; imperforate tracheary elements libriform
wood fibers, average length 668 p, range 574 u-968 u; fibriform vessel
members absent; vascular rays mainly uniseriate, 5-12 cells high, and
biseriate, 5-12 cells high, heterocellular; axial parenchyma diffuse,
banded in places; crystals absent; storying absent; stem normal;
pith flecks absent.

Paropsia madagascariensis: Growth rings absent; pores mainly in
radial multiples, rounded, average diameter 85 yw, range 64 y-127 p;
average vessel element length 468 », range 420 u-780 u; vessel wall
thickness 2 y-3 u; perforation plates simple; vessel element end wall
inclination oblique to transverse; intervascular pitting alternate;
imperforate tracheary elements libriform wood fibers, average length
580 pw, range 394 y-803 yu; fibriform vessel members absent; vascular
rays multiseriate without any uniseriate extensions, 2-3 cells wide,
5-11 cells high, heterocellular; axial parenchyma diffuse; crystals
present in ray cells; storying absent; stem normal; pith flecks absent.

Paropsia schliebenii: Growth rings present; pores mainly in radial
multiples, rounded, average diameter 45 yw, range 27 u-62 uw; average
vessel element length 590 », range 398 y-772 y; vessel wall thickness
2 u-3 u; perforation plates simple; vessel element end wall inclination
oblique to transverse; intervascular pitting alternate; imperforate
tracheary elements libriform wood fibers, average length 775 py, range
424 y-907 uw; fibriform vessel members present; vascular rays multi-
seriate, 2-3 cells wide, 13-28 cells high, few with uniseriate wings 4-8
cells high, largely homocellular, cells mainly procumbent but with a
few upright cells; axial parenchyma diffuse; crystals absent; uniseriate
rays vaguely storied; stem normal; pith flecks absent.

Paropsia vareciformis: Growth rings inconspicuous; pores mainly in
radial multiples, rounded, average diameter 70 yw, range 62 u-110 pu;
average vessel element length 480 yw, range 382 u-697 u; vessel wall
thickness 2 u-3 yu; perforation plates simple; vessel element end wall
inclination almost transverse; intervascular pitting alternate; im-
perforate tracheary elements fiber-tracheids and libriform wood fibers,
average length 668 u, range 511 »-939 yu; fibriform vessel members
absent; vascular rays multiseriate 2-3 cells wide, 18-34 cells high, few
with uniseriate wings 4-8 cells high, mainly homocellular, consisting
solely of procumbent cells, other rays with some upright cells; axial
parenchyma diffuse; crystals present in ray cells; uniseriate rays storied
in places; stem normal; pith flecks absent.

56 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Passiflora arborea: Growth rings present; pores 50 percent solitary,
30 percent radial multiples, 20 percent clusters, rounded, average
diameter 130 yu, range 98 y-152 yw; average vessel element length 343 yp,
range 295 u-587 uw; vessel wall thickness 2 u-3 yu; perforation plates
simple; vessel element end wall inclination oblique to transverse;
intervascular pitting alternate; imperforate tracheary elements fiber-
tracheids and libriform wood fibers, average length 481», range
394 u-647 yu; fibriform vessel members absent; vascular rays biseriate
with long uniseriate wings 9-22 cells high, heterocellular; axial paren-
chyma mostly diffuse, diffuse-in-aggregates to banded and vasicentric;
storying absent; stem normal; pith flecks present.

Passiflora auriculata: Growth rings absent; pores mainly solitary,
rounded, average diameter 210 yu, range 198 u-230 u; average vessel
element length 362 u, range 303 u-479 uw; vessel wall thickness 2 u-5 y;
perforation plates simple; vessel element end wall inclination trans-
verse; intervascular pitting opposite to alternate; imperforate trache-
ary elements fiber-tracheids, average length 472 u, range 403 »-723 yu;
fibriform vessel members present; vascular rays mainly multiseriate
2-8 cells wide, 7-30 cells high, few with uniseriate wings 4-9 cells
high, some uniseriate rays 5-12 cells high, heterocellular; axial paren-
chyma diffuse; crystals present in ray cells; uniseriate rays storied;
stem anomalous (included phloem); pith flecks absent; tyloses present.

Passiflora coceinea: Growth rings absent; pores mainly solitary,
rounded, average diameter 210 uw, range 182 y-231 u; average vessel
element length 385 yu, range 307 »-478 yu; vessel wall thickness 2 u-7 y;
perforation plates simple; vessel element end wall inclination oblique
to transverse; intervascular pitting alternate; imperforate tracheary
elements tracheids and fiber-tracheids, average length 437 y, range
410 p-629 y; fibriform vessel members present; vascular rays mainly
uniseriate 1-18 cells high, some multiseriate without any uniseriate
extensions, 2-8 cells wide, 6-13 cells high, heterocellular; axial paren-
chyma diffuse with local banding; crystals absent; uniseriate rays
storied; stem anomalous (interrupted); pith flecks present; tyloses
present.

Passiflora edulis: Growth rings absent; pores mainly solitary,
rounded, average diameter 131 yu, range 101 u-168 yu; average vessel
element length 185 y, range 143 y—208 yu; vessel wall thickness 2 u-7 yp;
perforation plates simple; vessel element end wall inclination trans-
verse; intervascular pitting opposite to alternate; imperforate trache-
ary elements fiber-tracheids, average length 315 yu, range 298 u472 yp;
fibriform vessel members present; vascular rays mainly uniseriate 1-14
cells high, mostly 6-12 cells high, some multiseriate 2-3 cells wide,
8-12 cells high, uniseriate wings 4-8 cells high, few very high multi-
seriate rays, heterocellular ; axial parenchyma mainly diffuse with some

AYENSU AND STERN—STEM IN PASSIFLORACEAE 57

banding; crystals present in ray cells; uniseriate rays storied; stem
anomalous (interrupted); pith flecks absent.

Passiflora elliptica: Growth rings absent; pores 30 percent solitary,
62 percent radial multiples, 8 percent clusters, rounded, average
diameter 117 uw, range 96 u-184 yu; average vessel element length 316 u,
range 299 y-437 yw; vessel wall thickness 2 u-5 »; perforation plates
simple; vessel element end wall inclination oblique to transverse;
intervascular pitting opposite to alternate; imperforate tracheary
elements fiber-tracheids, average length 420 4, range 413 p-489 uy;
fibriform vessel members present; vascular rays mainly multiseriate
without any uniseriate extensions, 2-8 cells wide, 7-12 cells high, few
uniseriate 4-9 cells high, heterocellular; axial parenchyma diffuse;
crystals absent; uniseriate rays vaguely storied; stem normal; pith
flecks present; tyloses present.

Passiflora fuchstiflora: Growth rings absent; pores mainly solitary,
rounded, average diameter 135 yu, range 97 u-154 yw; average vessel ele-
ment length 396 yu, range 284 y-473 yw; vessel wall thickness 2 u-5 y;
perforation plates simple; vessel element end wall inclination oblique
to transverse; intervascular pitting opposite to alternate; imperforate
tracheary elements fiber-tracheids, average length 4714, range
435 y-622 u; fibriform vessel members present; vascular rays mainly
multiseriate 2-8 cells wide, 6-13 cells high, uniseriate wings 4-6 cells
high, few very high multiseriate rays, some uniseriate 4-8 cells high,
heterocellular ; axial parenchyma diffuse and diffuse-in-aggregates with
a few narrow bands; crystals present in ray cells; uniseriate rays and
parenchyma strands vaguely storied; stem normal; pith flecks present.

Passiflora glandulosa: Growth rings vague; pores 60 percent solitary,
40 percent radial multiples, rounded, average diameter 200 yp, range
142 »-254 yw; average vessel element length 341 uw, range 311 y-441 py;
vessel wall thickness 2 u—-7 uw; perforation plates simple; vessel element
end wall inclination oblique to transverse; intervascular pitting alter-
nate; imperforate tracheary elements fiber-tracheids, average length
403 uw, range 361 y—-547 y; fibriform vessel members present; vascular
rays mainly uniseriate 1-7 cells high, some multiseriate 2-8 cells wide,
4-9 cells high, few very high multiseriate rays, heterocellular; axial
parenchyma diffuse and in narrow bands; few crystals present in ray
cells; uniseriate rays storied; stem anomalous (interrupted) ; pith flecks
present.

Passifiora laurifolia: Growth rings absent; pores mainly solitary,
rounded, average diameter 124 p, range 98 u-155 yw; average vessel ele-
ment length 318 yp, range 264 u—410 u; vessel wall thickness 2 y—-5 py; per-
foration plates simple; vessel element end wall inclination oblique to
transverse; intervascular pitting opposite to alternate; imperforate
tracheary elements fiber-tracheids, average length 474 yu, range

58 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

401 »-498 uy; fibriform vessel members present; vascular rays mainly
multiseriate 2-5 cells wide, 4-13 cells high, few uniseriate 1-15 cells
high, few very high multiseriate rays, homocellular, consisting entirely
of upright cells; axial parenchyma diffuse; crystals present in ray cells;
uniseriate rays vaguely storied; stem normal; pith flecks present.

Passiflora maguirei: Growth rings absent; pores 85 percent solitary,
radial multiples 10 percent, clusters 5 percent, rounded, average
diameter 75 uw, range 42 w-114 uw; average vessel element length 211 yp,
range 160 y-351 uw; vessel wall thickness 2 u-4 yu; perforation plates
simple; vessel element end wall inclination oblique to transverse; inter-
vascular pitting alternate; imperforate tracheary elements fiber-
tracheids and libriform wood fibers, average length 488 p, range
408 u-730 u; fibriform vessel members absent; vascular rays mainly
uniseriate 1-12 cells high, some multiseriate without uniseriate exten-
sions, 2-8 cells wide, 6-12 cells high, homocellular, composed solely of
upright cells; axial parenchyma diffuse, diffuse-in-aggregates, and in
narrow bands; crystals absent; uniseriate rays vaguely storied; stem
normal; pith flecks absent.

Passiflora mollissima: Growth rings absent; pores 90 percent soli-
tary, 10 percent radial multiples, rounded, average diameter 111 pg,
range 92 u-170 uw; average vessel element length 368 yu, range 284 p-
433 yw; vessel wall thickness 2 u-5 yw; perforation plates simple; vessel
element end wall inclination oblique to transverse; intervascular
pitting opposite to alternate; imperforate tracheary elements fiber-
tracheids, average length 432 u, range 401 u—-520 u; fibriform vessel
members present; vascular rays mainly multiseriate without uniseriate
extensions, 2-5 cells wide, 6-14 cells high, few uniseriate 1-10 cells
high, homocellular, comprising only square cells; ray cells with simple
perforations; axial parenchyma diffuse; crystals present in ray cells;
uniseriate rays vaguely storied; stem normal; pith flecks absent.

Passiflora multiflora: Growth rings absent; pores mainly solitary,
rounded, average diameter 128 pn, range 102 y-233 yw; average vessel
element length 345 yp, range 210 u-445 pu, vessel wall thickness 2 y-5 p;
perforation plates simple; vessel element end wall inclination trans-
verse; intervascular pitting alternate; imperforate tracheary elements
fiber-tracheids, average length 422 uy, range 354 u-591 uw; fibriform
vessel members present; vascular rays mainly uniseriate 2-15 cells
high, some biseriate 25 or more cells high, homocellular, entirely of
procumbent cells; axial parenchyma apotracheal and paratracheal;
crystals present in ray cells; uniseriate rays storied, fibrous elements
somewhat storied; stem anomalous (dispersed); pith flecks absent.

Passiflora nitida: Growth rings absent; pores solitary, rounded,
average diameter 138 4, range 104 u-157 u; average vessel element
length 234 u, range 219 u-447 p; vessel wall thickness 2 u-4 »; vessel

NAT HERBARIUM

fiddle {

CONTR. U.S. NAT. HERBARIUM, VOL. 34 PART 3, PLATE 2

& ag th {
re t, /

,

lic. 3.— Passifiora coccinea Nubl. ‘Vransverse section of wood showing solitary pores char-
acteristic of Hianous stems in Passifloraceac, & 150.) Fic. 4. Passiflora sp. fonker
Daniels 1227. Transverse section of wood showing tangential pore disposition and

vasicentric parenchyma,» 150.

CONTR. U.S. NAT. HERBARIUM, VOL. 34 PART 3, PLATE 3

Pic. 5.—Passiflora sp. Jonker S Daniels 1227. ‘Vransverse section of wood showing mainly

solitary pores, short apotracheal bands and vasicentric axial parenchyma, 130.0 Vie.
6 —Crossostemma laurtfolium Planch. ‘Transverse seetion of wood with vasieentric
parenchyma arrangement, and s litary, rounded pores, >< 150.

720 982 —64

CONTR U.S. NAT. HERBARIUM, VOL. 34 PART 3. PLATE

CONTR. US NAT HERBARIUM, VOL. 34 PART 3, PLATE 5

Mic. 9--Passiflura arborea Spreng. ‘Transverse section of wood with radial pore multiple
and several kinds of axial parenchyma arrangements: diffuse, ditluse-in-aggregates with a
tendency toward banded, and vasicentric, K 140.) Fie, 10—Passtflora arborea Spreng.
Tangential section of wood showing biseriate heterocellular vascular rays with long

uniseriate wings. Libriform wood fibers have thick walls, < 200.

3 FLATE

PAR TI

34

VO

1M.

NAT HERBAE

US

CON TER.

sechion

rontial

SHOW The

CONTR. US NAT. HERBARIUM. Vol 34 PART 3, PLATE 7

Big. b4.—Passiflora mollisstina (H. B. KL) Bailey. Tangential section of wood illustrating
oblique vessel element end walls, >< 180. Fic. 15.—Passiffora elliptica Gardn. Radial
section of wood showing perforated ray cell, & 900. Fic. 16.—Passifora arborea Spreng

Transverse section of xylem showing a large pith fleck, & 150.

PART 3, PLATE 8

S. NAT. HERBARIUM, VOL. 34

CONTR. U

lnterface

CONTR. U.S. NAT. HERBARIUM, VOL. 34 PART 3. PLATE 9

Pic. L2—Passiflora coccinea Aubl. ‘Uransverse section of stem along xylem-phloem inter-
face as scen in the interrupted type of anomalous development; x =xyvlem, p =phloem,
150.) Bic. 20. —-Paropeia madagascariensis (Baill.) TL. Perrier. “Pransverse section of
xylem ilustrating the occurrence of abundant axial parenchyma in difuse-in-agerevates
arrangement, 150. Bro. 21.—Passiffera arborea Spreng. ‘Vangential section of wood

showing alternate intervascular pitting, >< 900.

CONTR. U.S. NAT. HERBARIUM, VOL. 34 PART 3, PLATE 10

Mic. 22, Passifl ra fuchstiflora Hemsl. ‘Transverse sectl ma of stem st wine hort al Type

of development, & 2S. Wie. 23. -Passtitlora muliitiora I. ‘Vransverse section of stem
illustrating dispersed tvpe of anomalous structure, )

CONTR. U.S. NAT. HERBARIUM, VOL. 34

se &
tee

oo

Vic. 24.—Passiflora glandulosa Cav.

‘Transverse section of stem showing interrupted type
of anomalous structure; stem shape follows xylem outline, 2.8. Bic. 25.—Passiflora
sp. B. Maguire 28327, ‘Vransverse section of stem. illustrating interrupted type of
anomalous structure; stem outline hardly retleets xylem configuration, 3,3

S44
van .

PART 3, PLATE 11

CONTR, U.S. NAT. HERBARIUM, VOL. 34 PART 3, PLATE 12

Fic. 26.—Passifiora glandulosa Cav. Ontogeny of the stem in serial transverse sections:
A, an early internode depicting normal development at this stage; B, a later internode
illustrating incipient formation of furrows in the vascular cylinder; C, D, E, phases
leading to development of steplike configuration which results from unequal production
of secondary xylem and phloem. BA=bark, SP=secondary phloem, SX=secondary

xvlem, PI = pith, P= pore.

cd

AYENSU AND STERN—STEM IN PASSIFLORACEAE 59

element end wall inclination oblique to transverse; intervascular
pitting alternate; imperforate tracheary elements fiber-tracheids,
average length 388 u, range 208 u-482 uw; fibriform vessel members
present; vascular rays mainly uniseriate 1-15 cells high, few multi-
seriate 2-4 cells wide, few very high multiseriate rays, heterocellular;
axial parenchyma diffuse; crystals present in ray cells; uniseriate rays
vaguely storied; stem anomalous (interrupted); pith flecks present.

Passiflora quadriglandulosa: Growth rings absent; pores mainly
solitary, rounded, average diameter 161 yu, range 73 y-207 u; average
vessel element length 283 yu, range 203 u-417 uw; vessel wall thickness
2 u-5 uw; perforation plates simple; vessel element end wall inclination
oblique to transverse; intervascular pitting alternate; imperforate
tracheary elements fiber-tracheids, average length 420 y, range 374 p-
513 uw; fibriform vessel members present; vascular rays uniseriate 1-15
cells high, and multiseriate 2-many cells wide and 2—many cells high,
heterocellular; axial parenchyma diffuse; crystals absent; uniseriate
rays storied; stem anomalous (interrupted); pith flecks present.

Passiflora securiclata: Growth rings absent; pores 80 percent
solitary, 20 percent radial multiples, rounded, average diameter 125 y,
range 101 »-150 u; average vessel element length 268 y, range 217 p—
301 uw; vessel wall thickness 6 u-7 u; perforation plates simple; vessel
element end wall inclination transverse; intervascular pitting opposite;
imperforate tracheary elements fiber-tracheids, average length 461 y,
range 328 y—424 u; fibriform vessel members present; vascular rays
mainly multiseriate without uniseriate extensions, 2-14 cells wide,
8-16 cells high, few uniseriate 1-8 cells high, mostly 3-6 cells high,
heterocellular; ray cells with simple perforations; axial parenchyma
diffuse, and with a few narrow bands; crystals present in ray cells;
storying absent; stem normal; pith flecks present.

Passiflora vitifolia: Growth rings absent; pores mainly solitary,
rounded, average diameter 160 u, range 104 u-196 uw; average vessel
element length 372 yp, range 214 y-472 uw; vessel wall thickness 3 p-
5 u; perforation plates simple; vessel element end wall inclination
transverse; intervascular pitting alternate; imperforate tracheary
elements fiber-tracheids, average length 455 u, range 411 u-549 u;
fibriform vessel members present; vascular rays multiseriate, 2-many
cells wide and few uniseriate 1-8 cells high, few very high multiseriate
rays, heterocellular; ray cells with simple perforations; axial paren-
chyma diffuse; few crystals present in ray cells; storying absent; stem
anomalous (interrupted) ; pith flecks present.

Passiflora sp. Jonker & Daniels 1227: Growth rings absent; pores
50 percent solitary, 30 percent radial multiples, 20 percent clusters,
rounded, average diameter 125 yw, range 94 u-178 uw; average vessel

720-982—64—_4

60 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

element length 344 y, range 311 u-433 yw; vessel wall thickness 2 u—4 yu;
perforation plates simple; vessel element end wall inclination trans-
verse; intervascular pitting opposite to alternate; imperforate tra-
cheary elements fiber-tracheids, average length 487 , range 431 p-
571 uw; fibriform vessel members present; vascular rays mainly multi-
seriate 2-4 cells wide, few uniseriate 1-9 cells high, some very high
multiseriate rays, homocellular, consisting solely of upright cells;
axial parenchyma diffuse with a few narrow bands; crystals absent;
uniseriate rays storied; stem apparently anomalous (included phloem) ;
pith flecks present.

Smeathmannia laevigata: Growth rings absent; pores mainly in
radial multiples, rounded, average diameter 84 yw, range 56 w-101 y;
average vessel element length 570 u, range 466 y-691 uy; vessel wall
thickness 2 u-3 u, perforation plates simple; vessel element end wall
inclination oblique; intervascular pitting alternate; imperforate
tracheary elements fiber-tracheids and libriform wood fibers, average
length 472 », range 422 y-801 yu; fibriform vessel members absent;
vascular rays mainly uniseriate 1-9 cells high, few biseriate 4-8 cells
high, heterocellular; axial parenchyma diffuse; crystals absent;
storying absent; stem normal; pith flecks absent.

Smeathmannia pubescens: Growth rings vague; pores mainly in
radial multiples, rounded, average diameter 87 yw, range 62 u-113 yp;
average vessel element length 590 », range 520 u-674 uw; vessel wall
thickness 2 u-3 y; perforation plates simple; vessel element end wall
inclination oblique; intervascular pitting alternate; imperforate tra-
cheary elements mostly libriform wood fibers, few fiber-tracheids,
average length 431 4, range 410 u-792 yu; fibriform vessel members
absent; vascular rays mainly multiseriate without uniseriate exten-
sions, 2-3 cells wide, 3-8 cells high, some uniseriate 8-13 cells high,
heterocellular; axial parenchyma diffuse, and in narrow bands; crys-
tals present in ray cells; storying absent; stem normal; pith flecks
absent; gummy deposits present in ray cells and vessel elements.

Ontogeny of the Stem

Ontogenetic investigations are based on series of transverse sections
from the stems of four species of Passiflora: P. auriculata, P. fuchsw-
flora, P. glandulosa, and P. multiflora. Prepared slides of the last
species were utilized from the earlier study by Stern and Brizicky.

Passiflora fuchsiiflora: Ontogenetic series in this species illustrate
only NorMAL (fig. 22) stem development, even in the very mature
portions.

Passiflora glandulosa: Transverse sections of the youngest portions
of the stem are quite normal and consist of concentric cylinders of

AYENSU AND STERN—STEM IN PASSIFLORACEAE 61

secondary xylem, vascular cambium, and secondary phloem around a
central pith (fig. 26A). The vascular cambiumi, in these early stages,
produces xylem and phloem in the usual fashion. At the fifth or sixth
node from the tip of the stem, the vascular cylinder becomes furrowed ;
four, five, or six such furrows occur around the stem as a result of
unequal production of xylem and phloem by the vascular cambium,
more phloem than xylem being produced in the furrows (fig. 26B).
As the stem ages and expands, tangential fragments of the vascular
cambium remain behind to form steps at the margins of the furrows
between the xylem and phloem (figs. 26C-E). These fragments of
cambium along the margins of the furrows continue to produce
secondary xylem and phloem in such a manner that a steplike config-
uration (fig. 26E) is produced as in some species of Bignonia (Schenck
1893). An interface between xylem and phloem as it occurs along
the margin of a furrow is illustrated in figures 18 and 19. This stem
configuration (figs. 24, 25) is designated the ivrerRUPTED type, after
Pfeiffer (1926, 1927).

Passiflora multiflora: A detailed ontogenetic study of the stem of
this species was made by Stern and Brizicky. Essentially, the xylem
cylinder is quite normal when young, as is the case in the other species
studied. However, as growth of the stem parenchyma cells proceeds,
the tracheary elements of xylem and phloem become separated into
irregular masses by the dilation of the thin-walled pith and cortical
cells, and cells of axial and radial xylem parenchyma. Each tracheary
mass appears to be associated with a fragment of vascular cambium,
and as a result of this irregularity the production of xylem and phloem
proceeds in different directions and at different rates. The resultant
anomalous developmental pattern (fig. 23) resembles that which occurs
in some species of Bauhinia (Schenck 1893). The prspersup natureof
xylem and phloem strands results from the rupturing of the vascular
cambium by parenchyma cells, and the subsequent action of the dis-
oriented bits of vascular cambium (Pfeiffer 1926, 1927; Stern and
Brizicky 1958).

Passiflora auriculata: This species is generally normal in its ontog-
eny. In the mature portions of the stem, however, isolated phloem
strands are formed in the secondary xylem which appear as patches
on the transverse section. The production of IncLUDED phloem is
quite erratic and phloem islands often appear radially oriented within
the xylem mass. Phloem islands occur when the cambium, sporadi-
cally and briefly, produces phloem centripetally, followed by longer
periods of normal xylem production. This type of anomalous develop-
ment corresponds to the “foraminate type” of Pfeiffer (1926, 1927).

62 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM
Discussion

Any interpretation of xylem anatomy and taxonomy in Passiflo-
raceae must take into consideration the interrelationships between
microscopic structure and plant form. That anatomical structure
may be especially modified in response to habit has been clearly
pointed out by Bailey (1957) and described in certain tribes of Com-
positae by Carlquist (1959, 1960). Habit-related anatomical modi-
fications may be superimposed upon specializations related to
phylogeny, and the separation of these two facets of structural change
is not always clear cut.

The Passifloraceae under study are either lianas or trees. Lianas
are specialized plants among dicotyledons and have developed certain
easily seen external modifications in response to this habit, such as
tendrils and recurved hooks. The stem is attenuated; that is, narrow
in transverse section in relation to the extreme length of the plant.
In their study of the water-conducting system in climbers, Wester-
maicr and Ambronn (1881) asserted that this feature represented an
effective impediment to the flow of water in the stem. They argued
that the relatively small cross section of stem available for the onto-
genetic deposition of conducting elements and the great length of the
conducting region were related to the anatomical peculiarities often
evident in lianous stems. To counteract these morphological ob-
stacles, anatomical development has been modified accordingly. Thus,
in the case of vessels, a serious hindrance to rapid flow is the ad-
hesion of water to the vessel walls. Broadening of vessels and
elimination of scalariform perforation plates are two anatomical modi-
fications which would tend to offset this limitation. A groundmass of
thin-walled, conductive tracheids, as opposed to thick-walled, sup-
portive libriform wood fibers, would also act to increase the area
available for water transport.

Table 2 contains data from which a number of interesting cor-
relations between anatomical structure and habit may be drawn.
Without exception, all trees have libriform wood fibers as well as
fiber-tracheids; none of the lianous species possesses libriform wood
fibers, and the groundmass of the xylem consists of fiber-tracheids
and tracheids. Fibriform vessel members occur in the lianous species
of Passiflora and in Paropsia schliebenit, a tree. Noteworthy in this
instance, is that these peculiarly modified cells have not been seer in
the rare Passiflora maguirei, a vine, nor do they occur in the lianous
species of Crossostemma, Dilkea, or Mitostemma which were available
to us. All lianous species are characterized by predominantly solitary
pores; only among certain tree species are the pores mostly in groups.
An exception is Passiflora elliptica. The mean diameter of pores in

AYENSU AND STERN—STEM IN PASSIFLORACEAE 63

TaBLE 2—Salient anatomical characteristics in the stems of Passifloraceae

[+ = present; — = absent; + = more or less]

3 g » Vessels
= }SS 2
8 |83 Pores 3
3 TEs] é End wall a
S |oe} 43 ~ angle %
& |e3]§ Distribution | § s
Species s |SSi 8 g Zs
3 i se 3 s 3 > |B > M 3 8
S|SzISS/ SE |e st IsJje |§ Is |S] es
Syoe<lsel] & a EVise AS gy] F = S o awd
BlSssss| 2S [Ses LssSelsele [8 Elo | &
S |SS\ES| E/S [ES|SESs/ce/esiSsiss| = |S] 3
o | [= [S3/Es| = esis eigsisclesssiesice| S] S| 2
Pls] 8 ($s/88| 8 [SS SP Ss PSSSs8lSslSslSs] so] SF] &
BIR (A SPR IS ATR TRUITT IE a] a |S
Crossostemma laurifolium +/+ )+ — | 198) + 424) + ~A)—]+
Dilkea johannesil +/—-/]+ — | 205) + 378] + zl]—-|]-
Mitostemma glaziovii +)—-/+ — | 112) + 416 -|j|-|-
Paropsia adenostegia + - +}—]| 50) + 450 +)—-—-]-{-
Paropsia madagas- + + 85 + | 468 + -|-|j-
cariensis
Paropsia schliebenti + _ + 45 + | 590 + +/-—|-
Paropsia vareciformis + - +i]- 7 + | 480 zAl-|—-
Passiflora arborea + - + | — | 130 + 343 + —-|-|-
Passiflora auriculata +/+ )+ + | 210) + 362 +)/—|]+
Passiflora coccinea +)/+/4+ + | 210) + 385 + +)/—-]J—
Passiflora edulis +);+i/4+ + | 181) + 185 +;/-J]t+
Passiflora elliptica +)—]+ +] 117 + | 316 + +~}—|+
Passiflora fuchsiiflora +)/—-J4+ + | 135) + 396 + +~>/—{+
Passiflora glandulosa +/+ {+ + | 200) + 341 + +/)-|]-—-
Passiflora laurifolia +)/—-/]+ + | 124) + 318 + a~l}]— {+
Passiflora maguirei + +) - 75| + 211 + +/)/-|-
Passiflora mollissima +/—]+ + | 111) + 368 + t~y/+ t+
Passiflora multiflora +/+ ]+ + | 128) + 345} + +}-]-
Passiflora nitida +i +it+ + | 138) + 234 + +tzi-|=
Passiflora quadriglan- +/+/+ + | 161) + 283 + +) -J]-
dulosa
Passiflora securiclata +)/—-—J+ + | 125) + 268} + —i|+ i+
Passiflora vitifolia +i/+y+ + | 160} + 372| + —-|+]-
Passiflora sp. (Jonker +/+/+ + | 125 + 344) + +)—|+
Daniels 1227)
Smeathmannia laevigata || + - +)]- 84 + | 570 + —-|-
Smeath mannia pube- + =_ +|-— | 87 + | 590 +/-|];-|-
scens

the lianous species of Passifloraceae is 148 u, species averages ranging
from 75 y-210 w. On the other hand, pores average 78 uv in diameter
in the trees, the species averages ranging from 45 u-130 4. Two
points should be raised in this connection: Passiflora arborea, an
arboreal species, has an average pore diameter which exceeds that in
some of the lianous species, and Passiflora maguiret, a lianous species,
has an average pore diameter less than that in some of the arboreal
species. Vessel element lengths also appear to be correlated with
habit; that is, average vessel element length in tree species is longer
(498; range 343 u-590,) than in lianous species (330 4; range 185 u—
424). Strictly transverse vessel element end walls occur only in

64 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

certain lianous species, and strictly oblique vessel element end walls
are confined to certain arboreal species. Most Passifloraceae exhibit
both types of end walls. The only case of scalariform perforation
plates is found in Paropsia adenostegia, a tree. Two further correla-
tions may be noted: Opposite intervascular pitting and anomalous
stem structure have been seen only in lianous species.

Overlooking the few obvious exceptions mentioned above, some
important generalizations can be stated concerning stem anatomy in
Passifloraceae, Vines have shorter vessel elements and broader ves-
sels which are solitary in distribution. Furthermore, they possess
fiber-tracheids and tracheids, along with fibriform vessel members.
Trees, on the other hand, have longer vessel elements and narrower
vessels which are inclined to be grouped in distribution; fibriform
vessel members are absent. These generalized correlations of ana-
tomical structure with habit can be interpreted reasonably only on a
physiological, rather than on a phylogenetic, basis. Considered in
conjunction with previous statements on the water-conducting
system in lianas, and the special anatomical problems associated with
these attenuated stems, it is easy to see that the xylem anatomy of
these vines is particularly modified to increase the flow of water.
Besides the evident facilitation of this process by the broad vessels
composed of short elements, it is altogether likely that the fiber-
tracheids, tracheids, and fibriform vessel members play an active
part in support of this function. These anatomical modifications,
coupled with others mentioned above, are doubtlessly related to water
conduction in the lianous species. It is in the wood structure of the
trees, therefore, that a search must be made for those anatomical
modifications related to phylogenetic specialization in Passifloraceae.

The most reliable indicators of evolutionary trends in the xylem
of dicotyledons are the development and subsequent specialization of
vessels (Bailey 1957). Long, attenuated, angular vessel elements
with many-barred scalariform perforation plates and scalariform in-
tervascular pitting most closely resemble tracheids, and on this basis
are regarded as most primitive. Short, truncated, rounded vessel
elements with simple perforation plates and alternate inter-
vascular pitting are least like tracheids and are considered most
advanced. Correlations between trends of vessel specialization and
concurrent modification of associated tissues have been employed in
reaching conclusions relative to the phylogenetic development of
imperforate tracheary elements, and radial and axial parenchyma, for
example.

The totality of evidence from the xylem of arborescent Passi-
floraceae points to attainment of a moderate stage of phylogenetic
specialization by these plants. Vessel elements average about

AYENSU AND STERN—STEM IN PASSIFLORACEAE 65

500 uw in length, less than the 649 w average for 600 dicotyledons.?
Vessel diameters average 80 », somewhat less than the 94 uw average
for 1500 dicotyledons. Imperforate tracheary elements are considera-
bly shorter on the average (582 ») than the 1317 » average for 534
dicotyledons. Vessels are rounded in transverse section and pores
are either predominantly grouped, or comprise grouped and solitary
distributions. Intervascular pitting is alternate, and perforations are
simple in all species; however, scalariform perforations also occur in
Paropsia adenostegia and perhaps in other species. Imperforate
tracheary elements are libriform wood fibers and _fiber-tracheids;
some species possess libriform wood fibers exclusively, but none
exhibits solely fiber-tracheids. Vascular rays are largely heterocel-
lular, although in Paropsia schliebenii and P. vareciformis homocel-
lular rays have been recorded. Axial parenchyma is generally diffuse
in nature, but the vasicentric configuration occurs in Passiflora
arborea in conjunction with the diffuse and diffuse-in-aggregates
types. Storied structure is absent in most species. Interestingly,
the two species which show homocellular vascular rays also exhibit a
tendency for storying in the rays. Storying and homocellular rays
are often considered as advanced characteristics. Among the arboreal
Passifloraceae on hand for this study, there does not appear to be
any trend in xylem specialization to indicate an evolutionary series
of any kind. Because Metcalfe and Chalk mention the occurrence
of scalariform intervascular pitting and scalariform perforation
plates in species and genera of which we have no representation
(e.g., Androsiphonia and Barteria), it is likely that with a broader
range of material trends of specialization in the xylem of the arboreal
Passifloraceae would become evident.

Three types of anomalous stem development have been observed
from ontogenetic studies. Material for these studies was available
only in the genus Passiflora, although Crossostemma laurifolium also
exhibits an anomalous stem. The anomalous growth in Passiflora
multiflora is quite distinct, and is caused by the dilation of the paren-
chymatous elements of the xylem which disrupts the cambium,
subsequently to form irregular masses of tracheary tissue. Thin-
walled xylem parenchyma is necessary for the accomplishment of
such a process. Although this type of anomalous development also
occurs in Bauhinia—Leguminosae and in other plants it would
be ludicrous to assume any genetic relationship here. Bailey (1957)
has cautioned, that because ‘‘. . . similarities due to parallel evolution
are surprisingly common in the xylem of dicotyledons” they cannot
be utilized in positive assertions of relationship or phylogenetic

2 Data on general anatomical averages are from appendices in Metcalfe and Chalk (1950).

66 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

derivation without corroborative evidence from other parts of the
plant.

The unique stem in Passiflora multiflora appears to lend credence
to Killip’s original suggestion that the species be segregated into a
genus on the basis of its sessile ovary. Along these lines it might be
well to point out here the occurrence of anomalous stems and storied
rays among several subgenera of Passiflora. These two features,
thus, probably have little taxonomic significance on the species
level.

Stern and Brizicky noted that anomalous stem structure, somewhat
similar to that in Passiflora multiflora, also occurs in Passiflora
coccinea (subgenus Distephana). Present studies indicate, however,
that the anomalous stem growth in Passiflora coccinea results from
tangentially unequal production of secondary xylem and phloem by
the vascular cambium (interrupted type) and not from disrupted and
displaced layers of cambium (dispersed type).

Earlier investigators (Masters 1871, Schenck 18938, Solereder 1908,
Killip 1938, Metcalfe and Chalk 1950, Boureau 1957) may have
overlooked the anomalous stem condition in Passifloraceae because it
ordinarily becomes apparent only in the older portions of the stem
not normally included in herbarium specimens. However, in Passi-
flora glandulosa, the unequal development of xylem and phloem pro-
ceeds very early in ontogeny. The time of attainment of the mature
ontogenetic pattern is not consistent among the species studied, and
it may be strictly genetic, or influenced by edaphic or other environ-
mental stimuli.

From the above descriptions of the ontogeny of the stem in Passi-
floraceae, it appears that the most specialized and highly modified is
the dispersed type of configuration exemplified by Passiflora multiflora.
It might be of interest to speculate on the phylogenetic development
of this type of stem in Passifloraceae, and upon its relationship, if any,
to the other types described here. Reference to figures of this paper,
and to the developmental sequences depicted in the Stern and
Brizicky paper, points to the possibility that the dispersed type of
vascular cylinder may have developed phylogenetically from the
interrupted type by continued distention of cells in the parenchy-
matous groundmass to rupture the vascular cambium completely.
This phenomenon would have to be coupled with, and preceded by,
unequal production of secondary xylem and phloem. The ontogeny
of the stem of P. multiflora shows stages of development representative
of the interrupted type of configuration as seen in P. glandulosa. On
the other hand, the production of included phloem in Passifloraceae
might be viewed as another offshoot of the unequal production of
secondary xylem and phloem. In this case, the unequal production

AYENSU AND STERN—STEM IN PASSIFLORACEAE 67

never became associated with rupturing of the vascular cylinder
brought on by distention of cells in the parenchymatous groundmass,
and the basic integrity of the vascular cylinder was continually
maintained.

In the course of this investigation we observed cells that Woodworth
(1935) termed “fibriform vessel members.” Originally (1934), he
assigned the name “perforated fiber-tracheids” to these same struc-
tures, which perhaps best describes these elements. Woodworth
explained later, however, that “for purposes of definition, this descrip-
tive phrase is unfortunate,” because in effect it means, perforated,
imperforate cells. Basically, these are elongate elements, similar in
most respects to fiber-tracheids, except for the presence of simple
perforations. From the structural standpoint, fibriform vessel mem-
bers resemble fiber-tracheids on the one hand, and vessel elements on
the other. They incorporate within the same cell the greater length
and fusiform shape of the fiber-tracheid with the perforation plates
of vessel elements.

Such a combination of features calls for a reconsideration of the
basic distinction between the two cell types; that is, imperforate
element vs. vessel (perforate) element. According to the [nternational
glossary of terms used in wood anatomy (Committee on Nomenclature
1957), a vessel is an axial series of cells where the cells have coalesced
to form an articulated, tubelike structure of indeterminate length.
This designation implies that a vessel may consist of but two cells
interconnected by perforations; by definition, therefore, two axially
aligned fibriform vessel members must be considered a vessel.

Even though the occurrence of the fibriform vessel member is
greatly restricted taxonomically, and use of the term is correspondingly
limited on this account, the name is inadequately descriptive, and
more importantly, it has misleading phylogenetic implications.
Woodworth was aware of these factors, and considered them in
justifying the use of his term. In every respect these cells resemble
imperforate tracheary elements except for the presence of perforations;
they bear little resemblance to normal vessel elements in the same
wood. Furthermore, the perforation is ordinarily intercalated be-
tween the ends of the cells, and the terminations of these cells have
little if any similarity to the ligulate ends of normal vessel elements
where ligulae occur. The perforation itself is aberrant, usually small
in size, poorly formed, and unlike those in normal vessel elements.
These cells are twice as long as the fusiform cambial initials from
which they have been derived, and differ strongly from normal vessel
elements in this way. It is extremely dubious that these cells are
allied in any way, ontogenetically or phylogenetically, with true vessel
elements. Therefore, to call these cells vessel members of any kind

68 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

would be to relate them, even if in name only, with normal vessel
elements upon whose phylogenetic modification rests our entire
scheme of evolution in the xylem. Because of these factors, we
strongly urge the disuse of fibriform vessel member and the reassump-
tion of Woodworth’s original term—perforated fiber-tracheid—to
describe these cells despite the obvious ambiguity, viz, a tracheid,
by definition, is an imperforate cell.

Although no extensive series of interconnected cells of this type was
seen in the species studied, there are interconnections involving two,
occasionally three, or even five such cells. Axial continuity between
fibriform vessel members and fiber-tracheids also occurs, although no
continuity between fibriform vessel members and true vessel elements
was noted. That there is actual conduction of fluid between con-
tiguous cells of fibriform vessel members and between fibriform vessel
members, fiber-tracheids, and tracheids, is probable in view of the
peculiar water transport problems of lhanas. Woodworth, on the
other hand, remarked that it seems unlikely that the fibriform vessel
members contribute significantly to water movement in such plants.
Regardless, it does seem entirely reasonable to assume that lianous
plants require additional anatomical modifications, such as perfora-
tions in fiber-tracheids, to ensure a more efficient water conducting
system even if the contribution of these modifications to total water
transport is comparatively small. Woodworth examined the xylem
of other lianas for fibriform vessel members and located them only
in some species of Celastraceae. Notwithstanding the apparent
advantage to water conduction in lianas of anatomical modification
which would tend to facilitate this phenomenon, it is obvious that the
genetic potential for such modification must exist as a preliminary
requisite.

The perforated ray cell, which occurs in some species of Passi-
floraceae, has also been observed by Chalk and Chattaway (19383) in
73 species of 16 unrelated plant families. Different specimens of the
same species vary considerably in the number of these cells present.
Perforated ray cells are produced when axially oriented vessels bend
tangentially to pass through one or more cells of a ray. Perforations
in the ray cells usually correspond in kind to the normal perforations
in associated vessel elements. Chalk and Chattaway have suggested
that from the position of the rays, these cells are developed from ray
initials instead of fusiform initials. Since perforated ray cells have
such a wide distribution, and are so variable in occurrence even within
species, their taxonomic significance is limited.

Pith flecks occur commonly in the species of Passiflora; that is,
more regularly than if the presence were merely in response to a
chance trauma. Although the occurrence of pith flecks is commonly

AYENSU AND STERN—STEM IN PASSIFLORACEAE 69

attributed to outside agents, such as the larvae of cambium miners,
their consistent occurrence within the genus Passiflora may have
minor taxonomic significance. Passiflora bark may be especially
attractive to certain organisms which seek it out in preference to other
food sources. The mutualistic relationship between ant species and
myrmecophytes is an example of this kind.

Paropsia, considered previously in this study, has been variously
placed in Flacourtiaceae and Passifloraceae. The wood anatomy of
Paropsia shows considerable similarity to some genera of Passi-
floraceae, especially Smeathmannia, a genus comprising mostly shrubs
and trees. Pores mainly in radial multiples, rounded; perforation
plates simple in some species; vessel element end wall inclination
oblique; alternate intervascular pitting; libriform wood fibers; the
occurrence of abundant axial parenchyma; and fibriform vessel
members, indicate affinity of Paropsia with Passifloraceae. The
occurrence of axial parenchyma in Paropsia is significant, for this
tissue is generally lacking in woods of the Flacourtiaceae (Record and
Hess 1943; Tupper 1934). Pollen grain analysis provides further
substantiation for aligning Paropsia with Passifloraceae. Erdtman
(1952) has shown that pollen of both Passiflora maculifoli and
Paropsia brazzeana has a reticulate sexine and measurements of the
subprolate grains are 524 X 44 and 53 X 41 yn, respectively.
He further suggests that the tribe Paropsieae of Flacourtiaceae should,
at least in part, be referred to Passifloraceae. Convincing resolution
of this problem will depend upon further studies in other botanical
disciplines.

Summary

1. Growth rings are absent or inconspicuously developed; rounded
pores are mainly solitary and in radial multiples; simple perforation
plates and alternate intervascular pitting occur in all species; vessel
element end walls vary from oblique to transverse. Imperforate
tracheary elements are tracheids, fiber-tracheids, and libriform wood
fibers; fibriform vessel members occur in some species. Perforated
ray cells are found in a few species. Vascular rays are both hetero-
cellular and homocellular, uniseriate and multiseriate; uniseriate rays
are commonly storied. Axial parenchyma is diverse in form and
includes a range of types in both apotracheal and paratracheal series.

2. Certain anatomical modifications are correlated with plant form;
lianas have shorter vessel elements and broader, solitary vessels;
fibriform vessel members are present. Trees show longer vessel
elements and narrower, grouped vessels; fibriform vessel members
are generally absent. The anatomical modifications in the xylem

70 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

of lianas arerelated to physiology and habitrather than to phylogenetic
position.

3. Xylem anatomy of tree species indicates that Passifloraceae is
only moderately advanced phylogenetically.

4. Anomalous stem development occurs in Crossostemma and
Passiflora. Ontogenetic series show four types of mature stem
configurations in the family: (1) normal type; (2) included phloem
type; (3) interrupted type; and (4) dispersed type. It is postulated
that the dispersed type of configuration is the most highly modified
anomalous stem form in Passifloraceaec. It is doubtful that the
anomalous stem condition here constitutes a valid taxonomic feature
in assuming genetic relationships between Passifloraceae and other
families, or that its occurrence provides a basis for taxonomic arrange-
ment within the family.

5. The presence of abundant axial parenchyma and of fibriform
vessel members in Paropsia schliebenii supports the position that
Paropsia belongs to Passifloraceae rather than Flacourtiaceae.

6. The substitution of ‘perforated fiber-tracheid” for “fibriform
vessel member” is recommended to avoid possible phylogenetic
misconceptions related to the use of the word vessel in this descriptive
sense.

Literature Cited

Baiey, I. W.
1957. The potentialities and limitations of wood anatomy in the study of
phylogeny and classification of angiosperms. Journ. Arnold Arb.,
vol. 38, pp. 243-254.
BentuaM, G., AND Hooker, J. D.
1867. Passifloreae. Genera plantarum, vol. 1, part 3, pp. 807-816.
Berc_er, L. G. DEN.
1928, Beitrage zur Kenntnis der Anatomie des sekundiren Holzes der
niederlandisch indischen Baumarten. I. Bull. Jard. Bot. Buiten-
zorg, ser. 3, vol. 9, pp. 223-248.
BourgEau, Hi.
1957. Anatomie végétale. Vol. 3, pp. 633-651.
Brizicky, G. K.
1961. The genera of Turneraceae and Passifloraceae in the southeastern
United States. Journ. Arnold Arb., vol. 42, pp. 204-218.
CaRLQUIST, 8.
1959. Wood anatomy of Helenieae (Compositae). Tropical Woods, vol.
111, pp. 19-39.
1960. Wood anatomy of Cichoricae (Compositae). Tropical Woods, vol.
112, pp. 65-91.
Cuax, L., AND Cuatraway, M. M.
1933. Perforated ray cells. Proc. Roy. Soc. London B., vol. 113, pp. 82-92.
1934. Measuring the length of vessel members. Tropical Woods, vol. 40,
pp. 19-26.
1935. Factors affecting dimensional variations of vessel members. Tropical
Woods, vol. 41, pp. 17-37.

AYENSU AND STERN—STEM IN PASSIFLORACEAE 71

Committee on Nomenclature. International Association of Wood Anatomists.
1957. International glossary of terms used in wood anatomy. Tropical
Woods, vol. 107, pp. 1-36.
ERpDTMAN, G.
1952. Pollen morphology and plant taxonomy. Angiosperms. Pp. 178-180.
FERNANDES, A., AND FERNANDES, R.
1958a. Contribuigdo para o conhecimento das Passifloraceae de Mogambique.
Garcia de Orta, vol. 6, pp. 241-262.
1958b. Contribuigéo para o conhecimento das Passifloraceae de Angola.
Garcia de Orta, vol. 6, pp. 649-671.
Guia, E.
1925. Flacourtiaceae. Jn Engler and Prantl, Die natiirlichen Pflanzen-
familien, vol. 21, pp. 377-457.
Harms, H.
1893. Ueber die Verwertung des anatomischen Baues fiir die Umgrenzung
und Ejinteilung der Passifloraceae. Bot. Jahrb., vol. 15, pp. 548-633.
1925. Passifloraceae. Jn Engler and Prantl, Die natiirlichen Pflanzen-
familien, vol. 21, pp. 470-507.
Kip, E. P.
1938. The American species of Passifloraceae. Publ. Field Mus. Nat.
Hist., Bot. Ser., vol. 19, pp. 1-613.
Kriss, D. A.
1935. Salient lines of structural specialization in the wood rays of dicotyledons.
Bot. Gaz., vol. 96, pp. 547-557.
Masters, M. T.
1871. Contributions to the natural history of the Passifloraceae. Trans.
Linn. Soc. London, vol. 27, pp. 593-645.
MetcatFE, C. R., anp CHALK, L.
1950. Anatomy of the dicotyledons. Vol 1, pp. xlix—xl, 674-680.
Oxpaton, M.
1960. Les lianes ligneuses a structure anormale des foréts denses d’ Afrique
Occidentale. Ann. Sci. Nat. Bot., ser. 12, vol. 1, pp. 1-220.
PreIFFER, H.
1926. Das abnorme Dickenwachstum. Jn Linsbauer, Handbuch der
Pflanzenanatomie, vol. 9, pp. 1-268.
1927. Beitrige zur kausalen Begriindung der abnormen Verdickungsschemata
der Pflanzenachsen. Scheininsland Festschrift. Pp. 41-44.
Recorp, S. J., AND Huss, R. W.
1943. Timbers of the New World. P. 171.
ScHENCK, H.
1893. Beitrige zur biologie und anatomie der Lianen. II. Beitrage zur
Anatomie der Lianen. Jn Schimper, Botanische Mittheilungen aus
den Tropen, vol. 5, pp. 1-271.
ScHWENDENER, 9.
1874. Das mechanische princip im anatomischen bau der Monocotylen mit
vergleichenden ausblicken auf die iibrigen Pflanzenklassen. P. 150
SoLEREDER, H.
1908. Systematic anatomy of the dicotyledons (Trans. L. A. Boodle and
F. EK. Fritsch). Vol. 1, pp. 383-388.
Stern, W. L., AND Brizicry, G. K.
1958. The woods and flora of the Florida Keys. Passifloraceae. Tropical
Woods, vol. 109, pp. 45-53.

72 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Stern, W. L., AND Coamsers, K. L.
1960. The citation of wood specimens and herbarium vouchers in anatomical
research. Taxon, vol. 9, pp. 7-13.
Tipro, O.
1941. A list of diagnostic characteristics for descriptions of dicotyledonous
woods. ‘Trans. Ill. Acad., vol. 34, pp. 105-106.
Tupper, W. W.
1934. Preliminary report on the wood structure of the Flacourtiaceae.
Tropical Woods, vol. 38, pp. 11-14.
WEsSTERMAIER, M., AND AMBRONN, H.
1881. Beziehungen zwischen Lebensweise und Struktur der Schling- und
Kletterpflanzen. Flora, vol. 64, pp. 417-430.
Wermors, R. H.
1932. The use of celloidin in botanical technic. Stain. Tech., vol. 7, pp.
37-62.
Woopworts, R. H.
1934. Perforated fiber-tracheids in the passion flowers. Science, vol. 80, pp.
449-450.
1935. Fibriform vessel members in the Passifloraceae. Tropical Woods, vol.
41, pp. 8-16.

INDEX

Index to anatomical descriptions of species

Crossostemma laurifolium, 54, 63
Dilkea johannesii, 54, 63
Mitostemma glaziovii, 54, 63
Paropsia adenostegia, 55, 63
madagascariensis, 55, 63
schliebenii, 55, 63
vareciformis, 55, 63
Passiflora arborea, 56, 63
auriculata, 56, 61, 63
coccinea, 56, 63
edulis, 56, 63
elliptica, 57, 63
fuchsiiflora, 57, 60, 63
glandulosa, 57, 60, 63

Passiflora—Continued
laurifolia, 57, 63
maguirei, 58, 63
mollissima, 58, 63
multiflora, 58, 61, 63
nitida, 58, 63
quadriglandulosa, 59, 63
securiclata, 59, 63
vitifolia, 59, 63

Passiflora sp. Jonker & Daniels 1227,

59, 63

Smeathmannia laevigata, 60, 63

pubescens, 60, 63

73

U.S. GOVERNMENT PRINTING OFFICE:1964

BULLETIN OF THE UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED StatEs NATIONAL HERBARIUM

VotumE 34, Parr 4

TAXONOMIC STUDY OF SUBGENUS
PODOSEMUM AND SECTION EPICAMPES
OF MUHLENBERGIA (GRAMINEAE)

By Tuomas R. SopERstrom

SMITHSONIAN PRESS e WASHINGTON, D.C. e 1967

Contents

Introduction .

Historical considerations .

Leaf anatomy and epidermis .

The embyro .

Lodicules

Chromosomes .

The disposition of Epicampes
Systematic treatment .

Species described as Epicampes but excluded from section Epicampes .

New species and new combination
Voucher specimens examined .
Literature cited

Index .

Tir

Page

75
77
84
97
101
108
110
111
177
178
178
182
187

TAXONOMIC STUDY OF SUBGENUS
PODOSEMUM AND SECTION EPICAMPES
OF MUHLENBERGIA (GRAMINEAE)

By Tuomas R. SopErstrom

Introduction

Within the genus Muhlenbergia (Gramineae) is a group of tall,
rather robust perennials that some botanists have treated as a distinct
genus, H’picampes. Since Presl (1830) proposed Epicampes, many
species have been added. All are found in the Western Hemisphere,
the majority occurring in northern and central Mexico, the center of
distribution.

The characters originally employed to delimit Epicampes have
been interpreted by botanists in different ways. Some have placed
considerable emphasis on the dorsal position of the awn of the lemma,
while others have regarded the relatively long glumes of the spikelet
as particularly characteristic. Another feature of value is the com-
pressed-keeled nature of the basal sheaths. No one character has
been found to distinguish all species of E’picampes from all other
genera. Authors who have described species in the group have had
different concepts of the genus, to the extent that Epicampes has
never been clearly circumscribed. Unfortunately, no recent author
has listed the species considered to comprise the genus Epicampes,
and not only is the generic status and affinities of the group a problem,
there is also no general agreement regarding the number of species.

Traditionally, the classification of grasses has been based on
characters of gross morphology, with special reference to the structure
of the spikelet. In such a classification Epicampes has been treated
as a member of the tribe Agrostideae, which includes grasses with
single-flowered spikelets, with close affinities to such genera as
Agrostis and Cinna.

Starting before the turn of the century, but more particularly
within the last decade, studies of features other than gross morphology
have shed much light on the true relationships within the Gramineae.
These studies have included investigation of the structure of the grass
embryo, lodicules, chromosomes, and the anatomy of the leaf and
epidermis. The results of these investigations are correlated remark-

75

76 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

ably and demonstrate that in many cases the characters of the spikelet
and other features of gross morphology are not reliable indexes to
the natural relationships of grasses. These studies for the most
part have dealt with representative species of genera, in an attempt
to find their natural affinities within the grass family. Relatively
little effort has been made to utilize these studies at the infrageneric
level to clarify relationships among the species within a single genus.

The first phase of the present study was to determine the relation-
ship of the Epicampes group at the genus level, and studies of the
embryo, lodicules, and leaf anatomy and epidermis were useful in
this regard. The results of these investigations provide strong evi-
dence that EHpicampes is not related to such genera as Agrostis and
Cinna of the Agrostideae but rather to grasses of the ‘“chloridoid”’
alliance, which includes many grasses traditionally placed within the
tribe Chlorideae.

The second phase was a taxonomic survey of the species belonging
to the Epicampes group. Field studies were made in Mexico in 1959
and 1960 and herbarium specimens examined included the majority
of type specimens or fragments of types of the species involved.

In addition to the large series of specimens studied at Yale Uni-
versity (YU) and the U.S. National Herbarium (US), collections
were borrowed from the following herbaria: University of Arizona
(ARIZ); California Academy of Sciences (CAS); Escuela Nacional
de Agricultura, Chapingo, Estado de México (CHAP); Stanford
University (DS); Field Museum of Natural History (Ff); Gray
Herbarium of Harvard University (GH); University of Ilinois (ILL);
Instituto Politéenico Nacional, México, D.F. (IPN); Texas Re-
search Foundation (LL); Universidad Nacional de Mexico (MEXU);
University of Michigan (MICH); Missouri Botanical Garden (MO);
New York Botanical Garden (NY); Pomona College (POM); Natur-
historiska Riksmuseum, Stockholm (S); Southern Methodist Uni-
versity (SMU); A.&M. College of Texas (TAKES); University of
Texas (TEX); University of California (UC); University of New
Mexico (UNM); and Instituto Boténico, Ministerio de Agricultura
y Cria, Caracas (VEN); The abbreviations of herbarium names
are those of Lanjouw and Stafleu (Index Herbariorum, ed. 5, 1964)
except for CHAP and IPN which are the author’s.

This paper is based on a doctoral dissertation prepared at Yale
University under the guidance of Dr. John R. Reeder, whose advice,
encouragement, and perseverance will always be an inspiration to
me. To his wife, Charlotte Goodding Reeder, my sincere appreci-
ation for her assistance during the entire study at Yale. The work
was completed at the U.S. National Herbarium and I am most grateful
to Dr. Jason R. Swallen for his advice, assistance, and generosity in

SODERSTROM—PODOSEMUM AND EPICAMPES V7

many ways. Mrs. Susan Colby McKanna of the Smithsonian deserves
very special thanks for giving so much of her time to type, proofread,
and carry out many other necessary tasks connected with the final
writing. Of my colleagues at the Smithsonian, a word of appre-
ciation to Drs. William L. Stern, Mason E. Hale, Floyd A. McClure,
and Mr, Conrad V. Morton, each of whom has been of special help.
To Dr. Frank W. Gould, Texas A & M University, who read the
manuscript before its final revision, I express my sincere thanks for
his constructive criticisms.

Historical Considerations

The description of the genus Epicampes by Jan S. Presl (1830) was
based on a single collection from southern Mexico. In K. B. Presl’s
introduction is the statement (translated): ‘In the month of November
Thaddaeus Haenke went alone to the capital city of Mexico, and
returned in December to Acapulco.” It was on this trip in 1791
that Haenke collected the type specimen of Epicampes. Although
no locality other than “Mexico” is given, Presl’s description and
illustration of the specimen resemble most closely other specimens
of Mpicampes from the central Mexican plateau. Furthermore,
Haenke’s specimen was probably collected higher on the plateau in
the state of Morelos or México, rather than in the lower and more
tropical elevations of Guerrero, of which Acapulco is the main port
of entry on the Pacific.

The name Lpicampes may have been derived from the classical
Greek word “kampes,” an adjective meaning “curved.” It is unclear
why Presl chose a word with such a meaning as a name for this genus.
Unfortunately, he gives no indication of its etymology in the original
description.

The type species, H’picampes strictus Presl (Muhlenbergia robusta
[Fourn.] Hitche.), is widely distributed in the higher elevations
from the state of Jalisco southward into Central America. His
description of the genus, translated from the Latin, follows:

Panicle contracted, in the form of a spike, with alternate solitary branches.
Glumes two, ovate, convex, nerveless, subequal, obtuse, entire. Floret a
little longer than the glumes. Glumes two, convex, ovate, the lemma sur-
rounding the palea at its base, entire, with the median nerve extending into
a Straight awn from beneath the apex, the palea 2-nerved, very obtuse, en-
closing the genitalia. Stamens 3. Ovary ovate, emarginate at apex. Styles
2. Stigmas divided into dense branchlets. Lodicules . . . caryopsis. .. .
It seems to differ enough from Agrostis by the entire lemma and palea,
with a straight awn neither twisted nor plicate, from Apera by the awned
lemma, To this genus belong also Agrostis pubescens and A. lanata.

Of the several features that Presl enumerated for Fpicampes, the
nerveless condition of the glumes and the awn rising from just be-

78 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

low the tip of the lemma have been regarded by subsequent authors
as important diagnostic characters for the genus.

As originally proposed, Epicampes comprised three species. Be-
sides EH’. strictus, Presl included Agrostis lanata H. B. K. and A.
pubescens H. B. K. These species had been described by Humboldt,
Bonpland, and Kunth (1815) from specimens collected by Humboldt
in the state of Guanajuato, north of where Haenke had collected.
Three other species described in the same work, Crypsis macroura,
C. phleoides, and Podosaemum distichophyllum, were later included
in Epicampes by other workers. Thus, the species assigned to
E’picampes had been described under the genera, Agrostis, Crypsis,
and Podosaemum.

Kunth, who originally named the two Humboldt collections from
Guanajuato as species of Agrostis, later accepted Presl’s contention
that they belonged to Epicampes. In 1833, Kunth’s Agrostographia
was published in Germany, where Hpicampes was included in the
Agrostideae, a tribe established by Kunth in 1815. Incorporated
in this tribe were all grasses characterized by single-flowered spikelets:

V. Agrostideae. Spikelets single-flowered, rarely with a rudimentary
subulate flower above the other. Glumes 2, lemma and palea, membrana-
ceous-herbaceous; lower often awned. Stigmas mostly spreading. Through

Muhlenbergia affinities with Stipa.

Although recent research has demonstrated the unnaturalness of the
Agrostideae as interpreted by Kunth, many contemporary taxono-
mists still recognize the tribe and place in it Muhlenbergia, including
the Epicampes group.

Kunth’s description of Hpicampes was similar to Presl’s. He
placed the genus between Cinna L. and Sporobolus Brown, men-
tioning that it ‘differs from Cinna principally by its convex, non-
keeled glumes.” The genus Afuhlenbergia, in which many recent
authors have incorporated Epicampes, evidently was not considered
by Kunth to be as closely allied, for he separates the two by several
genera.

Trinius (1841) recognized E’picampes essentially as Presl and Kunth
had before him but described the species in more detail and added
one new species, FE’. gracilis (Muhlenbergqia lindheimeri Hiteche.). He
described the glumes as 1-nerved, in contrast to Presl’s and Kunth’s
statements that the glumes are nerveless. Yet, in his description of
I. strictus and FE. pubescens, Trinius noted that the glumes are nerve-
less. The new species Trinius described was from Texas, the first of
the group from the United States.

Until 1874 Hpicampes was treated as comprising only four species,
E. strictus, F. lanatus, E. pubescens, and E. gracilis, all with a habit
and panicle similar to the first species that Presl described, and as a

SODERSTROM—PODOSEMUM AND EPICAMPES 79

genus ranging from southwestern United States (Texas) to southern
Mexico. This concept of Hpicampes was expanded by Grisebach
(1874) in a description and discussion of the collections of Argentine
plants of Professor Lorentz. In this work Grisebach described a
new species, E. coeruleus (Muhlenbergia angustata [Presl] Kunth),
with spikelike panicles, 1-nerved, awn-tipped glumes, and a long
acute ligule. At the same time he transferred two other species
with spikelike panicles to Epicampes: Cinna phleoides Kunth (based
on Crypsis phleoides H. B. K.) and C. stricta Kunth (based on Crypsis
stricta H. B. K.). By incorporating these species from Argentina
into Epicampes, the range of the genus was considerably extended.
George Bentham (1881) in the first really critical comments con-
cerning the generic limits of Hpicampes recognized four subtribes
within the Agrostideae, one, the Euagrosteae, included H’picampes.
About 16 genera comprised this subtribe, of which he concluded:
. . . the general character is a dorsal usually twisted awn on the flowering
glume, the grain neither so closely enveloped in the fruiting glume as in
Stipeae, nor so readily exposed as in Sporoboleae, and the spikelets usually
small, loosely paniculate, very rarely condensed into a head as in Phleoideae;
but there are exceptions to every one of these characters, and the limits of
the larger genera are so vague as to render this portion of the genera of
Gramineae the least satisfactory of the whole series.

Bentham stated that Hpicampes was composed of “about 16
species,” ! but that this number was possibly too large and most
probably reducible by one-third, and that it “is a genus most embar-
rassing to the systematist; for it seems to connect Muhlenbergia and
Sporobolus with Agrostis.” One of the characteristics he regarded as
distinctive of Epicampes was the long, narrow, dense panicle composed
of numerous small spikelets. He differentiated Epicampes from
Muhlenbergia by the awns on the lemmas which, when present in the
former, are short and not quite terminal, while in the latter are longer
and terminal. The dorsal position of the awn was one of the chief
distinguishing features recognized by Presl in establishing the genus.
Bentham distinguished L’picampes from Sporobolus by the fact that
in the former the glumes are thinner and more membranous than the
lemma and palea but in the latter they are thicker and less mem-
branous. Bentham admitted his unfamiliarity with many of the

1 A survey of the literature reveals only 9 published species of Epicampes up
to and including those described in Bentham’s paper. For his estimate of 16
species, Bentham doubtless relied on Fournier’s treatment of the group in Mezi-
canas Planitas. Although the title page was not printed until 1886, Fournier had
sent copies to several botanists as early as 1880. In the first part of his ‘‘Notes’’
(p. 20) Bentham stated that Fournier had supplied him with a copy, remarking,
“T feel bound, insofar as I am concerned, to treat it as having already taken date.’’

80 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

published species, adding that, “a further study may require consider-
able modification of the generic character and limits.”

Following the example of Grisebach, Bentham included in Epi-
campes additional species with spikelike panicles. He transferred
Fournier’s Crypsinna and Kunth’s Cinna macroura and C. stricta to
Epicampes macroura (Muhlenbergia macroura [H.B.K.] Hitehce.).
Cinna macroura Thurber was placed in Epicampes to become Epi-
campes rigens Benth. (Muhlenbergia rigens [Benth.] Hitche.), a species
closely related to H. macroura. The genus Bauchea Fourn., repre-
sented by a single Mexican species, B. karwinskyi, which Bentham had
not seen, was also considered by him to be an Epicampes (although
later synonymized with Sporobolus wrightii Munro ex Scribn.).

In Bentham and Hooker (1888) ? four subtribes of the tribe Agros-
tideae are listed, with the genera enumerated under each. The four
subtribes are: Stipeae, Phleoideae, Sporoboleae, and Euagrosteae.
E’picampes was placed within Euagrosteae and Afuhlenbergia in Stipeae.
Sporobolus, considered by some as closely allied to Hpicampes, was
the basis of subtribe Sporoboleae.

Of the four subtribes listed, the Sporoboleae, containing the
single genus Sporobolus, is distinctive in that the grain is not a true
caryopsis but rather the seed is loose from the ovary wall (pericarp).
Furthermore, the fruit is loosely enclosed within the lemma and palea
and drops out easily; for this reason the popular name ‘“dropseed”’
has been applied to the genus. The chief distinction of the Kuagros-
teae appears to be the position of the awn on the lemma, dorsal in
some cases, ‘‘rarely arranged subterminally to a mucro, or variously
awned” (Bentham, 1883). However, grasses with awned lemmas are
also found in the Stipeae and Sporoboleae. The position of the awn
on the lemma in the Euagrosteae does not easily separate this subtribe,
and the Stipeae and Euagrosteae show no distinctive features to
separate them clearly.

According to Bentham’s description of Muhlenbergia (Stipeae), the
major character to differentiate this genus from Hpicampes (Kuagros-
teae) was the relatively small size of the glumes as compared with the
lemma and palea. Yet, even here species are included with the
glumes subequal to the floret. The awn of the lemma in Epicampes
was stated as minute and subterminal, although in Muhlenbergia the
awn was described only as ‘rarely obsolete.’””? There was no reference
to its position on the lemma. Although Bentham gave these con-
trasting characters for the two genera, none was distinctive enough
to separate all species of Hpicampes from all species of Muhlenbergia.

2 The treatment of Gramineae in Genera Plantarum (1883) was produced by
Bentham whose “Notes on the Gramineae’ (1881) was based on observations
made during its preparation.

SODERSTROM—PODOSEMUM AND EPICAMPES 81

Following the lead of previous authors, Vasey (1885) placed
Epicampes in the Agrostideae. Employing the system of Bentham
and Hooker, he included Muhlenbergia in the subtribe Stipeae and
Epicampes in the subtribe Euagrosteae. According to Vasey’s
interpretation, the only essential character that differentiated the
two subtribes was the position of the awn on the lemma. He said of
the Stipeae, “beard of the flowering glume terminal,’ and of the
Euagrosteae, “flowering glume usually with a more or less twisted
dorsal awn; rarely mucronate or awnless.” Like Bentham and Hooker,
he included the genera Agrostis, Polypogon, Arctagrostis, and Cinna
in the same group of Euagrosteae with E’picampes.

In his description of H’picampes, Vasey recognized three species:
E. distichophyllus Vasey, LE. macroura Benth., and FE. rigens Benth.
He stated that the spikelets of Epicampes are much like Sporobolus
but contracted in a long, narrow, and dense panicle. He added,

.. . Outer glumes somewhat unequal, membranaceous, convex on the back,

scarcely keeled, obtuse, three-nerved; flowering glume mostly equalling the

outer ones, sometimes three to five nerved, entire or sometimes awned from

the apex; palet hyaline, about equalling the flowering glume, two nerved or

two-keeled.
Vasey seems to have considered long dense panicles and glumes a
little longer than the lemma as the important features of Hpicampes.
He erred, however, in stating that the glumes are 3-nerved; previous
agrostologists had emphasized the nerveless or l-nerved condition
of the glumes in species of this group. He also said that the lemmas
(“flowering glumes’’) are entire or awned from the apex, but did not
recognize the dorsal position of the awn as a diagnostic feature.
Nevertheless, in E. distichophyllus Vasey (M. emersleyt Vasey) the
awn is clearly dorsal, or subterminal, rather than terminal.

Hemsley received a copy of Fournier’s manuscript of Mezicanas
Plantas before its publication and drew upon it in Biologia Centrali-
Americana (1885). In the introduction, Hemsley remarked, “with
regard to many of the new species described by Fournier which we
have had an opportunity of examining, we consider them as varieties
undeserving of distinctive names even.” Hemsley’s treatment in-
cluded many of Fournier’s species, and he comments that of the 16
species proposed up to that time, several were not distinguishable as
such. The genus /picampes followed Sporobolus in his treatment, in
accordance with contemporary taxonomic opinion. Both Hemsley’s
and Fournier’s works merely contain listings of genera and species,
with a brief description of each species but no descriptions of the
genera. Fournier and Hemsley followed the treatment of Gramineae
as proposed earlier by Bentham (1881).

82 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Hackel (1890) in The True Grasses (a translation by F. Lamson
Scribner and E. Southworth) also followed Bentham and _ placed
Epicampes in the Euagrosteae. Hackel mentioned 12 species in con-
trast to the 13 recognized by Fournier.

Vasey (1892) listed two species of Hpicampes, E. ligulatus, described
as new, and I. rigens Benth. (Muhlenbergia rigens [Benth.] Hitchc.).
One species that Vasey included in the genus Muhlenbergia, M.
emersleyi Vasey? was later transferred to Epicampes by A. S. Hitch-
cock. Vasey’s concept of the generic limits of Zpicampes did not fully
coincide with the general established concept, for he certainly would
have included M. emersleyt within E’picampes if he had been familiar
with the characters used to delimit the genus. Except for the glabrous
lemmas of FE. ligulatus and E. rigens, no characters would exclude them
from his concept of Muhlenbergia—but, even then, under Muhlen-
bergia he remarked that the flowering glumes, or lemmas, are “fre-
quently” pubescent below. Although Vasey recognized Muhlenberqia
and Epicampes, he failed to select good characters to distinguish these
two genera.

A key to nine species of Epicampes was given by Beal in 1896.
In his discussion, Beal remarked that all species of Hpicampes seemed
near Agrostis, but some closest to Cinna, others to Muhlenbergia, and
yet others to Sporobolus. He stated:

. the chief general feature is the long narrow dense panicle with very
numerous rather small spikelets, the awn of the floral glume, when it exists,
much smaller than in Muhlenbergia and often not quite terminal; the unawned
species are distinguished from Sporobolus by the fruiting glume and grain
which are nearly those of Agrostis.

The notes on Epicampes by Marcus Jones (1912) point up the
confusion that had developed in establishing the boundaries for
Epicampes. Jones’ conclusions, based solely on morphological
observations, are corroborated to a large extent by anatomical studies
made by the present author. Jones considered as the most important
character of Hpicampes the specialized awn of the lemma that arises
from just below the tip, that is, dorsally or from an emarginate tip.

Although Jones erred in associating Bealia mexicana Scribn.
(Muhlenbergia biloba Hitche.), M. argentea Vasey, and M. clomena
Trin. with Hpicampes, he did recognize that certain other species
associated with H’picampes belong to a separate group. This group
includes the species with spikelike panicles, coarse ribbed leaves, and
firm ligules. Jones followed Fournier in assigning such species to
the genus Crypsinna. He also included Epicampes macroura Benth.

8 Vasey also listed another Muhlenbergia, M. distichophylla Kunth. The speci-
men referred to, however, was M. emersleyi and is not to be confused with M,
distichophylla of Mexico and Guatemala.

SODERSTROM—PODOSEMUM AND EPICAMPES 83

in the genus, as Fournier had done, and further considered E. rigens
Benth. to be a member.

Fournier originally established the genus Crypsinna based in part
upon Crypsis H. B. K. and in part on Cinna Kunth (not L.), and in-
cluded species with 1-flowered spikelets and dense spikelike panicles.
Bentham chose to reduce Crypsinna to Epicampes, but Jones accepted
Fournier’s genus and did not recognize those species with spikelike
panicles as belonging to Hpicampes.

Hitchcock in 1914 and in earlier works recognized the genus
Epicampes, although later he united it with Muhlenbergia. His
description of Muhlenbergia indicated that ‘it grades on the one hand
into Sporobolus, from which it differs in having an awned or mucronate
lemma, and on the other into E’picampes, from which it differs in
having a relatively firmer lemma.”’ Bentham used the firm lemma
in Hpicampes as a character in contrast with the thinner lemmas of
Sporobolus.

Hitchcock (1920) based his description of Hpicampes on the
characters of the equal glumes, the awn of the lemma which, when
present, characteristically rises from just below the tip, the caespitose,
perennial habit of the plants, and the open, narrow, or spikelike
panicles. Hitchcock regarded Crypsinna as a synonym of Mpicampes.
He also transferred Muhlenbergia emersleyi Vasey and M. vaseyana
Scribn. (M. distichophylla sensu Vasey) to EHpicampes, bringing the
number of species to 15.

Bews (1929) regarded the texture of the lemmas as a significant
character in differentiating genera within the Agrostideae. Under
the first category of the tribe, “lemma usually hyaline or mem-
branaceous at maturity, at least as delicate as the glumes, or more
so,” he included Sporobolus, Agrostis, and Epicampes, in addition to
many other genera. MJuhlenbergia is found under the second category,
“lemma indurated at maturity, or at least firmer than the glumes.”
Bews listed the same species of Hpicampes that Hitchcock had listed
in his 1920 publication.

A significant change was made in the disposition of Hpicampes by
Hitchcock in 1934. In transferring Epicampes ligulatus Scribn. to
Muhlenbergia (M. longiligula Hitchc.), he commented:

. . . the character which I have used to differentiate the two, the awn from
the back of the lemma, just below the apex in EHpicampes and from the tip in
Muhlenbergia, has several exceptions in the latter genus. Since it is impos-
sible to find a clear line of division to separate the genera, I am uniting
Epicampes with Muhlenbergia.

Hitchcock (1935) contributed a complete treatment of all North
American species of Muhlenbergia, in which he transferred all species

$4 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

of Epicampes to Muhlenbergia. is disposition of the group has been
followed subsequently by most American taxonomists.

The inclusion of Epicampes within Muhlenbergia, however, has
not been universally accepted. The most recent system of classifi-
cation of the grasses treating all known genera was that of Robert
Pilger in 1954, with a more complete form in 1956. Pilger placed
Epicampes in the tribe Eragrosteae which he divided into six sub-
tribes. Subtribe 5, Sporobolinae, included not only Sporobolus, but
also E’picampes, Crypsis, Heleochloa, Urochondra, Sphaerocaryum, and
Blepharoneuron. Muhlenbergia, excluding the Epicampes species,
formed subtribe 6, the Muhlenbergiinae. A translation of the section
of his key separating the Sporobolinae and Muhlenbergiinae follows:

I. Spikelets often broad; glumes thin; lemmas awnless; caryopsis more or less
wide 2... .. eee et ee ee ee es Subtribe 5. SPOROBOLINAE
II. Spikelets small; glumes firm; glumes often more or less shorter than the
lemmas, these often more or less long-awned; caryopsis small, cylindrical.
Subtribe 6. MUHLENBERGIINAE
Pilger did not enumerate the species he recognized under E’picampes,
but most of the species traditionally regarded as members of this
genus possess awned lemmas. The type specimen of the genus, £.
strictus Presl, is figured with short-awned lemmas. Moreover, since
Presl established Hpicampes the position of the awn on the lemmas
has been used as a generic character of great importance. It is
therefore surprising that Pilger should have placed Hpicampes in a
tribe composed of members with awnless lemmas.

Hubbard, in his treatment of Gramineae (1959, rev.), recognized
Epicampes. Sporobolus, often regarded as showing a relationship
with Hpicampes, was placed in the tribe Sporoboleae, along with
Blepharoneuron. Muhlenbergia and Epicampes, in addition to Agros-
tis, Calamagrostis, Lagurus, Gastridium, Triplachne, L[chinopogon,
Dichelachne, Polypogon, Phleum, and Alopecurus, among others, con-
stituted the tribe Agrosteae.

It is apparent that Hpicampes never has been clearly understood.
Different characters have been used by botanists to delineate the
genus and few of their descriptions fully agree. The American, A. 5.
Hitchcock, incorporated the species of Epicampes into the genus
Muhlenbergia, and since then most American taxonomists have ac-
cepted his treatment.

‘Leaf Anatomy and Epidermis

The first suggestion that the anatomy of grass leaves might be
useful in systematics was made by Duval-Jouve (1875) who found
differences in the distribution of bulliform cells among species of
different tribes. Additional anatomical differences were pointed out
by Schwendener (1890). In 1898, Pée-Laby recognized five groups

SODERSTROM—PODOSEMUM AND EPICAMPES 85

based on characters of leaf anatomy, although these groups were based
on poor characters (Brown, 1958). Avdulov’s system (1931) incorpo-
rated the characters of leaf anatomy and chromosome size and number.
Prat (1936) found that characters of the leaf epidermis are useful
from a systematic standpoint and that the results based on these
characters correlate well with those derived from a study of the
internal anatomy. He referred to Avdulov’s major groups as
“nanicoid” and “festucoid” and segregated the former into two
subgroups, the ‘“eupanicoid” and ‘“‘chloridoid.”

Brown (1958), in a study of the leaf anatomy of 101 species in 72
genera, found that six groups can be delimited by characters of the
internal anatomy. Various combinations of these characters make
it possible to differentiate the six basic groups that Brown designated
bambusoid, festucoid, arundoid, panicoid, aristidoid, and chloridoid.

Tateoka et al. (1959) surveyed the leaf epidermis of 238 species of
grasses and found that two groups, which differ in the type of
microhairs present, correspond to the panicoid and chloridoid groups
of Prat. They also reported that the bicellular microhairs of several
genera have distinctive features which suggest that leaf epidermal
characters may be useful at the generic level.

The most detailed survey of leaf anatomy and epidermis of grasses
was made by Metcalfe (1960). He compiled the diagnostic features
of the leaf epidermis and internal anatomy of 206 genera, representing
413 species. In addition, he included information on numerous other
species, summarized from the literature. The survey by Decker
(1964) of 135 genera traditionally placed in the tribe Festuceae
demonstrates the value of leaf anatomy in reinterpreting a large,
long-recognized tribe.

In the present study, the leaf anatomy and epidermis were studied
by the author in about 50 species of Muhlenbergia, including those
traditionally placed within Epicampes. About 50 additional species
of Muhlenbergia were studied by Reeder at Yale University, and the
author was fortunate to have the results of Reeder’s unpublished
studies for comparison with his own findings.

LEAF ANATOMY

Comparative studies of leaf anatomy were made from transverse
sections of leaves of herbarium specimens. Mature blades were
selected from about the middle of the culm and sections taken from a
point in the blade approximately 1 cm. from the ligule (at this point
the vascular bundles and associated structures are well developed;
more distally the blades become narrower and certain features become
altered). The dry blade was placed in boiling water for several
minutes to soften and to bring it back as much as possible to the

86 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

pliable condition of a fresh leaf. In some cases the blades were
soaked in a 0.3-0.4 percent solution of trisodium phosphate in a 60°
paraffin oven for 1%-2 hours, although equally good results were
obtainable without this additional treatment.

Freehand sections were made in all cases. The leaves were
heavily sclerified and their rigidity permitted thin sections to be
made by hand, thus obviating the use of a microtome. The blade
was placed between halves of a block of celloidon and sections cut
with a sharp single-edged razor blade. The sections were stained
with Chlorazol Black E for % hour and then washed and brought
through the alcohol series to absolute ethyl alcohol, and made perma-
nent by mounting in diaphane.

Muhlenbergia species traditionally treated as Epicampes possess
panicles that are either spikelike or open and pyramidal or columnar.
The leaves in the former group become inrolled upon drying (M/.
rigens); in the latter group the leaves become folded upon drying (\/.
gigantea). Anatomical studies of the leaves of those two species
reveal striking structural differences and the leaves of the other
species of the Epicampes group are of one type or the other. For
purposes of discussion, species with the leaf anatomy of MV. rigens are
referred to as having a “Rigens” type, and species with a leaf anatomy
of M. gigantea are referred to as having a “Gigantea’’ type.

Plates 4-14 contain photomicrographs of leaf cross sections of a
number of species of Muhlenbergia, including those of the Epicampes
group. <A partial leaf cross section of Af. rigens appears in plate 5,
figures 1 and 2, and of AZ. gigantea in plate 7, figure 2. The anatomical
features of Epicampes leaves are illustrated on plate 4 in cross sections
of M. lindheimeri and M. palmeri leaves. These two species have
a leaf anatomy that corresponds, respectively, to that of Mf. gigantea
and M. rigens.

Anatomical Structure of Leaves of the Epicampes Group 4

Units: The leaf consists of a series of units of varying. sizes,
each unit representing a single rib of the leaf. The individual unit
contains a vascular bundle and associated structures between and in-
cluding the upper and lower epidermis. On plate 4, figure 2 are

4The author is grateful to Dr. C. R. Metealfe of the Roval Botanic Gardens,
Kew, for his aid in interpreting the anatomical structures of the cross section of
the leaf of M. lindheimeri from the photomicrograph sent to him.

5T am employing the term ‘‘unit’’ as used by Brown (1958, p. 176) to describe
leaves of chloridoid grasses—“‘In these groups each bundle, sheath, and associated
radial chlorenchyma constitute a diserete unit of structure separated from the
similar adjacent units by large, empty, bulliform cells’’—but I consider also the
upper and lower epidermis and include sclerenchyma as part of this unit. The
words “primary,’’ “secondary,’”’ and ‘tertiary’? to distinguish units are mine.

SODERSTROM—PODOSEMUM AND EPICAMPES 87

shown four units of a blade of Muhlenbergia lindheimeri and below,
figure 3, seven units of a blade of M. palmeri.

Units are designated primary (1°), secondary (2°), and tertiary
(3°), from largest to smallest. In the Gigantea type, only 1° and 2°
units are found. The median unit is heavily sclerified at the base.
This 1° unit and adjacent units are “embedded” in a mass of paren-
chyma, the whole arrangement constituting the ‘‘midvein area,” or
keel. The keel structure is shown on plate 4, figure 1. The 1° units
alternate with one to three 2° units throughout the leaf. In the
Rigens type, 3° units often are present, in addition to 1° and 2° units.
As seen in M. palmeri one 2° and two 3° units occur between the two
1° units. Such an alternation of units occurs throughout the leaf
and the central units are not specialized into a keel.

VASCULAR BUNDLES: The phloem of the 1° vascular bundles of
M. lindhewmert and M. palmeri appears as darkly-stained groups of
cells among nonstained thick-walled cells. All of the phloem is
darkly stained in the 2° and 8° bundles of these species. The phloem
of the 1° bundles is sclerosed, i.e., partly converted to lignified fibers.
In cross section the active parts of the phloem (sieve tubes and
parenchyma) appear as darkly-stained strands embedded in a fibrous
ground tissue that is not stained. Sclerosis of the phloem is less
pronounced, or lacking, in the 2° and 3° bundles.

The xylem of the 1° bundles is characterized by large metaxylem
vessels on either side of the protoxylem. (The line marking the
xylem in the photomicrograph of M. palmeri, plate 4, points to the
metaxylem.) These large metaxylem vessels are lacking in the 2°
and 3° bundles; in these bundles both the xylem and phloem stain
black and are indistinguishable in the photomicrographs.

The 1° units are defined as those having metaxylem vessels, which
are lacking in the 2° and 3° units. In cases where the 1° and 2°
units are about equal in size, they are distinguished on the basis of
whether or not they possess such metaxylem vessels.

Bunpie sHEatus: Two distinctive layers of cells, or ‘“sheaths,’’
surround the vascular bundle. The innermost, variously termed the
“mestome sheath,” “inner sheath,” or “‘endodermis” is composed of
thick-walled cells. The cells of the inner sheath adjacent to the
xylem are larger; the cells adjacent to the phloem fibers are smaller
and often indistinguishable from the latter.

The inner sheath is surrounded by a single layer of conspicuously
larger, thin-walled cells, termed the “parenchyma sheath” or ‘outer
sheath.” The outer sheath may be continuous around the inner
sheath, or interrupted below, above, or both. In the 1° units of M.
lindheimert the outer sheath is interrupted above and below, but
only above in the 2° units. It is continuous in the 2° and 3° units

221-352-672

88 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

of M. palmeri. (The closed condition is typical of every unit of the
leaf in some species of Muhlenbergia—cf. plate 12, figures 1, 2, and
plate 13, figure 1.)

CuiorencuyMa: The chlorenchyma lies adjacent to the outer
sheath and both are stained black in the photomicrograph. Although
not evident in the photomicrograph, the cells are radially arranged.
The chlorenchyma assumes the same pattern as the outer sheath in
the unit, being continuous around the inner sheath or interrupted.
(Photosynthetic activity is confined to this narrow layer of chlor-
enchyma and, presumably, the outer sheath also; together, they
constitute a relatively small part of the whole unit [Brown: 1958,
p. 176]).

ScLERENCHYMA: The individual units, especially the 1° ones,
possess caps of sclerenchyma cells subjacent to the upper and lower
epidermis. The amount of sclerenchyma varies, but it is especially
well developed at the base of the central unit where it reinforces the
keel of the blade. In a species with the Rigens type of leaf anatomy
the sclerenchyma often forms a continuous layer subjacent to the
lower epidermis.

THICK-WALLED PARENCHYMA: Between the vascular bundle and
subepidermal sclerenchyma is a mass of large parenchymatous cells.
They are referred to here as ‘thick-walled’ parenchyma,® as the
walls of these cells appear to be thicker than in normal parenchymatous
cells. These cells constitute a large part of the 1° units, a smaller
part of the 2° units, and are lacking in the 8° units.

In some cases the thick-walled parenchyma of the units is con-
fluent between them. Such a condition is found in the keel of leaves
of the Gigantea type in which the vascular bundles and surrounding
sheaths give the appearance of being embedded in a mass of paren-
chyma cells. In such cases there is no observable difference between
parenchyma cells of the units and those that occur between them
as they are confluent.

CoLoRLEss CELLS: Between the units are found one to several
rows of cells that look similar to thick-walled parenchyma. Met-
calfe refers to these as colorless cells. In some grasses bubble-like
cells that are part of the epidermis are referred to as bulliform cells.
Technically, the cells marked (be) in the photograph of AZ. lind-
heimert are bulliform cells. The cells subjacent to these bulliform
cells are the colorless cells (cc). In species of the Epicampes group

6’ According to Metcalfe (personal communication), “It is quite in order to
refer to the cells as thick-walled parenchyma unless they can be shown in Ls.
to be appreciably longer than wide. Even if they are somewhat elongated it is
best to refer to them as ‘elongated parenchymatous cells’ as they are not fibrous
or sclereids.”’

SODERSTROM—PODOSEMUM AND EPICAMPES 89

there is no obvious difference between the bulliform and colorless
cells apart from the position they occupy.

The Rigens and Gigantea Types of Leaf Anatomy

The leaf anatomy of the following species, traditionally placed in
E’picampes, was examined: Muhlenbergia distans, M. distichophylla,
M. emersleyi, M. gigantea, M. grandis, M. lindheimeri, M. longiligula,
M. macrotis, M. macroura, M. pubescens, M. rigens, M. robusta, M.
speciosa, and M. virletit.

All the above species possess the following anatomical features:
(1) The phloem of the 1° vascular bundles is partly sclerosed. (2)
The vascular bundle is surrounded by an inner sheath and an outer
sheath. Adjacent to the outer sheath is the chlorenchyma, appearing
under high power to consist of a layer of narrow cells, radially arranged.
(3) A large amount of thick-walled parenchyma occurs adaxial, some-
times abaxial, to the vascular bundle. (4) A thick cap of scleren-
chyma lies below the upper and lower epidermis of the 1° units. Apart
from shape, the units of all species of E’picampes are of similar struc-
ture. The major differences are in the kinds of units present and
in their arrangement.

The Rigens type of leaf anatomy is characterized by the following
features: (1) A similar alternation of units occurs throughout the leaf.
The 1° units alternate with one or two 2° units or three to five smaller
units (consisting of two to four 3° units and a 2° unit). There is no
differentiation of central units into a keel. (2) The lower surface of
the blade is flat, or ribbed and furrowed; the upper surface is ribbed
and furrowed. (3) The 1° units are rounded above in M. macroura
(cf. Mf. dubia, pl. 5, fig. 3), obtriangular in M. rigens (pl. 5, figs. 1, 2),
and in some species more or less rectangular (e.g., MZ. elongata, pl. 11,
fig. 2).

The following species of the Epicampes group possess a Rigens
type of leaf anatomy: MZ. macroura and M. rigens.

The Gigantea type of leaf anatomy is characterized by the fol-
lowing features: (1) Only 1° and 2° units are present. The central
units of the blade are specialized into a keel. The thick-walled
parenchyma of these units is confluent and a large cap of sclerenchyma
is developed at the base of the central unit. (See M. gigantea, pl. 9,
fig. 1, for illustrations of the central and adjacent units of the keel.)
Beyond the midrib area is a regular alternation of 1° and 2° units.
The 1° units are separated by one to three 2° units. (2) The upper
and lower surfaces of the blade are ribbed and furrowed. Beyond
the midrib area the units are separated by several layers of colorless

90) CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

cells. The sclerenchyma subjacent to the lower epidermis is con-
fined to the individual units and therefore does not form a continuous
layer between them. (8) The 1° units are more or less rectangular.
The central 1° unit of the keel is rounded or obtuse below.

The following species of the Epicampes group possess a Gigantea
type of leaf anatomy: MM. distans, M. distichophylla, M. emersleyi,
M. gigantea (pl. 7, fig. 2, and pl. 9, fig. 1), AZ. grandis, M. lindheimeri
(pl. 7, fig. 3, and pl. 8, fig. 3), AZ. longiligula (pl. 10, fig. 1), MZ. macrotis
(pl. 7, fig. 1), M. pubescens, M. robusta, M. speciosa, M. virletii.

The Rigens or Gigantea Type in Species of Muhlenbergia

Apart from the anatomical differences that separate the species of
the Epicampes group into the Rigens and Gigantea types, the two
groups have the same basic unit structure in the leaves and share the
following gross morphological features: Plants perennial, caespitose
(usually strongly so); culms erect, not branching at the nodes; and
panicle usually densely flowered.

The species with a Rigens type of leaf anatomy have rounded basal
sheaths, a firm ligule, and the leaves become inrolled upon drying.
In contrast, most species with a Gigantea type of leaf anatomy have
compressed-keeled basal sheaths, a membranous ligule, and the leaves
become folded upon drying.

Many species of Muhlenbergia share the morphological features
common to all of the species of the Epicampes group. These species
may be grouped into those having compressed-keeled basal sheaths
and a membranous ligule, and those with rounded basal sheaths in
association with a firm ligule. Because of the close morphological
similarities to the Epicampes group, a study of the leaf anatomy was
made, with the following results:

Muhlenbergia species with the Rigens type of leaf anatomy:

M. angustata M. holwayorum

M. arenicola M. jaliscana

M. articulata M. lucida

M. capillaris (pl. 10, fig. 2) M. metcalfei

M. dubia M. nigra

M. dubioides M. reverchonii (pl. 11, fig. 1)
M. elongata (pl. 11, fig. 2) M. rigida

M. expansa (pil. 6, fig. 2) M. setifolia

M. firma M. subaristata

M. glabrata M. xerophila

All of the above species have rounded basal sheaths and a firm ligule.
(The rather variable species M. rigida sometimes may have a long
membranous ligule.)

SODERSTROM—PODOSEMUM AND EPICAMPES Q]

Species that have the Gigantea type of leaf anatomy:

M. aurea M. longiglumis (pl. 10, fig. 3)
M. breviligula M. mutica

M. gooddingii M. pubigluma

M. inaequalis M. reederorum

M. involuta M. scoparia

M. tridifolia M. versicolor

M. lehmanniana M. xanthodas

These species have compressed-keeled basal sheaths and membranous
ligules with the exception of M. involuta and M. reederorum in which
the sheaths are rounded (rarely compressed) and ligules firm below
and membranous above.

Some species of Muhlenbergia exhibit certain anatomical features
that require special mention:

M. longiligula (pl. 10, fig. 1). The several 1° units of the midrib
area are of equal size. The central unit does not possess a larger
basal cap of sclerenchyma than the adjacent units of the keel. The
basal sheaths are either rounded or sometimes compressed, but never
strongly keeled. However, the structure of the units, the sclerosed
phloem of the 1° bundles, and the midrib area of thick-walled paren-
chyma are characteristic of the Gigantea group of species.

M. longiglumis (pl. 10, fig. 3). The central unit is a 1° bundle with
sclerosed phloem and a large basal cap of sclerenchyma; both are
characters of the Gigantea type of leaf anatomy. The central
units, however, are not embedded in a mass of thick-walled paren-
chyma. On the basis of leaf anatomy, this species does not conform
well to either anatomical type but shows more similarity to the
Gigantea type.

M. aff. montana (pl. 11, fig. 3). The units have an arrangement
similar to that of the Rigens type. However, the phloem of the 1°
units is not sclerosed.

M. involuta. All of the units of the leaf are embedded in a con-
tinuous layer of thick-walled parenchyma, unlike the Gigantea species
in which such a condition is confined to the keel. The phloem of the
1° bundles is sclerosed and the central units possess a large basal cap
of sclerenchyma, as in the Gigantea type. The anatomy of the leaf
most closely approaches the Gigantea type.

M. stricta. The units are all very large and are connected by a
continuous layer of thick-walled parenchyma and the phloem of the
1° bundles is sclerosed. The central unit does not possess an addi-
tional amount of basal sclerenchyma. On the basis of leaf anatomy
this species exhibits greater similarity to the Gigantea type, but its
gross morphology is more like that of species in the Rigens group.

92 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

M. torreyana (pl. 9, fig. 3). The leaves of this species have a keel
structurally like that of the Gigantea group, but the phloem of the 1°
bundles is not sclerosed. Morphologically it is unlike species of the
Gigantea group in having short, very scaly rhizomes and branching
culms, and a minute, ciliate, firm ligule.

Muhlenbergia Species Unlike the Rigens and Gigantea Types

Most species of Muhlenbergia have a leaf anatomy quite different
from the Gigantea and Rigens types. Although the anatomical struc-
ture of all cannot be discussed here, several points should be men-
tioned to illustrate how these species compare with the Epicampes
group and allies.

Photomicrographs of 11 species of Muhlenbergia with a leaf anatomy
distinct from the Rigens or Gigantea types are shown on plate 9,
figure 3, plate 11, figure 3, and plates 12-14.

All 11 of these species of Muhlenbergia differ from species of the
Rigens and Gigantea types in the condition of the phloem. The
phloem of all the bundles, including the 1° ones, when these are
differentiated, is darkly stained throughout, indicating it 1s com-
pletely functional and not sclerosed. Special reference is made to
M. aff. montana (pl. 11, fig. 3) and M. torreyana (pl. 9, fig. 3). In these
species the 1° units are similar to those of the Epicampes group but
the phloem of the 1° units is not sclerosed.

The annual species are characterized by a relatively simple unit
structure, with little additional sclerenchyma or thick-walled paren-
chyma. Note, for example, the annuals M. confusa (pl. 13, fig. 1),
M. crispiseta (pl. 14, fig. 1), M. peruriana (pl. 12, fig. 1), and M. pul-
cherrima (pl. 12, fig. 2). In M. peruviana and M. pulcherrima the
units are of equal size; the outer sheath and chlorenchyma are con-
tinuous above. Additional sclerenchyma and parenchyma are
lacking in the units and the phloem is not sclerosed. M. crispiseta
has sclerenchyma developed in the central unit and some in each of
the other units. The outer sheath and chlorenchyma entirely circle
the vascular bundle without interruption and the phloem in the
bundles is not sclerosed.

Among perennial Muhlenbergias, plates 12-14, are M. arizonica
(pl. 12, fig. 3), M. asperifolia (pl. 14, fig. 2), Af. plumbea (pl. 13, fig. 3),
and M. richardsonis (pl. 13, fig. 2). The structure of the individual
units in M. richardsonis and M. asperifolia approaches that of the
Epicampes group in its specialization. However, the phloem of
the 1° bundles in these two rhizomatous perennials is not sclerosed.
Muhlenbergia plumbea, another rhizomatous perennial, has units
mostly equal in size, except for the central unit which has a basal

SODERSTROM—PODOSEMUM AND EPICAMPES 93

cap of sclerenchyma. Note that the outer sheath and chlorenchyma
are continuous around the vascular bundles. The phloem of the
bundles is not sclerosed. The leaf of M4. arizonica, a perennial that
branches at the nodes, is not too unlike that of M. plumbea. The
units are mostly equal in size, and the median one is heavily scleri-
fied. The outer sheath and chlorenchyma are continuous around the
vascular bundles outside of the central unit. The phloem of the
vascular bundles is not sclerosed.

Various anatomical configurations in the leaves of the remaining
species of Afuhlenbergia examined are not illustrated here.’ In all of
these species the phloem of the vascular bundles is not sclerosed.

Conclusions

The leaves of species traditionally placed in Epicampes are char-
acterized by an inner sheath (or endodermis), a parenchyma (outer)
sheath, and chlorenchyma, consisting of cells radially arranged.
All are features of the chloridoid type of leaf anatomy, as pointed out
by Brown (1958). On the basis of leaf anatomy these species are
allied to Muhlenbergia, Sporobolus, Lycurus, and other genera of the
chloridoid alliance. Jn contrast, the leaves of Agrostis and Cinna are
of the festucoid type, as Brown has shown, and therefore do not
show a relationship to Epicampes. On the basis of leaf anatomy,
Epicampes should be removed from the Agrostideae and placed
with other members of the chloridoid alliance. The present investi-
gations have shown that E’picampes is most closely related to species of
Muhlenbergia which not only have similar gross morphological features,
but leaf anatomical features; most noteworthy is the unique condition
of sclerosed phloem in the 1° vascular bundles. These findings
support those of Schwabe (1949) who in a study of the foliar anatomy
of 22 species of Muhlenbergia, including two species of the Epicampes
group, concluded that the latter should probably be included in the
former.

LEAF EPIDERMIS

Sections were taken from the upper and lower epidermis of the
leaf. In all cases the preparations were made from leaves taken
from herbarium specimens. Sections of the blade, 1-1%’’ long,
were placed in boiling water until softened, just as in preparing

7 The reader is referred to the treatment of Schwabe (1949) in which features
of the leaves of many additional species of Muhlenbergia are illustrated.

94 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

the whole leaf for cross sections. To remove the epidermis of the
abaxial surface the blade was placed with the abaxial surface down
on a clean glass slide. With the aid of the 10 X power of a binocular
microscope, the tissue of the leaf was scraped away with a scalpel
until only the epidermis remained. The same process was repeated
for the adaxial surface with the adaxial surface down.® The epi-
dermal scrapes were stained with methylene blue and counterstained
with ruthenium red. They were then washed in water to remove the
excess dye, brought through the alcohol and xylene series, and
mounted in Permount.

Constituents

The epidermis of the grass leaf contains a number of different cell
types, among them microhairs, prickle-hairs, papillae, long cells,
stomatal cells, interstomatal cells, and silica cells. The bicellular
microhairs ® and silica cells exhibit the greatest differences between
the groups of species within Epicampes.

Micronairs: All species studied of Muhlenbergia (including
Epicampes) have bicellular microhairs. Differences are found in the
shape and in the ratio of the length of the basal to the upper (distal)
cell. In some species the microhairs are short and broad; the basal
cell is equal to, or a little longer than, the distal cell. In other species
the microhairs are longer and narrower, and the basal cell is much
longer than the distal cell.

Sinica CELLS: Silicon-containing cells in the epidermis are referred
to as “silica cells” or “siliceous cells.” Characterized by a wall
heavier than that of adjacent cells with small granules within the cell,
they occur in rows and are a major constituent of the epidermis in
species of the Epicampes group. In some species all of the silica cells
are saddle-shaped and rather simple in outline; in others most of them
are dumbbell- or cross-shaped; in still others the majority are saddle-
shaped, with cross-shaped cells occurring infrequently among them.

8 In many species, e.g., M. macroura, the blade is characterized by a flat abaxial
surface and an adaxial surface of ribs and furrows. In such cases the whole
epidermis scrapes off easily from the abaxial surface but only strips of epidermis
can be removed from the adaxial surface, representing the top surfaces of the ribs
only.

9 Metealfe (1960, p. xliii) remarks, “Micro-hairs are not always easy to examine,
but it is generally rewarding to persevere with them because they are valuable
indicators of the affinities of the grasses in which they occur.”’

95

SODERSTROM—PODOSEMUM AND EPICAMPES

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96 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Following is a summary from epidermal studies in Epicampes

species:
Shape of silica cells:
s=saddle-shaped

Average ratio of basal to x=cross-shaped
Species distal cell of microhair d=dumbbveli-shaped
M. distans 6.6:1 S
M. distichophylla 5.1:1 s
M. gigantea 4.0:1 8
M. grandis 3.2:1 8
M. lindheimeri - 8
M. macrotis 5.7:1 8
M. macroura 2.2:1 mostly x or d
M. pubescens 4.7:1 s
M. rigens 2.3:1 mixed, sand d
M. robusta 4.3:1 8
M. speciosa - s
M. virletii 4.0:1 s

Species of the Epicampes group with a Rigens type of leaf anatomy
(M. macroura and M. rigens) exhibit certain characteristics in their
epidermis different from the other species of Epicampes. The
bicellular microhairs are short and broad in these species and silica
cells are dumbbell-shaped, cross-shaped, or saddle-shaped. In the
species that have a Gigantea type of leaf anatomy, the bicellular
microhairs are generally long and narrow, and the upper cell is usually
much shorter than the basal cell. In these species the silica cells are
always saddle-shaped.

The epidermis of other species of Muhlenbergia with a Rigens or
Gigantea type of leaf anatomy also was studied. In these, likewise,
the epidermis of members with a Gigantea type is characterized by
mostly long and narrow bicellular microhairs and rectangular or
saddle-shaped silica cells. And in the epidermis of species with a
Rigens type the silica cells are cross-shaped, saddle-shaped, or rec-
tangular, and the bicellular microhairs relatively short and broad. In
figure 1 are illustrated the major types of silica cells and bicellular
microhairs encountered.

Conclusions

Epidermal features of the species of Muhlenbergia, including
Epicampes, are panicoid in nature; microhairs are present and the
silica cells are rectangular, saddle-shaped, cross-shaped, or dumbbell-
shaped. Relationship with the chloridoid group is apparent in the
club-shaped or globose microhairs and membranes of equal thickness
on the upper and lower cells (Prat, 1936; Tateoka, Inoue, and Kawano,
1959; Metcalfe, 1960).

SODERSTROM—PODOSEMUM AND EPICAMPES 97

The Embryo

Until recently the structure of the grass embryo has been of passing
interest to grass taxonomists. Bruns (1892), who dealt with about
60 species, was the first worker to publish on grass embryos. Later,
Van Tieghem (1897) and Kennedy (1899) added to the understanding
of the grass embryo, the work of the former being more significant
from the standpoint of systematics. Not until Yakolev (1950) and
Reeder (1953) was the value of the embryo in grass systematics again
stressed. Reeder (1957) in a paper of great significance presented his
findings based on a study of the caryopses of about 300 species of
grasses. ‘These included over 150 genera, representing all of the
tribes in the traditional Hackelian system. Reeder showed that
several embryo groups could be recognized, and the criteria he used
to distinguish them are the relative size of the embryo to endosperm
in the caryopsis, as well as the features observable in the transverse
and median sagittal sections of the embryo. He listed six groups of
grasses based on various combinations of four embryo characters:
festucoid, panicoid, chloridoid-eragrostoid, bambusoid, arundinoid-
danthonioid, and oryzoid-olyroid. Reeder later (1962) reinterpreted
the oryzoid-olyroid genera as bambusoid but segregated a further
group, the centothecoid.

In the present study, mature caryopses were selected from herbarium
specimens, placed in boiling water for a few minutes, then soaked over-
night in FAA. They were then transferred to a 3 percent solution of
KOH for 1-2 hours to soften the endosperm. After removing the
endosperm, the embryo was embedded in agar and the agar block,
containing the embryo, run through the butyl alcohol series and
embedded in paraffin. Serial sections 104 in thickness were made,
transversely and longitudinally, in the sagittal plane. Safranin and
Fast Green were used in staining.

Embryos of 10 species of Muhlenbergia were sectioned, of which 9
species have been traditionally associated with Hpicampes. Of these,
7 have a Gigantea type of leaf anatomy: MM. emersleyi, M. gigantea,
M. grandis, M. longiligula, M. macrotis, M. pubescens, and M. robusta.
All are illustrated in figure 2 except M. emersleyi. Two have a Rigens
type: M. macroura (fig. 3) and M. rigens (fig. 2). Additional species,
represented in figure 3, are M. nigra, with a Rigens type, and M.
schreberi, the type species of Muhlenbergia. For comparison, diagrams
also of the embryos of five genera that have been suggested as bearing
a relationship to Epicampes appear in figure 3: Agrostis, the type genus
of the classical tribe Agrostideae, Cinna, Polypogon, Sporobolus, and

Stipa.

98 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

a

f

GV

(

Ficure 2.—Outline drawings of caryopses, median sagittal sections, and transverse sections
through the coleoptile region of Muhlenbergia embryos: a, M. macrotis (Pringle 2360);
b, M. robusta (Standley 59136); c, M. robusta (Pringle 2321); d, M. grandis (Pringle 2765);
e, M. gigantea (Pringle 3335); f, M. pubescens (Arséne); g, M. longiligula (Reeder &3 Reeder
2699); h, M. rigens (Blumer 1491). Drawings are diagrammatic and not drawn to scale.

§

=>

te

SODERSTROM—-PODOSEMUM AND EPICAMPES 99

M A

oy, Y EY

w

e f g h

Ficure 3.—Outline drawings of caryopses, median sagittal sections, and transverse sec-
tions through the coleoptile region of embryos of Muhlenbergia and other genera: a,
Muhlenbergia macroura (Arséne); b, M. nigra (Cook); c, M. schreberi (no voucher); d, Agro-
stis scabra (Goodding); e, Cinna arundinacea (Tweedy); f, Polypogon monspeliensis (Hortus
Botanicus Bruxellensis); g, Sporobolus wrightii (Goodding); h, Stipa avenacea (Reeder €3
Reeder 233), Drawings are diagrammatic and not drawn to scale.

100 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

The caryopses and embryos of Muhlenbergia species of the Epi-
campes group have the following features:
Caryopsis: The embryo is relatively large in comparison with the
endosperm.
Embryo: Median sagittal section.
1. A distinct internode is present in the vascular system between
the point of divergence of the trace to the scutellum and the
trace to the embryonic leaves (P).
2. An epiblast is present (+).
3. A cleft is present between the lower part of the scutellum and
the coleorhiza (P).

Embryo: Transverse section.

4. The margins of the embryonic leaf contain few bundles and
the margins do not overlap (F).

The embryo formula (see Reeder, 1957) for these species is P+ PF,
the chloridoid-eragrostoid type: Muhlenbergia and Sporobolus also are
chloridoid-eragrostoid, but Agrostis, Cinna, Polypogon, and Stipa are
festucoid.

Muhlenbergia macroura and M. nigra differ from the other species
in lacking an epiblast, unusual for chloridoid-eragrostoid species. On
the basis of gross morphology and leaf anatomy, however, they are
quite similar to M. rigens and an epiblast is present in that species as
well as in M. dubioides.

Of 12 caryopses examined from a single specimen of Muhlenbergra
pubescens (Soderstrom 693), 7 had double embryos. However, other
specimens of M. pubescens from different localities and all other species
examined had single embryos.

Conclusions

The embryos of Muhlenbergia do not show basic differences charac-
teristic of individual groups, except Mf. macroura and M. nigra which
differ in the absence of an epiblast. As emphasized by Reeder, the
embryo is most useful in indicating the placement of a grass within
one of six natural groups of genera. The species of the Epicampes
eroup are like other species of Muhlenbergia and Sporobolus and are
related to grasses of the chloridoid-eragrostoid group. They do not
show a relationship to Agrostis and other festucoid genera of the
Agrostideae with which they have been traditionally associated. A
summary of embryo characters follows (see also figs. 2 and 3):

SODERSTROM—PODOSEMUM AND EPICAMPES 101

Embryo/ | Internode | Epiblast| Cleft in Embryonic leaf
endosperm| in trace | present | scutellum margins meet,
Genus and/or species large (P) | present (+) present bundles few (F) Formula
amall (F)|(P) absent} absent |(P) absent| margins overlap,
(F) (-—) (F) bundles many (P)
Muhlenbergia
Gigantea type of
leaf anatomy
M. emersleyi P P + P F P+PF
M. gigantea P P + Pp F P+PF
M. grandis P P + P F P+PF
M. longiligula P P + P F P+PF
M. macrotis P P + P F P+PF
M. pubescens P P + P F P+PF
M. robusta P P + P F P+PF
Rigens type of
leaf anatomy
M. macroura P P — P F P—PpF
M. nigra P P — P (Not seen) | P—P
M. rigens P P + P F P+PF
Type species
M. schreberi P P + P F P+PF
Agrostis F F + F F F+FF
Cinna F F + F F F+FF
Polypogon F F + F F F+FF
Sporobolus Pp P + P F P+PF
Stipa F F + F F F+FF
Lodicules

Lodicules are small hyaline or fleshy scalelike structures subsessile
between the lemma and palea of a grass floret. At time of anthesis
when they reach their maximum development they become swollen
and force the lemma and palea apart, thereby allowing the stamens
to become exserted from the spikelet. They are thought to repre-
sent a reduced inner whorl of perianth parts, the outer whorl having
been lost completely through evolution. Although only two lodicules
are found in most genera, three are present in the floret of Stipa
and most bamboos; lodicules are lacking entirely in Anthozanthum,
Cenchrus, and Pennisetum. The lodicules are usually separate, but
are fused in Melica and Glyceria. Not only are differences in lodi-
cules among genera found with respect to the number present, but
also in size and shape, manner of thickening, and vascularization.

Zuderell (1909) examined the lodicule anatomy of about 50 species
of grasses and in the same year Krause mentioned that lodicules
might be of systematic value. Apparently, not until recently has
real emphasis been placed on these structures as indicators of natural

102 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ficure 4.—Lodicules of Muhlenbergias in the Podosemum section, all times 46: a, M.
angustata (Camp E-2519); b, M. glabrata (Amer. Gr. Nat. Herb, 1356); ¢, M. dubia (Soder-
strom 509); d, M. rigida (Soderstrom 640); ¢, M. macroura (Soderstrom 551); f, M. nigra
(Pringle 11739); g, M. palmeri (Pringle 1417).

OV 1 6

no Wy oe

Ficure 5.—Lodicules of Muhlenbergias in the Podosemum and Epicampes sections, all
times 45: a, M. firma (Amer. Gr. Nat. Herb, 1351); b, M. rigens (Soderstrom 731); c, M.
capillaris (Kral & Godfrey 54211); d, M. xerophila (Goodding A-9493); ¢, M. expansa
(Godfrey 8151); f, M. arenicola (Pringle 479); g, M. articulata (Pringle 3913); h, M.
reverchonii (Silveus 2478); i, M. longiglumis (Pringle 2365); j, M. stricta (Soderstrom 642);
k, M. involuta (Silveus 780); 1, M. emersleyt (Soderstrom 746).

SODERSTROM—PODOSEMUM AND EPICAMPES 103

a b Cc

d

k |

Ficure 6.—Lodicules of Muhlenbergias in the Epicampes section, all times 46: a, M.
distans (Pringle 5374); b, M. speciosa (Reeder @ Reeder 2476); c, M. grandis (Pringle
2765); d, M. distichophylla (Pringle 2346); e, M. robusta (Arséne); f, M. versicolor (Smith
927); g, M. macrotis (Mexia 9098); h, M. pubescens (Soderstrom 483); 1, M. lindhermeri
(Burr 513); 7, M. robusta (Soderstrom 382); k, M. longiligula (Reeder & Reeder 3222);
l, M. gigantea (Reeder @ Reeder 2488).

relationships. Stebbins (1956), in a study of intergeneric hybrids
of Lolium and Festuca, pointed out that cytological findings agree
with those based on studies of the grass flower and caryopsis. He
illustrated four types of lodicules which he designated panicoid,
chloridoid, festucoid, and bambusoid. In the panicoid and chlori-
doid types the lodicules are truncate at the apex, but pointed in the
festucoid and bambusoid types. Church (1949) also mentioned the
value of using characters of the grass flower, including lodicules, in
systematics. Although Hubbard (1934, 1959 rev.) figured the lodi-
cules of many species, he did not discuss them from a comparative
viewpoint. Some understanding of the diversity of structure among
genera can be gained from his sketches of the lodicules of nearly
every species of grass from the British Isles (1954). Bor (1960)
likewise included sketches of the lodicules of many grasses.

Reeder and Ellington (1960) demonstrated that the genus Ca-
lamovilfa is chloridoid and should be removed from the Agrostideae.
Their conclusions were based on lodicule anatomy along with embryo
structure, leaf anatomy and epidermis, and chromosome size and
number. Tateoka (1960a) also used lodicules, among other floral

221-352-673

104 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

g h
Picure 7.—Lodicules of Muhlenbergias other than the Podosemum or Epicampes sections,
all times 50: a, M. quadridentata (Hinton 4389); b, M. montana (Arséne 5811); c, M.
schreberi (no voucher); d, M. wrightii (Goodding & Hardies A-8498); e, M. uniflora

(Miller 130); f, M. biloba (Pringle 819); g, M. depauperata (Reeder @ Reeder 2279); h, M.
brevis (Reeder Reeder 2620); 1, M. torreyi (Goodding M-189).

e f g

Ficure 8.—Lodicules of Muhlenbergias other than the Podosemum or Epicampes sections,
all times 45: a, M. sylvatica (no voucher); b, M. racemosa (Cornman); c, M. frondosa
(no voucher); d, M. andina (Reeder G& Reeder 1808); e, M. glomerata (Reeder © Reeder
944); f, M. mexicana forma ambigua (no voucher); g, M. torreyana (Bissell).

SODERSTROM—PODOSEMUM AND EPICAMPES 105

| h 4
esl foots Oa,

h
Ficure 9,—Lodicules of genera that have been associated with Epicampes, all times 44:
a, Sporobolus wrightii (Toumey); b, Blepharoneuron tricholepis (Reeder @ Reeder 1807);
c, Stipa avenacea (Reeder & Reeder 233); d, Agrostis alba (Dudley); e, Cinna arundinacea
(no voucher); f, Polypogon monspeliensis (Swan); g, Lycurus phieoides (Reeder G Reeder
2352); h, Sporobolus asper (Nichols); i, Sporobolus heterolepis (Nichols).

features, to show the relationships of several genera of the Festuceae.
Reeder ?° reported on the preliminary results of his study of the lodi-
cules of over 300 species, representing 230 genera. Decker (1964),
in a study of 135 genera, utilized information from the lodicules,
along with other microcharacters, to determine the naturalness of
the classical tribe Festuceae. Hsu’s (1963) study is an example of
lodicules figured for several species in a single genus.

In the present study the lodicules of 47 species of Muhlenbergia
(including Epicampes) have been examined. Included are 13 species
with a Gigantea type of leaf anatomy, 15 with a Rigens type, 3 inter-
mediate between the two, the type species of Muhlenbergia, and 15
additional species. Line drawings of the lodicules of each of these
species, in addition to those of several other genera with which
Epicampes has been associated, appear in figures 4-9. The vascular
tissue of the lodicules and manner of thickening are indicated in these
drawings.

All lodicules were selected from spikelets of herbarium specimens
in which the anthers were fully developed but not yet exserted.

10 Paper read at meetings of the American Institute of Biological Sciences,
Stillwater, Oklahoma, 1960.

106 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

The spikelets were boiled in water for several minutes, transferred
to a watch glass, and the lodicules dissected out under the low power
of a binocular microscope. The shape of the lodicules was drawn at
this time and the areas of thickening noted, then the lodicules were
mounted in glycerine jelly without staining. From 2 to 3 weeks later
the excess glycerine jelly was removed and the slides ringed with
diaphane. The vascularization was drawn later from the slides under
the high power of a compound microscope.

Most of the species of Muhlenbergia examined possess lodicules
more or less rectangular in outline. The thickened portion of the
lodicules extends from the base to near the apex; the upper margins
are of a thinner texture. One to many vascular traces usually are
present in the form of annular tracheids—rarely are they lacking
entirely. According to Stebbins (1956) and Reeder and Ellington
(1960), these lodicules are of the chloridoid type.

The lodicules of species with a Gigantea type of leaf anatomy
(figs. 5¢ and 6a-j, /) possess one to four or five vascular traces. Apart
from small differences in size and amount of vascularization, the
lodicules appear to be rather similar. Lodicules of the species with a
Rigens type of anatomy (figs. 4a-g and 5a-h) usually possess two to
many traces. The lodicules of M. rigens (fig. 56) are atypical in
having no vascular traces; in M. arenicola (fig. 5f) the one or two traces
present are almost negligible. The considerable difference in lodicule
size is attributable to the difference in the size of the spikelets. The
large lodicules of M. macroura (fig. 4e), M. nigra (fig. 4f), and M.
palmerv (fig. 4g) reflect the large spikelet size in these species. Lodicule
shape, degree of thickening, and vascularization, with little exception,
are similar in all species with a Rigens type of leaf anatomy.

Species of Muhlenbergia not of the Rigens or Gigantea groups
exhibit the same kind of thickening in the lodicules and in degrees
of vascularization. The lodicules of many of them are like the
species with a Rigens or Gigantea type of anatomy—WM. torreyana
(fig. 89), Af. quadridentata (fig. 7a), M. montana (fig. 76), and M.
wrighti (fig. 7d).

In the type species of Muhlenbergia, M. schreberi (fig. 7c), the
upper margin of the lodicule is extended on one side into a small
“wing.” A similar lodicule is found in M. uniflora (fig. 7e). In
M. racemosa (fig. 8b), a species with scaly rhizomes, the upper corners
of the lodicules are prominently projected. Other species with scaly
rhizomes also have winglike extensions, to a lesser or greater degree—
M. sylvatica (fig. 8a), M. frondosa (fig. 8c), M. andina (fiz. 8d), M.
glomerata (fig. Se), and M. mexicana forma ambigua (fig. Sf).

Interesting differences are found among lodicules of various genera
that, in one system of classification or another, have been considered

SODERSTROM—PODOSEMUM AND EPICAMPES 107

as closely related to the Epicampes group. Of the festucoid type
Agrostis (fig. 9d), Cinna (fig. 9e), Polypogon (fig. 9f), and Stipa (fig. 9c)
have long narrow lodicules with only the lower part thickened.
Stipa further differs in having three, rather than two, lodicules.
Blepharoneuron (fig. 96) possesses truncate lodicules in which the
thickening extends from the base to the apex, and is thus of the
chloridoid type, as are the lodicules of Lycurus (fig. 9g) and Sporobolus
(figs. 9a,h,2). Blepharoneuron and Lycurus show no fundamental
differences from Muhlenbergia. However, the lateral unthickened
margins in Sporobolus are considerably wider than in any of the
species of Muhlenbergia examined.

Conclusions

On the basis of lodicules, the species of Muhlenbergia, including
Epicampes, are of the chloridoid type, along with such genera as
Blepharoneuron, Lycurus, and Sporobolus. They are unlike the
festucoid genera Agrostis, Cinna, Polypogon, and Stipa.

Differences of lodicules among the species of Muhlenbergia are of
comparatively little value in recognizing groups of related species.
However, the type species of Muhlenbergia and similar species with
scaly rhizomes do exhibit a difference in the shape of the lodicules,
and further studies on the remaining species of this genus may reveal
that other species have distinctive lodicules.

A summary of the lodicule characters for each of the genera dis-
cussed follows (see also figs. 4-9):

Genus veins Shape Thickening Vascular- | Lodicule
per ization group
floret
Muhlenbergia
Gigantea 2 truncate thickened from base | 1—many C*
type of leaf to near apex; traces
anatomy lateral margins
thin
Rigens type 2 truncate thickened from base | 1-many C
of leaf to near apex; traces
anatomy lateral margins
thin
Type species, 2 truncate, thickened from base | 2-4 Cc
M. schreberi with pro- to near apex, traces
and allies jections lateral margins
thin

*See footnote at end of table.

108 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Lodi-
Genus cules Shape Thickening Vascular- Lodicule
per izaton group
flor et
Agrostis 2 narrow, thickened below; lacking KF
pointed thin above
Blepharoneuron 2 truncate thickened from base | lacking C
to apex, lateral
margins thin
Cinna 2 narrow, lower % thickened, lacking F
rounded upper }4 thin
Lycurus 2 truncate thickened from base | 1 short C
to near apex; trace
lateral margins
thin
Poly pogon 2 narrow, lower 4 thickened, lacking F
pointed upper }6 thin
Sporobolus 2 very wide, | thickened from base | 1-several | C
more or to near apex; traces
less lateral margins
truncate thin
Stipa 3 narrow, thickened below, lacking F
pointed thin above
C=chloridoid. F=festucoid.
Chromosomes

Avdulov (1931) was the first worker to utilize cytological informa-
tion as a basis for a new system of classification of the Gramineae.
Subsequent workers, Prat (1936), Pilger (1954), and Stebbins (1956),
have also employed cytological in addition to other data in their
proposed systems. The role of cytology in grass systematics has been
thoroughly reviewed by Tateoka (1960b).

Although cytological studies were not undertaken in the present
investigation, several workers have published results of studies on
many species of Muhlenbergia (including Epicampes) and putative
allied genera, from which the following summary was made.

According to Darlington and Wylie (1956) x=7 is the basic chromo-
some number for Agrostis, Cinna, and Polypogon. These genera
traditionally have been placed with E'picampes in the tribe Agrostideae,
along with Stipa, Sporobolus, and Muhlenbergia. On the basis of
cytology, Agrostis, Cinna, and Polypogon are festucoid, in corrobo-

SODERSTROM—PODOSEMUM AND EPICAMPES 109

ration with the conclusions based on other studies. The basic numbers
Darlington and Wylie listed for Stipa are x=9, 10, 11, 12, 14, 16, or
17; for Sporobolus, x=9, 10, or 12; and for Afuhlenbergia, x=9, 10, or
21. A survey of the chromosome numbers published for Sporcbolus
indicates that most of the species have a basic number of 9 chromo-
somes, whereas most of the species of Muhlenbergia (including Epi-
campes) have a basic number of x=10. Stipa, in which a high degree
of aneuploidy occurs (Stebbins & Love, 1941, p. 379), does not appear
to have a consistent basic number. Stebbins (1956) considers this
genus to be a member of a specialized line, the Stipeae, the members
being neither completely festucoid nor panicoid. He lists x=11 or
12 as the basic number for this group.

Chromosome counts of seven species of the Epicampes group of
Muhlenbergia are listed below: M. emersleyi (2n=40 & 60); M.
gigantea (2n=20); M. lindheimeri (2n=20); M. macroura (2n=40);
M. pubescens (2n=40); M. rigens (2n=40); and M. reederorum (2n=
68). Following is a list of American species of Muhlenbergia for which
chromosome numbers have been determined:

Species of Muhlenbergia 2n Authority
Gigantea type of leaf anatomy
M. emersleyi 40 Brown (1950)
60 Reeder (1961, unpublished)
M. gigantea 20 Tateoka (1962)
M. lindhetmeri 20 Brown (1950)
20 Reeder (1965, unpublished)
M. pubescens 40 Reeder (1961, unpublished)
M. reederorum 68 Reeder (1965, unpublished)
Rigens type of leaf anatomy
M, macroura 40 Tateoka (1962)
M. reverchonit Vasey & Scribn. 40 Brown (1951)
M. rigens 40 Stebbins & Love (1941)
40 Reeder (1961, unpublished)
M. rigida (H.B.K.) Trin. 40 Gould (1958)
Other species
M. andina (Nutt.) Hitche. 20 Myers (1947; counted by
Stebbins)
M. asperifolia (Nces & Mey.) 20 Myers (1947; counted by
Parodi Stebbins)
M. brachyphylla Bush 40 Brown (1950)
M. filiformis (Thurb.) Rydb. 18 Myers (1947; counted by
Stebbins)
M. mexicana Trin. 40 Avdulov (1931)
M. monticola Buckl. 40 Brown (1951)
M. polycaulis Scribn. 20 Brown (1951)
M. porteri Scribn. 20 Brown (1951)
40 Gould (1960)
M. pungens Thurb. in A. Gray 42 Nielsen & Humphrey (1937)

60 Nielsen (1939)

110 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Species of Muhlenbergia an Authority
M. racemosa (Michx.) B.S.P. 40 Avdulov (1931)
M. repens (Presl) Hitche. 60 Brown (1951)
M. richardsonis (Trin.) Rydb. 40 Stebbins & Love (1941)
M. schrebert Gmel. 40 Gould (1958)
M. squarrosa (Trin.) Rydb. 40 Bowden (1960)
40 Stebbins & Love (1941)
M. sylvatica (Torr.) Torr. ex 40 Avdulov (1931)
Gray
M. tenuifolia (A.B.K.) Kunth 40 Tateoka (1962)
M. uniflora (Muhl.) Fern. ca. 42 Bowden (1960)

On the basis of cytology, Muhlenbergia (including the Epicampes
group) is panicoid (or chloridoid) rather than festucoid, as the chromo-
somes are small and in multiples of 10. The chloridoid genus Sporo-
bolus has a basic number of 9 and thus is distinct from Muhlenbergia.
Of the few species known cytologically, the diploid species of the
Epicampes group, M. lindheimeri and M. gigantea, are quite uniform
morphologically as would be expected. Two counts are available for
the M. emersleyi complex, one a tetraploid and the other a hexaploid;
further counts will probably reveal a range of numbers among the
various forms assigned to the species. The most interesting count is
of M. reederorum, evidently an aneuploid. From the gross morphology
MM. reederorum appears to have been derived in part from M. lon-
giligula, but we have no count for that species.

In regard to the disposition of Hpicampes itself, Stebbins & Love
(1941, p. 379) said:

The chromosome number and size obtained for Muhlenbergia richardsonis
(Trin.) Rydb. and M. rigens (Benth.) Hitehe. (2n=40, fig. 22) agrees with
that reported by Avdulov (1931) for other species of Muhlenbergia. Since
the chromosomes of M. rigens are closcly similar to those of the other species

of the genus, the union by Hitchcock of the genera Epicampes and Muhlen-
bergia is well supported by the cytological evidence.

The Disposition of Epicampes

The results of studies of the leaf anatomy and epidermis, embryos,
lodicules, and chromosomes agree that the species traditionally re-
garded as E’picampes are of the chloridoid alliance. Several genera of
the tribe Agrostideae, Agrostis, Cinna, and Polypogon, are of the
festucoid alliance and are not closely related to Epicampes as has
often been suggested.

Features of the embryos have been useful only in demonstrating
that the species of the Epicampes group are chloridoid but did not
serve to differentiate genera within this alliance. The lodicules of the
Epicampes group are also of the chloridoid type as are those of
Muhlenbergia but no fundamental differences were found among

SODERSTROM—PODOSEMUM AND EPICAMPES 111

them. The lodicules of Sporobolus were found to be somewhat dis-
tinctive in being larger and wider than those of Muhlenbergia (includ-
ing Epicampes). In the few species of the Epicampes group with a
known chromosome number, the basic number is the same as that of
other species of Muhlenbergia, but differs from that of Sporobolus.
This cytological information should indicate that species of the
Epicampes group are more closely related to Muhlenbergia than to
Sporobolus.

The studies most useful in clarifying the relationships of the Epi-
campes group have been those of leaf anatomy and, to a more limited
extent, the leaf epidermis. They have shown that the Epicampes
group is chloridoid and have pointed to the most closely related species
within Muhlenbergia.

The species of Muhlenbergia, including the Epicampes group,
possessing a Rigens or Gigantea type of leaf anatomy have the follow-
ing morphological features in common (all species examined with
other types of leaf anatomy also differ morphologically in one or
more respects from these):

1. Plants perennial, strongly caespitose (rhizomes never pro-
duced).
2. Culms stout, robust, erect, not branching or rooting at the
nodes.
3. First and second glumes nerveless or 1-nerved.
This group is well defined not only on the basis of gross morphology
but also in characters of leaf anatomy and epidermis; on this evidence
I feel this group of species should be treated as a subgenus of Muhlen-
bergia comprising two sections. The section E’picampes, characterized
by a Gigantea type of leaf anatomy, includes among others M.
robusta (including LE. strictus, the type of the genus E’picampes). The
section Podosemum, characterized by a Rigens type of leaf anatomy,
includes among others M. rigens, M. macroura, and M. capillaris
(based on P. capillare, the type species of the genus Podosemum).

Systematic Treatment

Muhlenbergia

Muhlenbergia Schreb. in L. Gen. Pl, ed. 8, 55. 1789.

Type species: Muhlenbergia schreberi Gmel. in L., Syst. Nat., ed. 13, 2:171.
1791.

Subgenus 1. Muhlenbergia.

Subgenus 2. Podosemum (Desv.) Soderstrom, stat. nov.

Podosemum Desv., Nouv. Bull. Soc. Philom. Paris 2:188. 1810.
Type species: Podosemum capillare (Lam.) Desv.=Muhlenbergia
capillaris (Lam.) Trin. Originally described as Stipa capillaris
Lam. Tabl. Encycel. 1:158. 1791.

112 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

MorpronoacicaL Features: Plants perennial, caespitose, rhizomes
not produced; culms rather stout to robust, erect, not branching or
rooting at the nodes; glumes nerveless or 1-nerved (rarely an additional
faint nerve or nerves present).

ANATOMICAL FRATURES OF THE Lear: Phloem of the 1° vascular
bundles sclerosed; large caps of sclerenchyma present in the 1° units.

Section 1. Podosemum"!

MorpnotocicaL Fraturss: Basal sheaths rounded; ligule firm;
glumes equal or unequal, shorter to longer than the floret, membranous
to firm; awn of lemma (when present) arising terminally or dorsally,
from just below the tip.

AnatomicaL Fraturses oF THE Lear: Central 1° unit not differ-
entiated from other 1° units; no keel present; 3° units often present
in addition to 1° and 2° units, the units alternating in a like fashion
throughout the leaf.

Species INcLupED (asterisked species originally described as
Epiwcampes): M. angustata, M. arenicola, M. articulata, M. capillaris,
M. dubia, M. dubioides, M. elongata, M. expansa, M. firma, M. glabrata,
M. holwayorum, M. lucida, M. macroura, M. nigra, M. palmeri,
M. reverchonu, *M. rigens, M. rigida, M. xerophila.

Section 2. Epicampes (J. S. Presl) Soderstrom, stat. nov.

Basionym: Epicampes J. 8. Presl in K. B. Presl, Rel. Haenk. 1:235.
t. 39. 1830. Type species: Mpicampes strictus Presl, loc. cit. =M.
robusta (Fourn.) Hitche.

MorrenoioaicaL Frarurss: Basal sheaths compressed-keeled;
ligule membranous throughout (firmer below, membranous above in
a few); glumes about equal, generally longer than the floret, mem-
branous, nerveless or indistinctly 1I-nerved; awn of lemma (when
present) arising from a mucronate tip or dorsally, from just below the
tip.

ANATOMICAL FraturREes or THE Lear: Central 1° unit with a
large basal cap of sclerenchyma—this and adjacent units sunken in a
confluent mass of thick-walled parenchyma, forming a keel; only
1° and 2° units present, these alternating in a like pattern on either
side of the keel.

1 Since this section includes the type species of the next higher rank, the sub-
genus Podosemum, it appears without author citation. It does, however, coincide
with Section 7 Pedosemum (Desv.) Pilger in Die Natiirlichen Pflanzenfamilien,
2nd ed., 14d:71. 1956, except for Muhlenbergia distichophylla (Presl) Kunth which,
under the present treatment, belongs in section 2, Epicampes. Section 4 Cinnas-
trum, of Pilger’s work (p. 69), also belongs in section Podosemum of my treatment.

SODERSTROM—PODOSEMUM AND EPICAMPES 113

Species IncLtupEep (asterisked species originally described as
Epicampes): M. aurea, M. breviligula, M. distans, M. distichophylla,
*M. emersleyi, *M. gigantea, M. gooddingii, *M. grandis, M. inaequalis,
M. iridifolia, M. lehmanniana, *M. lindheimeri, M. longiglumis,
*M. macrotis, *M. mutica, M. pubescens, M. pubighuma, *M. robusta,
M. scoparia, M. speciosa, M. versicolor, *M. virletiit, M. xanthodas.

Species Intermediate Between Epicampes and Podosemum
M. involuta, *M. longiligula, M. reederorum, M. stricta.

Key to the major groups of Muhlenbergia

Phloem of the primary vascular bundles partially sclerosed; strongly caespitose
perennials; culms erect, simple usually stout and robust; glumes nerveless

or l-nerved ..... toe ee Subgenus Podosemum
Basal sheaths rounded; ligule firm (coriaceous) ; glumes equal or unequal,
shorter to longer than the floret. . . . . . . . Section 1. Podosemum

Basal sheaths compressed-keeled; ligule membranous (sometimes firm below) ;
glumes about equal, generally longer than the floret . Section 2. Epicampes
Phloem of the primary vascular bundles not sclerosed; morphologically different
in one or more of the above characters. . . . . . Subgenus Muhlenbergia

Key to the species of Section Epicampes
(Including species intermediate between sections Epicampes and Podosemum)

Ligule broadly decurrent, firm and brown below, often membranous above;
basal sheaths rounded, compressed, or compressed-keeled; old blades involute
and somewhat arcuate.

Basal sheaths rounded or compressed, but not strongly keeled.
Lemmas long-awned, slightly pubescent on the margins toward the base
(Arizona and Sonora, Mexico) ......... .. 1. M. gooddingii
Lemmas awnless or with an awn no longer than 2 mm., glabrous.

Spikelets mostly 2.5-3 mm. long; blades 3-5.5 mm. wide, flat or inrolled;
basal sheaths rounded (southwestern United States and northern
Mexico). ..... ... .. 2 M. longiligula

Spikelets mostly 3-4 mm. ‘long; blades \- 2 mm. wide, involute; basal
sheaths compressed and somewhat keeled.

Glumes equal to or longer than the floret; lemmas cuspidate or with an
awn to 1 mm. long; spikelets averaging 4 mm. long (Mexico:

Durango). ..... .... . & M. reederorum
Glumes shorter than the floret; ‘lemmas with an awn 1-2 mm. long;
spikelets averaging 3.5 mm. long (Texas) .. . . 4. M. involuta

Basal sheaths strongly compressed-keeled.
Glumes and lemmas glabrous to scaberulous; culms glabrous to scaberulous
below the nodes (Texas and Coahuila, Mexico) . . 5. M. lindheimeri
Glumes and lemmas pubescent to lightly villous; culms pubescent below the
nodes (Mexico: Coahuila and Nuevo Leén) .. . . 6. M. pubigluma

114 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Ligule not broadly decurrent, membranous (rarely somewhat firm below);
basal sheaths strongly compressed-keeled; old blades flat or folded, not
arcuate.

Sheath auricles present, long and pointed, often more than 1 cm. long.

Lemmas mostly glabrous.

Lemmas awnless or cuspidate, glabrous; spikelets mostly 2.5-3 mm. long;
sheath auricles usually 1 cm. or more long (Mexico [Sinaloa to
Oaxaca] and Guatemala). ... . ..... 7. M. macrotis

Lemmas long-awned, glabrous except for short hairs on the margins toward
the base; spikelets mostly 2-2.5 mm. long; sheath auricles usually
less than 1 cm. long (Mexico [Jalisco south] and Guatemala).

8. M. distichophylla

Lemmas villous (Michoaecdn, Mexico, south to Honduras). 9. M. versicolor

Sheath auricles lacking, or, if present, broad and rudimentary, not long and
pointed.

Branches of the very large panicles widely spreading (tending to open after
seed-set), long-naked below; lemmas completely glabrous; ligule usually
about 1 cm. or more long.

Glumes equaling the floret, rounded or acutish; spikelets generally 1.5—2.2

mm. long (Mexico: Durango to Chiapas). . . . . 10. M. gigantea
Glumes several mm. longer than the floret, acute; spikelets generally 2.5-3
mm. long (Mexico: Veracruz and Chiapas). . . . . 11. M. mutica

Branches of the panicle ascending or appressed (usually opening outward
at anthesis, tending to become appressed at seed-set), long-naked below
or floriferous to base; lemmas with some pubescence (except usually
M. robusta, this species with a short ligule unlike that of M. gigantea and
M. mutica).

Glumes awn-tipped.
Lemma short-pilose over the lower half (Colombia and Venezuela).
12. M. inaequalis
Lemma glabrous except for a tuft of short hairs at the base.
Spikelets 1-2 mm. long; glumes unequal; blades narrow, 2-3 mm.;
ligule 8-16 mm. long, hyaline (Mexico: Chihuahua to Michoacan).
13. M. scoparia
Spikelets about 4 mm. long; glumes about equal; blades wider, 4-7
m.; ligule 5-7 mm. long, firm (Mexico: Jalisco).
14. M. iridifolia
Glumes muticous, not awn-tipped.
Basal sheaths, blades, and culms moderately to densely villous.
Ligule about 1 mm. or less long; spikelets averaging 2 mm. long; awns
of lemma 1 cm. or more long (Mexico: Chihuahua to Jalisco).
15. M. speciosa
Ligule 1.5-6 (usually 2-4) mm. long; spikelets mostly 2-3 mm. long;
awn of lemma lacking or, when present, up to 7 mm. long (Mexico:
Chihuahua to Puebla). . ..... =... 16. M. pubescens
Bagal sheaths, blades, and culms glabrous to scabrous, but never villous.
Lemma all or mostly glabrous (the hairs confined to the base of
the margins or rarely present on the midnerve at the base),
awnless or rarely short-awned; plants robust.

Sheath auricles present, about 3 mm. long; ligule mostly 2-3.5 mm.
long; spikelets 2-3 mm. long, ashy gray or purple; lemma
glabrous, the margins rarely pubescent at the base (Mexico
[Nayarit south], Guatemala, and Nicaragua). .17. M.robusta

SODERSTROM—PODOSEMUM AND EPICAMPES 115

Sheath auricles lacking; ligule 4-12 (usually 4-8) mm. long;
spikelets 3-3.6 mm. or more long, dark green; margins of lemma
generally Short-pilose at the base (Mexico: San Luis Potos{to
Puebla) . a .... . 18. M. virletii

Lemma pubescent on the midnerve or r lower part to densely villous
throughout, if glabrous, then long-awned.
Ligule a short rim, generally not over 5 mm. long.

Blade much narrower than the sheath at their juncture; old basal sheaths
conspicuously brown, curled and fibrillose (Guatemala and Honduras).
19. M. breviligula
Blade not much narrower than the sheath at their juncture, old basal sheaths

not conspicuously brown, curled, and fibrillose.
Lemmas moderately pubescent or villous on and between the nerves (Mexico:
Chihuahua to Puebla)... . . ... . . 16. M. pubescens
Lemmas glabrous above, the pubescence confined to the very base of the

margins and midnerve.

Culms glabrous below the nodes; ligule 0.3-1.5 mm. long; spikelets
2-2.2 mm. long; glumes slightly awn-tipped (Guatemala: Quezal-
tenango). ... ... . 20. M. aurea

Culms pubescent below the nodes; ligule 1. 5-6 mm. long; spikelets usually
1.5-2 mm. long; glumes acute or slightly erose (Mexico: Sinaloa to
Jalisco) ... .... . 21. M. grandis

Ligule often 1 cm. or more long, at Teast more than 5 mm. long.
Lemmas pubescent or villous on and between the nerves; spikelets usually

3 mm. or more long, green or greenish brown.

Lemmas densely villous throughout, with awns over 1 em. long; panicles
lead-green or purplish green (Mexico [Michoacdn south] and Honduras).

9. M. versicolor

Lemmas sparsely villous, awnless or short-awned (awns much less than 1 em.
long when present); panicles greenish brown or tan (Mexico [Durango
south] and Guatemala). cone .... . 22. M. distans
Lemmas glabrous or pubescent on the midnerve and/or margins, but not

(or hardly) pubescent between the nerves.

Spikelets 5-7 mm. long; glumes sparsely pubescent, awn-tipped (Mexico:
Jalisco and Aguascalientes)... . ... . . 23. M. longiglumis
Spikelets 2-3 mm. long (rarely longer) ; glumes glabrous, muticous.

Palea pubescent between the keels; callus of floret truncate (southwestern
United States to Oaxaca, Mexico). . ... .. . 24. M. emersleyi

Palea glabrous between the keels; callus of floret truncate or acute.

Glumes glabrous, shining, almost translucent, nerveless or indistinctly
1-nerved; panicle branches short, appressed; callus of floret truncate

(Mexico: Chiapas) . . .... . 25. M. xanthodas

Glumes scabrous, distinctly 1 -nerved; panicle branches long, ascending
or spreading; callus of floret acute (Costa Rica to Colombia).
26. M. lehmanniana

1. Muhlenbergia gooddingii Soderstrom, sp. nov.

Perennis caespitosa, culmis erectis glabris, ca. 60-110 cm. altis,
nodis 4, vaginis inferioribus teretibus compressis vel paullo carinatis
glabris; ligula utrinque decurrente, basin versus firma fuscaque, alibi
membranacea, margine versus apicem lacerata, ca. 15 mm. longa,
laminis arcuatis ad apicem attenuatis, involutis, 50 cm. longis vel

116 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

longioribus, ca. 2 mm. latis, panicula densa nutante straminea vel
purpureo-viridi, axe scaberulo, ramis primariis 3-9.5 (plerumque 3-5)
cm. longis, secundariis plerumque 1-2 cm. longis, tertiariis ad 1 em.
longis, pedicellis scaberulis, 1-1.5 mm. longis, spiculis acutis 2-3 mm.
longis, glumis acutis, nonnullis minute aristatis, plerumque aequalibus,
quam flosculo longioribus, glabris vel scaberulis, nervis 0 vel 1 indis-
tinctis, lemmatibus acutis glabris basin versus margine (et plerumque
in nervo centrali) puberulis, palea glabra.

A M. emersleyi Vasey vaginis magis teretibus, ligula utrinque
decurrente, laminis involutis arcuatisque, lemmatibus minus pubes-
centibus, et palea glabra differt.

Caespitose perennial; culms 60-110 cm. tall; nodes 4, glabrous
below; basal sheaths rounded to strongly compressed, rarely slightly
keeled, glabrous, stramineous, inflated when old; ligule firm, brown,
and broadly decurrent at the base, becoming more membranous and
lacerate above, the firm lower part 4-7 mm. long, the upper membra-
nous part 1 cm. or longer; sheath auricles lacking; blades of the culm
long and narrow, becoming filiform, conduplicate or somewhat. in-
volute when dry, 50 cm. or longer, about 2 mm. wide when unfolded,
upper surface of blade scaberulous or scabrous, the lower surface scab-
rous, the blades when old firm, brown, and strongly arcuate at the base;
upper throat glabrous to scaberulous; collar glabrous,

Panicle narrow, nodding, the branches appressed or ascending,
dense, stramineous or light purplish green, 22-45 cm. long, 2-4 cm.
wide; axis scaberulous; primary branches mostly 3-5 (rarely as much
as 9.5) cm. long, floriferous to near the base, the secondary branches
mostly 1-2 em. long, tertiary branches lacking or, when present, 1
cm. or less long; pedicels 1-1.5 mm. long, scaberulous; spikelets
2-3 mm. long (averaging 2 mm. in the holotype); glumes equal or
the second a little longer than the first, acute, dentate, many bearing
a tiny awn, glabrous or punctate-scaberulous, nerveless or indistinctly
l-nerved; lemma a little shorter than the glumes, acute, mostly
glabrous except for sparse hairs toward the base of the lateral margins
and/or midnerve, 3-nerved, excurrent from below the tip as a flexuous
yellow or purple awn 1 cm. or longer; palea about equal to the lemma,
glabrous or rarely with sparse short hairs between the keels toward
the base.

Holotype in the herbarium of Yale University; collected in canyon
north of Moristo Canyon, Baboquivari Mountains, altitude 4,000
ft., November 19, 1945, by Leslie N. Goodding (No. 462-45). Iso-
types at ARIZ and NY.

DistriBuTion: Restricted in range—collected in mountains near
the Mexican border in the southernmost Arizona counties of Cochise,
Pima, and Santa Cruz, at elevations of 3,000-7,000 ft. Of the two

SODERSTROM—PODOSEMUM AND EPICAMPES 117

additional collections from Sonora, Mexico, one is questionably M.
gooddingir.

UNITED STATES: Arizona: cocHIsE county: West slope of Mule Moun-
tains, Goodding M-345 (ARIZ), M-346 (ARIZ); Garden Canyon, Huachuca
Game Reserve, Goodding 586-49 (ARIZ); Dragoon Mountains, Goodding 50-56
(ARIZ); Mule Mountains, Goodding 307-61 (ARIZ), 451-61 (ARIZ); Fort
Huachuca Military Reservation, Goodding 271-62 (ARIZ). Pima county:
Quinlin Mountains, Goodding 179-45 (ARIZ); Baboquivari Peak, Goodding
281-45 (ARIZ, NY, YU); Moristo Canyon, Baboquivari Mountains, Goodding
408-45 (ARIZ, NY, YU), 457-45 (ARIZ, NY, YU), 460-45 (ARIZ, YU),
468-45 (ARIZ, NY, YU); Canyon N. of Moristo Canyon, Baboquivari
Mountains, Goodding 462-45 (YU holotype, ARIZ, NY, isotypes); Coyote
Mountains, Goodding & Hevly 53-57; (ARIZ); Quinlin Mountains, Goodding &
Lusher 179-45 (NY, YU). SANTA cruz county: Sycamore Canyon, Goodding
CG-18 (US, YU), 2996 (UNM), M-394 (ARIZ, YU), G14-40 (YU).

MEXICO: Sonora: El Cation de la Bellota, Sierra de la Cabellera, Santos 2110
(SMU, TEX, US);(?) N. of Babiacora on road to Cumpas, Wiggins 7393 (ARIZ,
DS, MICH, UC, US).

This species superficially resembles some forms of M. emersleyi, as
most of the collections have been named. It differs in the rounded
or compressed, but rarely keeled, basal sheaths, firm ligules brown
below and membranous above, and long-attenuate blades becoming
threadlike or filiform. The old blades are usually arcuate at the
base. Of the spikelet features to distinguish the species, the lemma
and palea are significant; in M. gooddingii the hairs on the midnerve
and margins are sparse and confined to the lower part and the palea
is glabrous between the keels. The panicles of M. gooddingw are
narrow and densely flowered and the lemmas are always awned.
Such plumelike panicles among the filiform foliage probably create
a distinctive look to this species in the field.

Muhlenbergia gooddingii cannot be called a typical member of section
Epicampes because its features of the firm ligule, brown below, and
the more or less rounded basal sheaths are characteristic of the
Podosemum section.

Because of its combination of characters a possible ancestry is
suspected in M. emersleyi and M. longiligula, for the new species
occurs in the overlap of range of the other two species and M. good-
dingit has features of both. A comparison of many characters from
descriptions in this paper illustrates the intermediate position of
M. gooddingii, as shown below:

M. emersleyi M. longiligula M. gooddingii
culms 1-1.5 (occasionally 60-130 (100) cm. 60-110 cm.
to 2) meters
nodes 3-4, glabrous or 1 or 2 on lower 4, glabrous below
somewhat puberu- part, glabrous

lent below

118

basal
sheaths

ligule

blades

blade surface

old blades

panicle

panicle
dimensions
1° branches
2° branches
3° branches

spikelets

CONTRIBUTIONS FROM THE

M. emersleyi
compressed, keeled,
glabrous

membranous, usual-
ly a little firmer
at base than
above where thin
and usually
lacerated into
narrow strands

2-6 mm. wide, flat
or folded, 23-47
(mostly 20-40)
em. long

upper: glabrous to
scaberulous

lower: scabrous

brown, not especially
firm, not arcuate

typically lax above,
branches loosely
ascending or
spreading; light
purple to pur-
plish tan

20-40 (mostly 30-
40) cm. long, 4-7
em. wide (more
in field)

9-17 (9-12) cm.
long

3-9 (3-5) em. long

0.5-4.5 (0.5-1.5)
cm. long

2.2-3.2 (2.5-2.7)
mm. long

M., longiligula
rounded, rarely

compressed, but

never keeled;
scaberulous;
old sheaths
expanded

firm, brown, de-
current, some-
times becoming
membranous at

tip

long-attenuate to
a brown,
pointed tip,
flat or inrolled
at edges; to 65
(20-40) em.
long, 3-5.5
(usually about
4 mm.) wide

upper: glabrous
to scaberulous
(mostly scaber-
ulous)

lower: mostly
scabrous

brown, firm,
folded and
somewhat
arcuate

erect, narrow
greenish to
purple

15-55 (80-45) em.

long, 1-5 (gen-
erally 1.5-2)
cm. wide
to 13 (generally
3-5) em. long
to 5 (generally
1-2) em. long
lacking

2-3.5 (usually
2.5-3) mm.
long

NATIONAL HERBARIUM

M. gooddingii
rounded to strongly
compressed, rarely
slightly keeled;
glabrous; old
sheaths expanded

firm, brown, broadly
decurrent at base,
becoming mem-
branous and
lacerate above

long, narrow, be-
coming filiform;
conduplicate or
somewhat involute
when dry; 50 em.
long, about 2 mm.
wide

upper: scaberulous
or scabrous

lower: scabrous

firm, brown,
strongly arcuate
at base

nodding, the branches
appressed or as-
cending, dense,
stramineous or
light purplish
green

22-45 cm. long,
2-4 cm. wide

3-5 (to 9.5) em.
long
1-2 em. long

lacking or to 1 cm.
long
2-3 (2) mm. long

SODERSTROM—PODOSEMUM AND EPICAMPES 119

M., emersleyi M. longiligula M., gooddingti
glumes broadly acute acutish, some- acute, dentate,
times midnerve many bearing a
extending into tiny awn
a minute awn
lemma acutish; pubescent glabrous (rarely mostly glabrous
on midnerve and scaberulous) except for sparse
margins 4-4 hairs toward base
their length of margins and/or
midnerve
awn oflemma 0 to 1.5 cm. long rarely a minute 1 cm. or more long
awn present
palea pubescent between glabrous glabrous or rarely
the keels -% with sparse short
their length hairs between the
keels toward the
base

This species, with a few exceptions, has been collected by Mr. Leslie
N. Goodding, a long-time resident of Arizona, staunch conservationist,
and botanist keenly interested in the Arizona flora. It is a pleasure
to recognize Mr. Goodding’s interest in grasses and contributions to
Arizona botany by naming for him this predominantly Arizona species.

2. Muhlenbergia longiligula Hitchc. Amer. Journ. Bot. 21:136. 1934.

Epicampes ligulata Scribn.in Vasey, Contr. U.S. Nat. Herb. 3(1):58. 1892.
Basis of Muhlenbergia longiligula Hitche., not M. ligulata Scribn. and
Merr. (Pringle, Santa Rita Mts., Arizona, July 26, 1884, selected by A. 8S.
Hitchcock as lectotype).

Epicampes anomala Scribn. in Beal, Grasses N. Amer. 2:311. 1896.
(Pringle 1423, Arroyo Ancho, Sierra Madre, Chihuahua, Mexico). Not
Muhlenbergia anomalis Fourn. 1896.

Epicampes distichophylla var. mutica Scribn. in Beal, Grasses N. Amer.
2:308. 1896 (Toumey 740, July 23, 1892, Arizona).

Epicampes stricta var. mutica Jones. Contr. West. Bot. 14:6. 1912. Based
on E. distichophylla var. mutica Scribn.

Strongly caespitose perennial; culms glabrous, becoming scaberulous
toward the panicle, 60-130 cm. (usually about 1 meter) tall; nodes
1 or 2, on the lower part, glabrous; basal sheaths rounded, rarely
compressed, but never keeled, glabrous to scaberulous, loosely im-
bricate, the old sheaths expanded, yellowish brown, persistent; ligule
firm, brown, decurrent, sometimes becoming membranous at the tip,
especially in the uppermost blades of the culm, 1-3 (usually 1-1.5) cm.
long, distinctly 2-keeled when large and broad; sheath auricles lacking;
blades of the culm to 65 (usually 20-40) cm. long, long-attenuate to
a brown, pointed tip, flat or inrolled at the edges, 3-5.5 (usually
about 4) mm. wide when flat, the uppermost blade of the culm beneath
the panicle short, the upper surface of the blade glabrous to scabrous

221-352—67_—4

120 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

(mostly scaberulous), the lower surface mostly scabrous, the old blades
brown, firm, folded, and somewhat arcuate at the point of attachment
to the sheath; upper throat glabrous; collar glabrous (rarely scaberu-
lous).

Panicle erect, narrow, the panicle branches borne in bunches along
the axis, branches of the lowest cluster shorter than the clusters
above, an interruption usually present between the basal and re-
maining clusters, greenish to purple, 15-55 (usually 30-45) em. long,
1-5 (usually 1.5-2) cm. wide; axis glabrous to scabrous (mostly
scabrous), the primary branches to 13 (usually 3-5) em. long, the
secondary branches to 5 (usually 1-2) em. long, bearing spikelets
almost to the point of attachment of the secondary with the primary
branches, tertiary branches lacking; pedicels very short, glabrous to
scaberulous (rarely scabrous) ; spikelets 2-3.5 (mostly 2.5-3) mm. long;
glumes usually longer than the floret, but often shorter, especially in
the longer spikelets, usually equal or the second a little longer than the
first, glabrous to scaberulous, acutish at the tip, sometimes extending
into a minute awn, nerveless or 1-nerved; lemmas 2.5-2.9 mm. long,
glabrous (rarely scaberulous), indistinctly 3-nerved, rarely with a
minute awn; palea equal to or often a bit longer than the lemma,
glabrous.

Lectotype in the U.S. National Herbarium, no. 746686, collected
in the Santa Rita Mountains, Arizona, elev. 4,000 ft., July 26, 1884,
by C. G. Pringle (without number). Hitchcock has written on the
type sheet, ‘Selected as probable type. The Texas specimens are
different and do not agree with the description. Note long firm
ligule.” Isolectotypes at F, GH.

Distrrisution: Rocky mountain slopes and canyons, at elevations
of 5,000-9,000 feet, most frequently found in exposed areas of the
pine-oak zone at 6,000-8,000 feet, occasionally in lightly frosted
areas where more shaded. Rather common from New Mexico to
Arizona into northern Mexico, occurring in Sonora, Chihuahua, and
Durango. Also reported from western Texas and southern Nevada.

UNITED STATES: New Mexico: Mountains W. of Gray, Goodding M-252
(ARIZ, US), M-254 (ARIZ). catron country: Apache National Forest,
Laney 16 (UNM); Solheim & Solheim 2343 (GH). Mangas, Metcalfe, Sept. 14,
1897 (GH, NY, US). Grant county: Black Range, Goodding & Goodding
367 (ARIZ). Santa Rita del Calve, Wright 1973 (CAS, GH, MO, NY, UC,
US). GREENLEE county: Castle Creck, Apache, Utzat 14 (UNM). socorro
counTy: Mogollon Mountains, Metcalfe 556 (ARIZ, GH, MO, NY, POM, US).
ARIZONA: Emersley 47 (US). Upper Oak Creck, Fulton 7345 (ARIZ). Peja
Blanca Mountains, Goodding, Sept. 20, 1935 (ARIZ); Jones, Sept. 20, 1884 (US).
Rocky Canyon, Rothrock, in 1874 (F, GH, NY). Oak Creek Canyon, Weatherwaz
2782 (TAES). apacHE county: Diamond Creek, Gould c& Robinson 5030
ARIZ, NY, TAES). Reservation Ranch, Wasser, Aug. 1936 (ARIZ). cocuise
county: Chiricahua Mountains, Blumer 189 (ARIZ). Barfoot Park, Blumer

SODERSTROM—PODOSEMUM AND EPICAMPES 121

1424 (ARIZ, DS, F, GH, MO, NY, US). Herb Martyr Dam Area, “Chiricahua
Vegetation Team’ 59-803 (ARIZ). Carr Canyon, Darrow, Gould, Phillips, &
Pultz 1487 (MO). Rock Canyon, Huachuca Game Reserve, Goodding 884-49
(ARIZ). Fort Huachuca Military Reservation, Goodding 568-58 (ARIZ, YU),
720-58 (YU). Pinery Pass, Goodding & Goodding 463 (ARIZ, UC). Carr
Canyon, Gould, Darrow, Pultz & Phillips 2461 (CAS, UC, US). Divide between
Carr and Ramsey Canyons, Gould & Haskell 3363 (ARIZ, NY, UC, US). Rucker
Canyon, Gould & Haskell 4585 (ARIZ, GH). Bowie, Jones 4290 (ARIZ, POM,
SMU, US). Huachuca Mountains, Jones, Sept. 3, 1903 (POM); Lemmon &
Lemmon 2921 (GH), Sept. 1882 (US). Pine Canyon, Toumey, Sept. 10, 1896
(US). cocontno county: Long Valley Ranger Station, Coconino National
Forest, Darrow 3262 (ARIZ, CAS, US). Pine Flat Camp, Oak Creek Canyon,
Darrow, Sept. 16, 1943 (ARIZ). atta country: Mt. Ord, Goodding 123-47
(ARIZ). Young, Goodding & Goodding 341 (ARIZ). Long Valley, Pine-
Winslow Road, Goodding & Goodding 354 (ARIZ). 5 mi. N. of McFadden
Peak on Young Road, Gould 3915 (ARIZ, US). Workman Creek Canyon,
Gould & Hudson 3793 (ARIZ, MO, UC, US). Cassadore Range, Gould &
Robinson 4896 (ARIZ, UC). White Mountains, Griffiths 5367 (US), 5409 (GH,
US), 5434 (US). GraHam county: 3 mi. SW. of Point of Pines, 70 miles E. of
San Carlos, Bohrer 421 (ARIZ, CAS). Bluejay Ridge, Gandy 8 (UNM). Mt.
Graham, Hope 9999 (ARIZ, MICH, US). GREENLEE county: Red Bluffs road
above Blue, Goodding & Goodding 349 (ARIZ, US). Big Lue Range, Gould &
Haskell 4064 (ARIZ, UC). 15 mi. N. of Blue, Gould & Robinson 5143 (ARIZ,
TAES, UC). Maricopa county: Sugar Loaf Mt. Wonderland of Rocks, Darrow,
July 8, 1937 (ARIZ). Navaso county: Fort Apache, Mayerhoff 89 (F) ; Schroeder,
in 19387 (ARIZ). Pima county: Mt. Lemmon Road, Benson 9106 (ARIZ, DS,
POM). Manning Camp, Blumer 3412 (ARIZ, DS, GH, MO, UC). Santa
Catalina Mountains, Brown 135 (ARIZ), Darrow, Oct. 3, 1987 (GH). Prison
Road, Ginter, Oct. 6, 1941 (UC). Moristo Canyon, Goodding 451-45 (ARIZ,
NY). Santa Rita Mountains, Griffiths & Thornber 122 (US). Mt. Lemmon,
Bear Canyon, Hevly 1028 (ARIZ). Santa Rita Mountains, Kearney & Peebles
10543 (ARIZ, MICH, US). Bigelow Tower, Leader & Leader 276 (ARIZ).
San Luis Mountains, Mearns 373 (US). Rincon Mountains, Nealley 65 (NY,
US), Aug. 1881 (GH, MO). Upper Bear Canyon, Santa Catalina Mountains,
Parker 7099 (ARIZ). Mt. Lemmon, Peebles & Harrison 2294 (US); Peebles,
Harrison & Kearney 2539 (ARIZ, US). Santa Rita Mountains, Pringle, July
26, 1884 (US lectotype, F, GH, isolectotypes). Madera Canyon, Silveus 3468 (TAKS,
TEX, US). Marshall Gulch, Thornber 2767 (ARIZ). Box Springs, Thornber,
Aug. 10, 1906 (ARIZ). Tucson, Toumey 740 (DS, NY, US, isotypes of £.
distichophylla var. mutica). SANTA CRUZ coUNTY: White House (Madera)
Canyon, Benson 9780 (POM); Gould 2597 (ARIZ, UC, US); Leader & Leader
198 (ARIZ). Sycamore Canyon, Goodding M-111 (ARIZ), M-259 (ARIZ,
Us). Santa Rita Mountains, Griffiths 6074 (MO, US). Santa Catalina
Mountains, Thornber 7689 (ARIZ). YAVAPAI couNTY: Jerome, Benham M-34
ARIZ), M-105 (ARIZ); Goodding 75-46 (ARIZ). Mingus Mountain, Benham
43-B (ARIZ, MICH). Prescott, Purchase 513 (ARIZ).

MEXICO: Sonora: San José Mountains, Marshall 173 (ARIZ); Mearns
996 (US), 1649 (US). El Tigre, Rid de Bavispe, Santos 1963 (ARIZ, GH, MO,
NY, SMU, TEX, US). El Picacho del Pilar, Rio de Bavispe, Santos 2164
(GH, SMU) Curmvanva: Between Rio Chico and Rio Caballo, Barlow, Sept.
30, 1911 (F, UC, US). Sanchez, Hitchcock 7667 (NY, US). Marsh Lake,
Jones, Sept. 19, 1903 (POM). N. de Cd. Madera, Martinez F-2477 (CHAP,
Us). 35 km. SW. of Mifiaca, Pennell 18928 (MEXU). NE. of Colonia Pacheco,

122 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Pennell 19194 (US). Arroyo Ancho, Pringle 1423 (GH, US, isotypes of EF.
anomala) Sierra Madre, Pringle 1427 (F, GH, NY, US), 1703 (UC). 27 mi. NW.
of Cd. Madera, Reeder & Reeder 2663 (YU). Ca. 43 mi. SW. of Casas Grandes,
Reeder & Reeder 2692 (YU). 38 mi. 8. of Casas Grandes, Reeder & Reeder 2699
(YU). 31 mi. W. of Colonia Judrez, Reeder & Reeder 3222 (YU). Palimos
Cafion ca, 13 mi, E. of Chuhuichupa, Reeder, Reeder, & Goodding 1222 (YU).
S. of Hernindez, Reeder, Reeder, & Soderstrom 3505 (US, YU), 3509 (US, YU),
3531 (US, YU). Duranao: 6.5 mi. W. of El Salto, Reeder & Reeder 2639 (YU).
25 mi. W. of Durango, Reeder, Reeder, & Soderstrom 3439 (US, YU).

Muhlenbergia longiligula is distinguished by its long, firm, decurrent
ligule and rounded basal sheaths. On the basis of leaf anatomy, the
long glumes, the dorsal awn (when present), and the general habit in
the field, it seems to belong with the Epicampes section of Muhlen-
bergia. The firm ligule and rounded basal sheaths are also charac-
teristics of species in the Podosemum section. Thus, M. longiligula
possesses characters of both sections, and it appears to be more of a
link between them than a distinct member of either. The closely
allied species of the subgenus Podosemum, M. lindheimeri, M. involuta,
M. pubighuma, and M. reederorum, all possess the decurrent ligule.
Of these M. involuta and M. reederorum, like M. longiligula, lack
strongly compressed-keeled sheaths. All occur within the same general
range, M. longiligula ranging from southwestern United States into
northern Mexico, M. lindheimeri in south-central Texas and Coahuila,
M. wvoluta in Texas, M. pubigluma in Coahuila and Nuevo Leén,
and M. reederorum in Durango, Mexico.

3. Muhlenbergia reederorum Soderstrom, sp. nov.

Perennis caespitosa, culmis angustis scabris, ca. 1 m. altis, basin
versus nodis 3, puberulentis, vaginis inferioribus compressis et paullo
carinatis glabris; collis scaberulis; ligula utrinque decurrente, basin
versus firma et fusca, alibi membranacea et margine versus apicem
lacerata, ca. 10-15 mm. longa; laminis culmi usque ad 65 em. longis
laete viridibus involutis, attenuatis et ad apicem filiformibus, laminis
vetustis fuscis firmis arcuatis, ca. 1.5 mm. latis, utrinque scabris;
panicula erecta, cetera laxa, laete viridi 30-47 cm. longa, quando
appressa 2-3 cm. lata, axe scabro, ramis primariis 4-6 cm. longis,
secundariis ca. 2 cm. longis, tertiariis ca. 5 mm. longis, pedicellis
scaberulis, lateralibus 1.5-2.5 mm. longis, terminalibus 2-3 mm. usque
ad. 6 mm. longis; spiculis lanceolatis 3.5-4.6 (plerumque 4) mm.
longis, glumis acutis fere aequalibus, quam flosculo longioribus,
scaberulis, nervis 0 vel 1, indistinctis, lemmatibus scaberulis, infra
apicem minute aristatis, quam glumis paullo brevioribus.

A M. involuta Swallen vaginis magis carinatis, glumis quam lem-
matibus paullo longioribus, spiculis longioribus, paniculis longioribus,
et aristis quam lemmatibus brevioribus differt.

SODERSTROM—PODOSEMUM AND EPICAMPES 123

Caespitose perennial; culms rather slender, scabrous, erect, about
1 m. tall; nodes 3, puberulent below; basal sheaths compressed and
somewhat keeled or compressed only, glabrous, purplish toward the
base when young; collar scaberulous; ligule decurrent, the lower 14-34
firm and brown, the upper half membranous and lacerate, about 10-15
mm. long; sheath auricles lacking; blades of the culm up to 65 em.
long, light green, tightly folded, long-attenuate becoming filiform,
the old blades brown, firm and arcuate, about 1.5 mm. wide when flat,
the upper and lower surfaces scabrous; upper throat scabrous.

Panicle erect, grayish green, 30-47 cm. long, 2-3 em. wide when
appressed, more open at anthesis; axis scabrous; branches of the
panicle narrow, the primary branches 4-6 cm. long, secondary branches
1-2 cm. long, tertiary branches lacking or about 5 mm. long; pedicels
evenly scaberulous, those of the lateral spikelets 1.5-2.5 mm. long, of
the terminal spikelets 2-3 (to 6) mm. long; spikelets lanceolate,
3.5-4.6 (averaging 4) mm. long; glumes acute, about equal, scaberu-
lous, nerveless or indistinctly 1-nerved; lemmas a little shorter than
the glumes, scaberulous, 3-nerved, bearing a rudimentary awn 0.2-2.5
mm. long from just below the tip; palea glabrous.

Holotype in the Yale University Herbarium, collected in the
Sierra Madre Occidental (W. of Ciudad Durango), 7 miles E. of
Navios, Durango, Mexico, elevation 7,700 ft., Sept. 26, 1963, by
J. R. & C. G. Reeder (no. 3834). Isotypes at F, K, MEXU, MICH,
NY, P, UC, and US.

DistrisuTion: Known from Durango and possibly Jalisco, Mexico.
The type specimen was collected in the Sierra Madre Occidental west
of the city of Durango, where the large caespitose clumps are found
on rocky canyon walls in the pine-oak zone at elevations of 7,700-
8,800 ft.

MEXICO: Durango: 4 mi. W. of Navfos, Reeder & Reeder 4230 (US, YU);
7 mi. E. of Navios, Reeder, Reeder, & Soderstrom 3436 (US, YU). 2 mi. SW. of
Buenos Aires, Reeder, Reeder, & Soderstrom 3347 (YU). (?)Jautisco: Mountains
N. of Autlain, 3-5 mi. above Mina San Francisco, McVaugh 19669 (MICH, US).

Muhlenbergia reederorum exhibits greatest similarity to M. involuta,
known only in Texas. In M. involuta the basal sheaths are compressed,
but not keeled; in M. reederorum they are usually more keeled, but
the keel is not strong. In M. reederorum the glumes are a little longer
than the floret, but much shorter in M. involuta, the spikelets are
longer, averaging 4 mm., the panicle is longer, up to 47 cm., and the
glumes are acute and entire at the apex. Muhlenbergia longiligula is
also similar in many respects but its unawned lemmas and flat, wider
(3-5.5 mm.) blades separate it from the new species. The many
long, narrow blades of M. reederorum produce softer and more delicate

124 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

clumps in the field than the typical coarser clumps of other species of
the Epicampes section. A unicate specimen (Reeder, Reeder, &
Soderstrom 3347) collected near Buenos Aires, Durango, differs from
the material of the type locality in the more membranous ligule, the
longer awn of the lemma (up to 2 mm.), and the wider blades (up to
2.5 mm. wide).

This previously undescribed species is named in honor of Professor
John R. Reeder of Yale University and his wife, Charlotte Goodding
Reeder. Because of their interest in the Gramineae, especially of
Mexico, and the genus MudAlenbergia it is appropriate that a Muhlen-
bergia from that country bear their name.

4. Muhlenbergia involuta Swallen. Amer. Journ. Bot. 19: 436.f. 2. 1932.

Caespitose perennial; culms glabrous or scaberulous at the base,
to 135 (usually 70-100) cm. tall; nodes 4 or more, mostly glabrous
or puberulent beneath; basal sheaths compressed, but not strongly
keeled, tightly imbricate, yellowish brown, not becoming fibrillose
when old; ligule decurrent, firm and brown at the base, becoming
acute and membranous only toward the tip, often firm about three-
quarters of its length, membranous one-quarter of the length, to 1.2
em. long; sheath auricles lacking; blades of the culm to 45 cm. long,
pale green, often arcuate, long-attenuate to a firm point, tightly
folded, 1-2 mm. wide when unfolded, the upper surface of the blade
scabrous, bearing a» mat of numerous tightly appressed white hairs
(spicules), the lower surface of the blade scabrous; upper throat
scabrous; collar scaberulous.

Panicle erect, greenish to tan, to 40 (usually 20-30) em. long, 1.5-7
em. wide, branches of the panicle long; axis scabrous, the primary
branches 6-9 (usually 6-8) cm. long, the secondary branches 1-4
(usually 2-3.5) cm. long, the tertiary branches to about 2 (usually
1 or less) cm. long; pedicels of the terminal spikelets very long, equal-
ing the spikelet or, more often, much longer, pedicels of the lateral
spikelets usually shorter than the spikelet; spikelets 3-4 (averaging
3.5) mm. long; glumes equal or the second 1 mm. or more longer than
the first, slightly toothed at apex, occasionally bearing a rudimentary
awn, usually glabrous but sometimes scaberulous, 1-nerved; lemmas
3.2-4.2 (usually 3.5) mm. long, usually much longer than the glumes,
rarely about equal, glabrous or with a few hairs at the base of the
lateral margins, rather strongly 3-nerved, awned, the awns 1-2.4
(usually 1-2) mm. long; palea about equal to lemma, glabrous.

Type in the U.S. National Herbarium, no. 1501594, collected on
draws in hills, 20 miles NE. of San Antonio, Texas, Oct. 10-31 (year
not cited), by W. A. Silveus (no. 358).

SODERSTROM—PODOSEMUM AND EPICAMPES 125

Disrrispution: Rocky banks and draws of hills, usually on lime-
stone soil. Of very limited distribution, in south-central Texas.

UNITED STATES: Texas: BANDERA COUNTY: Bandera, Silveus 7393 (TEX).
BEXAR CouNTY: Hills NE. of San Antonio, Silveus 314 (TEX, US). 20 mi. NE.
of San Antonio, Silveus 358, = Amer. Gr. Nat. Herb. 1324 (US Holotype, CAS,
GH, LL, MO, SMU, UC, isotypes). BoeRNe country: Open calcareous hillsides,
Palmer 10829, in 1916 (US). BURNET county: S. of Burnet, Silveus 7676 (ARIZ,
CAS, DS, SMU, TAES, TEX). uays counry: W. of Cedar Valley, Edwards
Plateau, Gould 5336 (TAES). KENDALL county: Spanish Pass, Cory 26212
(TAES, US). SW. of Boerne on the Boerne-Bandera road, Silveus 780 (TEX,
US). Comfort, Silveus 2335 (CAS, TAES, TEX), 7339 (CAS, TAKES, TEX).
KERR counTY: NW. of Kerrville, Reed, Sept. 28, 1937 (US). TRAVIS couUNTY:
W. of Trading Post on cut off road to Pedernales Bridge, Brown 50-323 (SMU,
TAES, TEX). Glen Rose, W. of Austin, Moon 167 (TEX). Austin, Tharp 3968
(TEX), 5198 (TEX). Pedernales road, 8 mi. W. of Bee Cave, Tharp 53-250 (TEX).
W. of Austin, Warnock 45-27 (TEX, US). Hamilton Pool, Webster 1961 (TEX).

Muhlenbergia involuta is distinguished by its firm, decurrent ligule, in
combination with the compressed but not keeled basal sheaths, the
tightly folded arcuate blades, and glabrous lemmas that usually exceed
the glumes in length. Swallen, in describing this species, remarked,
“Muhlenbergia involuta is related to M. distichophylla (Presl) Kunth
but in that species the spikelets are only 1.5-2 mm. long, the awns are
10-15 mm. long, the blades are flat, as much as 5 mm. wide, and the
sheaths bear auricles 2-3 cm. long. In both species the glumes are
shorter than the floret.” However, AZ. involuta, is by no means closely
related to M. distichophylla. The latter is allied to M. macrotis and M.
robusta in which the sheaths bear rudimentary to very long auricles,
the lemmas are glabrous, and the ligule membranaceous throughout.
In M. involuta the ligule is decurrent, very firm and brown at the base,
becoming membranaceous only at the tip. This character, in addition
to the pale green, firm, tightly folded arcuate blades, indicates that it
is closer to M. lindheimeri, M. pubigluma, and M. reederorum. Muh-
lenbergia involuta is endemic to Texas, M. lindheimeri to Texas and
Coahuila, AL, pubigluma is known only from Coahuila and Nuevo
Leén in northern Mexico, and M. reederorum only from Durango (and
possibly Jalisco), Mexico. The distribution of these species in Texas,
Coahuila, and Durango separates them from M. distichophylla from
farther south, Jalisco, Mexico, into Guatemala.

5. Muhlenbergia lindheimeri Hitchc. Journ. Washington Acad. Sci. 24(7):291.
1934.

Epicampes gracilis Trin. Mem. Acad. St. Petersb. VI. Sci. Nat. 4:271. 1841.
Fragments of panicle of this specimen ex Trinius herbarium (LE), in type
collection at the U.S. National Herbarium marked ‘“‘Epicampes gracilis in
‘Mexico de Karwinsky.’ Hb. Reg. Monac (Munich) no. 595.” The
published locality of this specimen is Mexico, but the species is known only

126 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

from south-central Texas. This reference is doubtless made to territory
now Texas but which was still part of Mexico in 1841. Not Muhlenbergia
gracilis (H. B. K.) Kunth. 1829.

Strongly caespitose perennial; culms stout, robust, glabrous, 0.5-1.5
(mostly more than 1) meters tall; nodes 4, glabrous beneath the nodes;
basal sheaths compressed-keeled, glabrous, tightly imbricate and
usually yellowish or purplish brown; ligule decurrent, firm and brown
at the base, becoming membranous above, usually enclosed within the
tightly folded blade, up to 3.5 (mostly 1-1.5) cm. long; sheath auricles
lacking; blades of the culm 25-55 (usually 30-45) cm. long, narrow,
pale green, very firm and conduplicate, 2-4 (usually 2-3) mm. wide
when flat, the upper surface of the blade scabrous, bearing a tight mat
of dense short white hairs, the lower surface scaberulous to scabrous;
upper throat scabrous, bearing a dense mat of short white hairs; collar
glabrous to scaberulous.

Panicle erect, narrow, pale green or tan, the panicle branches
floriferous almost to the base, occurring in bunches at intervals along
the axis, the naked portion of the axis sometimes obvious, sometimes
obscured by the overlapping appressed panicle branches, to 50
(mostly 20-30) em. long, 1-3 (mostly 1.5-2) em. wide; axis scabrous,
the primary branches to 5 (mostly 2-4) em. long, the secondary
branches to 1.5 (mostly 0.5-1.5) em. long, tertiary branches lacking;
pedicels of the lateral spikelets to 0.5-1.2 (usually 0.5-0.8) mm. long,
those of the terminal spikelets to 3 mm. long, scabrous; spikelets
2-3.5 (usually 2.5-3) mm. long; glumes about equal to (sometimes
shorter than) the floret, the second a little longer than the first,
glabrous to scabrous (mostly scabrous), rounded at the tip or blunt,
rarely acute except in the longer spikelets, awnless, strongly 1-nerved;
lemmas to 3 (mostly 2.4-2.6) mm. long, glabrous or rarely puberulent
at the base, mostly rounded at the tip, 3-nerved (the nerves purple),
awnless or rarely with awns as much as 4 mm. in length; palea a
little shorter than the lemma, glabrous or occasionally with a few
minute hairs at the base between the purplish keels.

Holotype in the U.S. National Herbarium, no. 998949, collected
in 1847 by Ferdinand Lindheimer in Texas. Only the number 725 is
given, this being the number of the “Plantae Lindheimerianae”
series. According to Blankinship (1907) this represents Lindheimer’s
own collecting number of 465 (fasc. IV) and was collected in Fried-
erichsburg (now Fredericksburg), Gillespie County, Texas, in
September 1847. Isotypes at F, GH, MO, and UC.

DistRIBuTION: Dry sandy or limestone soil, in draws of hills and
open areas. Of limited distribution in south-central Texas and
Coahuila, Mexico.

SODERSTROM—PODOSEMUM AND EPICAMPES 127

UNITED STATES: Texas: Southern Texas, Lindheimer, in 1849-51 (NY);
Nealley, in 1889 (GH). Foothills, upper Seco, Reverchon 1610 (F, MO, NY,
TEX, US). Glen Rose Hills, Tharp 46293 (TAKES, UC), Tharp, Dec. 7, 1928
(MO, NY, SMU). BanpERa county: N. of Medina, Parks & Cory 24422
(TAES). W.-NW. of Bandera, Shinners 16880 (SMU). BELL county: Max-
dale, Wolff 2632 (TAES, US). Bexar county: Onion Creek, near San Antonio,
Brown 3487 (TEX). Pleasonton Road 15 mi. S. of San Antonio, Burr 513
(NY). Cibolo Creek, Liles 17 (TAES). North Loop, Silveus 11 (TEX, US),
355 (TEX, US). 20 mi. NE. of San Antonio, Silveus 354 (MICH, TEX, US).
BURNET COUNTY: NE. of Lake Buchanan toward Lampasas, Whitehurst, Nov. 21,
1956 (LL). comMat county: Comanche Spring, New Braunfels, etc., Lindheimer
1255 (ARIZ, F, GH, NY, TEX, UC, US). comancue county: Comanche
Spring, Lindheimer 176 (CAS, ILL, MO, TAES). epwarps county: Barksdale,
Palmer 11004, in 1916 (MO, US); Parks & Cory 26893 (TAES). GILLESPIE
county: Fredericksburg, Thurber 68 (NY). Lindheimer 465, = Plantae Lind-
hetmertanae 725 (US holotype, F, GH, MO, UC, isotypes). HAYS COUNTY:
Reagan Houston Ranch, Johnson 360 (TEX). KENDALL COUNTY: N. of Comfort,
Cory 20732 (GH, UC); Parks & Cory 20729 (TAKES), 20730 (TAES), 20731
(TAES). Spanish Pass, Palmer 10859, in 1916 (US). Comfort, Silveus 7334
(CAS, DS, SMU, TAES, TEX). xerr county: Kerrville State Park, Cory
50506 (DS, GH, NY, SMU, TAES, TEX, UC, US). 25 mi. NW. of Kerrville,
Gould 8468 (SMU, TAES, UC). SW. of Kerrville, Parks & Cory 24840 (TAES).
KIMBLE CouNTY: NE. of junction on Mason Highway, Landers 5223 (TAES).
KINNEY county: W. of Bracketville, Correll & Correll 24740 (LL). MASON
county: N. of Mason, Reardon 23 (TAES). reau county: W. Fork of West
Frio River, Cory 43240 (SMU, TEX). N. of Leakey, Parks & Cory 27385
(TAES). tTRavis county: Austin, Bodin, Oct. 20, 1891 (DS). Oak Hill,
Rogers 6520 (TEX). Barton Creek, Austin, Tharp 70 (TEX, US). Edwards
Plateau, Tharp 3076 (US). Watkins Ranch, Tharp 49009 (US); Tharp, Rogers,
Wilkins, Cowan, Clark & Jeffrey 49004 (TEX, US); Tharp et al. 51-516 (TEX).
Austin, Warnock W1065 (TEX, US). witutamMson county: Georgetown,
Bodin 287 (NY).

MEXICO: Coauurta: About 24.5 mi. SW. of Piedras Negras, Reeder & Reeder
3921 (US, YU). 8 mi. W. of Cuatro Cienegas, Reeder & Reeder 3945 (US,
YU).

Muhlenbergia lindheimeri is distinguished by the decurrent ligule,
long spikelets with glabrous lemmas, the glumes equal the floret,
and the long-attenuate firm blades somewhat arcuate when old.
It appears to be most closely related to M. involuta and M. pubigluma.
The former differs in having basal sheaths somewhat compressed but
not keeled and glumes shorter than the floret, the latter in having
puberulent glumes and lightly villous lemmas.

6. Muhlenbergia pubigluma Swallen. Proc. Biol. Soc. Wash. 56:78. 1943.

Strongly caespitose perennial; culms stout, glabrous, about 75-125
em. tall; nodes 3, pubescent below; basal sheaths compressed—
keeled, glabrous, tightly imbricate, the old basal sheaths firm, dark
brown, not becoming fibrillose; ligule decurrent, firm and brown at
the base, becoming membranous for the greater part of its length,
5-13 mm. long, longer on the innovations; sheath auricles lacking;

128 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

blades of the culm to 35 em. long, narrow, pale green, firm and
conduplicate, long-attenuate to a firm brown point, 1-2 mm. wide
when flat, the upper surface of the blade scabrous, bearing a tight
dense mat of tiny white hairs, the lower surface scabrous; upper
throat bearing a dense mat of short appressed hairs; collar scaberulous.

Panicle erect, narrow, the branches appressed, interrupted below,
olive to grayish green, 20-34 em. long, 1-2 cm. wide; axis scabrous;
lower primary branches 4-5 cm. long, the secondary branches to
1.5 em. long, tertiary branches lacking; pedicels much shorter than
the spikelets, scabrous; spikelets 2.5-3.5 (averaging 3) mm. long;
elumes rounded at the tips, mostly longer than the floret, the second
a little longer than the first, hyaline, nerveless or indistinctly 1-nerved,
lightly pubescent, awnless; lemmas 2.9-3.1 mm. long, acute at the
tip, indistinctly 3-nerved, awns 0.5-3.5 mm. long, the lower two-
thirds lightly villous, the upper one-third scabrous, the basal margins
more densely villous; palea a little shorter than the lemma, pubescent
on the lower two-thirds between the keels.

Holotype in the U.S. National Herbarium, no. 2209360, collected
in shrub zones of lower canyon where common, Sierra de la Madera,
Cafion del Agua, Municipio de Cuatro Cienegas, Sept. 10, 1939, by
C. H. Muller (no. 3264).

DisrripuTion: Known only from five collections made in canyons
of Coahuila and Nuevo Leon, northern Mexico.

MEXICO: Coanvuiva: La Cueva, Corte Blanco fork of Charretera Canyon,
Johnston 9072 (GH, MEXU, MO). Cafion del Agua, Municipio de Cuatro
Cienegas, Muller 3264 (US holotype, GH, UC, isotypes). San Lorenzo Canyon,
6 mi. SE. of Saltillo, Palmer 401, in 1904 (F, GH, MO, NY, UC, US). Nurvo
Leén: Between La Soledad and Escondida, Beetle M-406 (UC). 14 mi. W. of
Doctor Arroyo, Shreve & Tinkham 9651 (GH).

Muhlenbergia pubigluma is distinguished by the broadly decurrent
ligule, puberulent glumes and lightly villous lemmas. In the original
description, the author related this species to M. pubescens (H .B.K.)
Hitche. (=M. distans Swallen, according to the present treatment).
That species, however, does not have the narrow, firm, pale green,
somewhat arcuate blades or broadly decurrent, firm at the base,
ligules, characters that indicate a closer relationship to AM. lindheimert
and M. involuta.

7. Muhlenbergia macrotis (Piper) IHlitche. N. Amer. FI. 17(6):463. 1935.

Muchlenbergia distichophylla var. mutica Scribn. ex Urbina. Cat. Pl. Mex. 393.
1897. Nomen nudum. Of the two collections listed, Pringle 2360= M.
macrotis, and Pringle 5577=M. versicolor.

Epicampes macrotis Piper. Proce. Biol. Soc. Washington 18:144. 1905.
Basis of Muhlenbergia macrotis (Piper) Hitche. (Rose 3528, Zacatecas,
Mexico.)

SODERSTROM—PODOSEMUM AND EPICAMPES 129

Epicampes minutiflora Mez. Repert. Sp. Nov. Fedde 17:212. 1921. Basis
of Muhlenbergia meziana Hitche. 1935, not of M. minutiflora (Michx.)
Hitche. 1896. (Langlassé 750, El Canizal, ‘“‘Michoacdén et Guerrero,”
Mexico.)

Muhlenbergia meziana Hitche. N. Amer. FI. 17(6):461. 1935. Based on
Epicampes minutiflora Mez.

Strongly caespitose perennial; culms narrow to very thick, glabrous
to scaberulous, 14-24% meters tall; nodes 3-5, glabrous, or sometimes
pubescent below the nodes; basal sheaths compressed-keeled, gla-
brous, the old basal sheaths brown, persistent, becoming frayed;
ligule membranous throughout, lacerate from the base or near the
base, 3-15 (usually 5-8) mm. long; sheath auricles 1-10 (usually
about 3) em. long, the auricles of the sheaths higher on the culm
often longer than those of the lower ones, straight or strongly twisted;
blades of the culm very long, as much as 95 em., becoming fine and
threadlike at the tip, mostly folded, 2-6 mm. wide when flat, the
blades at the base narrower than the sheaths, edges of the blades
saw-toothed, the upper surface scaberulous to scabrous (mostly scab-
rous), the lower surface scabrous; upper throat scabrous; collar
glabrous.

Panicle erect, columnar, the branches humerous, appressed, gen-
erally interrupted below, light green to whitish green, 25-65 cm.
long, 1.5—6 em. wide; axis scaberulous to scabrous (mostly scabrous),
the primary branches 5-12 em. long, the secondary branches to 4.5
cm. long and spikelet-bearing to the base, the tertiary branches to
1 cm. in length; pedicels 0.3-1 mm. long, shorter than the spikelets,
scabrous; spikelets 2-3 mm. long, rarely as short as 1.5 mm. or longer
than 3 mm.; glumes equal, usually longer than the floret, glabrous to
scaberulous (mostly scaberulous), rounded at the tips, awnless, hya-
line, white or whitish green, nerveless or obscurely 1-nerved; lemmas
1.8-2.5 mm. long, acutish, indistinctly 3-nerved, awnless or rarely
cuspidate, completely glabrous, even on the lower margins, at most
the surface punctate; palea equaling the lemma or rarely a little
longer, glabrous.

Holotype in the U.S. National Herbarium, no. 302505, collected in
the Sierra Madre, Zacatecas, Mexico, Aug. 17, 1897, by J. N. Rose
(no. 3528).

Distrisution: Moist shaded canyon slopes, ravines, banks above
streambeds, moist ditches, and wooded hillsides, at elevations of
6,000-7,000 feet. Common from Sinaloa and Durango, south to
Guerrero and Oaxaca, Mexico. A single specimen has been collected
in Guatemala.

MEXICO: Sinatoa: Mazatlin, Ortega 5075 (US). Duranao: Sierra Tres
Picos, Gentry 5312 (ARIZ, DS, MO, NY); Harvey 5312 (GH). 23 mi. SW. of

130 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Buenos Aires, Reeder & Reeder 4218 (US, YU). Zacatecas: Sierra Madre,
Rose 3528 (US holotype). Nayarit: La Barranca, Jones 23445 (CAS, MO, NY,
POM). Stnatoa-Nayarir: San Ignacio, San Javier, Ortega 1204 (MEXU).
Jauisco: Tepatitlin-Guadalajara, Herndndez, Ruppert, & Guevara X-2669 (US).
SW. Autlin, Hernandez X. 4641 (CHAP). La Barranca, Guadalajara, Jones
27656 (POM), 27677 (DS, MO, NY). Orendain, Jones 2767 (UC). Nevado de
Colima, McVaugh 10170 (MICH). Sierra del Tigre, 3 mi. 8. of Mazamitla,
McVaugh 13085 (SMU, US). 1-2 mi. E. of Tapalpa, Mc Vaugh 20519 (MICH,
US). Real Alto, Trail to Tranquillas, Mexia 1718 (CAS, DS, F, GH, MICH,
MO, NY, UC, US). Guadalajara, Pringle 2360 (ARIZ, CAS, F, GH, MICH,
NY, TAES, UC, US), 2356 (F, GH, MO, NY, UC, US); Soderstrom 613 (US),
614 (US). Guanasuato: Duges, in 1897 (F). Ca. 8 mi. W. of San Felipe,
Cerro del Fraile, Sohns 452 (MICH, MO, US). Pussta: Mexique-Acatzingo,
Distriet de Tepeaca, Nicolas, July 7, 1909 (F). Distrito FrpErat: C. de
Progreso, Matuda 19676 (MICH). Micnoackn: Quinceo, Arséne 3218 (GH,
ILL, MO, NY, US). Loma Santa Marfa, Arstne 8581 (MO, US). Cerro San
Miguel, Arsene 9937 (MO, US). Atzimba National Park, Beetle M-128 (US).
Zacapu, Hernandez, Ruppert, & Guevara X-2822 (US). 18 mi. E. of Morelia,
King & Soderstrom 5011 (MEXU, MICH, SMU, TEX, US). 8-10 mi. NW. of
C. Hidalgo, McVaugh 9925 (MICH, US). Patzcuaro, Seler 1227 (GH, US).
Quiroga, Soderstrom 552 (US), 555 (US) ; Sohns 806 (US). GurErreERO: Petlacala,
Meria 9098 (CAS, F, GH, MO, NY, UC, US). Between Ayusinapa and Petatlin,
Nelson 2122 (US). “Micnoackn er Guerrero”: El Canizal, Langlassé 750
(GH, US, isotypes of M. meziana). Oaxaca: Rio Grande (N. of Niltepec) to
Finea “La Gloria,” Herndndez & Sharp X-1257 (US).

GUATEMALA: Hvurnurrenanco: 10 km. W. of Aguacatdin, Williams,
Molina, & Williams 21844 (US).

Muhlenbergia macrotis is distinguished by the prominent sheath
auricles that attain a length of 1 em. or more (as much as 10 em. in
Hernéndez X-4641!) and in the mostly glabrous and awnless or
cuspidate lemmas. It is closely allied to M. distichophylla which
has long auricles, but the lemmas bear a tiny tuft of hairs toward
the base of the margins and are usually long-awned. The culms
and panicles of MM. macrotis are usually thicker and more robust
than in M. distichophylla. Comparison may also be made with M7.
robusta, but that species has only rudimentary sheath auricles and
the lemmas, although mostly glabrous, sometimes have sparse short
hairs at the base.

Muhlenbergia macrotis was originally described by Piper in 1905 as
an Epicampes from a specimen collected by Rose in Zacatecas,
Mexico, in1897. It was known only from the type collection. Hiteh-
cock transferred EF. macrotis to Muhlenbergia in 1935, at the same
time he transferred E. minutiflorus to the genus naming the latter
M. meziana.

Muhlenbergia meziana Hitche. is considered as synonymous with
M. macrotis in the present treatment. The original description of
the former characterizes a specimen with very long auricles (to 45 mm.

SODERSTROM—PODOSEMUM AND EPICAMPES 131

in length), flat blades to 4 mm. in width, a dense panicle, and spike-
lets 2.5-2.75 mm. long.

The collection listed by Mez in his description of Epicampes
minutiflorus is “E. Langlassé, bei El Canizal, Michoacén (Mexico),”
without collection number. The specimen in the U.S. National
Herbarium marked as the type is E. Langlassé, no. 750, El Canizal,
Michoacén. This specimen, however, does not agree in all respects
with the original description, for the auricles are less than 1 cm. long
and the spikelets are 2-2.5 (averaging 2) mm. long. This leaves some
doubt as to the collection Mez had in mind when he described F.
minutiflorus. Nevertheless, the completely glabrous lemmas, the
white, almost translucent glumes, and relatively long auricles place
it with other specimens of M. meziana. The type appears, therefore,
to be an aberrant representative of the species.

The holotype of M. macrotis agrees with M. meziana, with the
exceptions that the spikelets are often a little longer and the florets
exceed the glumes in length. The glumes, however, are 2-2.5 mm.
long, rounded, almost translucent, and nerveless, like those of M.
meziana specimens. Many of the spikelets in the panicle of the holo-
type of M. macrotis are diseased. I have noticed that in diseased
panicles of other Muhlenbergia species the florets often become longer
than normal, exceeding the glumes where the florets were normally
contained within them. This perhaps explains why many of the
florets are so long in the spikelets of the type specimen of M. macrotis
which, in other respects, is similar to specimens of M. meziana.
Specimens similar in general to the type collection of Muhlenbergia
meziana occur in the same geographic region where the type of M.
macrotis was found, and therefore it seems likely that the holotype
of M. macrotis was only a diseased specimen of the species later
described as M. meziana. The following list, a comparison of the
major characters of specimens of M. meziana and the holotype of
M. macrotis, illustrates the similarity.

M. meziana M. macrotis
height 125-230 cm. 134 cm.
culm surface mostly glabrous glabrous
node surface glabrous to slightly glabrous

pubescent
sheath surface glabrous glabrous
upper throat scabrous scabrous
collar glabrous to scaberulous glabrous
blade width 2-6 (mostly 3-4) mm. 4-5 mm.
upper surface of blade — scaberulous to scabrous scaberulous
(mostly scabrous)
lower surface of blade scabrous scabrous

ligule length 3-13.5 (usually 5-8) mm. 11 mm.

132 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

M, meziana M. macrotis
auricle length 1-7 (usually about 3) cm. to 3.5 and 4 cm.
panicle length 26-65 cm. 38 em.
panicle width 1.5-6 cm. 1.5 cm.
panicle color light green or whitish greenish tan (old panicle

green only)
axis surface scaberulous to scabrous scabrous
length of panicle
branches:
primary 5-11.8 cm. 7 cm.
secondary 1-4.5 cm. 2 cm.
tertiary to ca. 1 cm. lacking
spikelet length 2-3 mm. 2.5-3.6 mm.
glumes equal, sometimes less than — equal, usually less than
floret floret
glume length 2-3 mm. 2-2.5 mm.
glume surface glabrous, nerveless or glabrous, nerveless
rarely obscurely 1-nerved
lemma length 1.8-2.3 mm. ca. 3.2 mm.
lemma surface glabrous glabrous
awn length usually lacking, lemma lacking

rarely cuspidate

8. Muhlenbergia distichophylla (J. 5. Presl) Kunth. Rev. Gram. Supp. xvi. 1830.
Podosaemum distichophyllum J. 8. Presl. Rel. Haenk. I: 231. 1830. (Mexico:
without precise locality, Haenke 173).
Epicampes stricta var. distichophylla (J. S. Presl) Jones, Contr. West. Bot.
14:6. 1912. Based on Podosaemum distichophyllum J. 5. Presl.
Muhlenbergia angustifolia Swallen. N. Amer, Fl. 17(6): 457.1935. (Mexico:
Jalisco, Pringle 2346).

Caespitose perennial; culms stout, glabrous, 1-1.75 m. tall; nodes 4,
elabrous or somewhat puberulent beneath ; basal sheaths compressed-
keeled, mostly glabrous, sometimes scaberulous; ligule membranous
throughout, entire at the base and lacerate above, or finely lacerate
throughout, 4-8 mm. long, longer on the innovations; sheath auricles
4-8 (usually 6-8) mm. long, the auricles of the upper culm sheaths
longer than those of the lower; blades of the culm to 90 em. long, long-
attenuate, becoming threadlike at the tip, flat or folded, 1.5-4.5
(mostly 2-2.5) mm. wide when unfolded, narrow in comparison with
the sheaths from which they depart, the upper surface of the blade
scaberulous to scabrous, lower surface scabrous, the edges of the blades
minutely saw-toothed; upper throat scabrous; collar glabrous to scab-
rous (mostly scaberulous).

Panicle erect, somewhat lax at the tip, light green to tan (rarely
purplish), the branches spreading widely, 35-60 cm. long, to 5 cm. or
more in width; axis scaberulous to scabrous (mostly scabrous), the
lower primary branches 6-13 cm. long, the secondary branches 1-6
cm. long, the tertiary branches 0.5-2 em. long; pedicles of the lateral

SODERSTROM—PODOSEMUM AND EPICAMPES 133

spikelets 0.2-0.8 mm. long, those of the terminal spikelets to 2 mm.,
glabrous to scaberulous (usually scaberulous); spikelets 1,5-2.5
(usually 1.7-2) mm. long; glumes longer than the floret, the first
glume often a little longer than the second, glabrous to scaberulous,
broadly acute, awnless, hyaline, nerveless; lemma 1.4—2.5 (usually
1.7-2) mm. long, glabrous throughout except for a tuft of short hairs
at the base of the margins, rounded at the tip, 3-nerved, the middle
nerve becoming purplish and thickened toward the tip, excurrent as an
awn 4-16 mm. (usually 1 em. or more) in length; palea equal to or
slightly shorter than the lemma, glabrous.

Fragment of type in the U.S. National Herbarium, collected in
Mexico, without precise locality, by Haenke (no. 173).

Disrrisution: Rocky hills and exposed areas of oak woods, at ele-
vations of 1,500-7,500 ft., from Jalisco, Mexico, southeastward into
Guatemala. Apparently infrequent.

MEXICO: Haenke 173 (US fragment of type of Podosaemum distichophyllum,
ex W). Jaisco: Barranca de Oblatos, Hitchcock 7335 (NY, US). SW. of La
Resolana, road to La Huerta, McVaugh 21106 (MICH, US). Above (N. of) La
Cuesta, road to Talpa de Allende, McVaugh 21224 (MICH, US), 21225 (MICH,
US). Guadalajara, Pringle 2346 (GH, NY, UC, US, isotypes of M. angustifolia).
Mexico: Tejupilco, Temascaltepec, Hinton 2719 (GH, LL, NY, US). Luvianos,
Temasealtepec, Hinton 5302 (ARIZ, NY, US). Amatepec, Matuda 29874 (US),
29875 (US). “Micnoacan rr GuERRERO”: Sierra Madre, Langlassé 607 (GH,
US). Cerro Mamegal, Langlassé 674 (GH, US). Guerrero: Carboneras-Filo
Mayor, Hinton 9910 (MICH, US). Vallecitos, Hinton 11635 (ARIZ, LL, MO,
NY, US). 15 mi. N. of Tierra Colorada, Reeder & Reeder 4159 (US, YU).
Curapas: Escuintla, Matuda 319 (MICH, US). Mt. Ovando, Matuda 322
(MICH, MO, US).

GUATEMALA: Hurnvurrenanao: E. of San Sebastian, Standley 81480 (US).
Near crossing of Rio San Juan Ixtdn, E. of San Rafael Pétzal, Standley 83016
(US). W. of Huehuetenango, Williams, Molina, & Williams 22309 (US).

Mulhlenbergia distichophylla is distinguished by long sheath auricles
and awned lemmas that are glabrous except for a small tuft of hairs on
the margins toward the base. It is apparently most closely allied to
M. macrotis, also possessing long sheath auricles, but lemmas that are
awnless and entirely glabrous. Moreover, it has a denser panicle than
M. distichophylla and a more robust habit. Label data for M. dis-
tichophylla indicates flowering from the end of November through
May; M. macrotis blooms earlier.

Although M. angustifolia Swallen was recognized in North American
Flora (1935) and has been maintained as distinct in the USS. National
Herbarium, it is represented only by specimens of the original collec-
tion of 1889. Additional collections of M. distichophylla have been
made since the description of M. angustifolia, and a critical comparison
of all specimens with the holotype of the latter revealed that the char-
acters that distinguish it are not of sufficient importance for specific

134

recognition. The following comparison of the two illustrates this

point.

height
culm surface
node surface

sheath surface
upper throat
collar

blade width

blade length

upper surface of blade
lower surface of blade
ligule texture

ligule length

auricle length

panicle length

panicle width

axis surface

panicle color
panicle branches:
primary, length
secondary, length
tertiary, length
spikelet length
glume surface
lemma length
lemma surface

awn of lemma, length

9. Muhlenbergia versicolor Swallen.

M. distichophylla specimens
135-173 cm.
glabrous
glabrous or puberulent
below
usually glabrous
scabrous
scaberulous to scabrous
(rarely glabrous)
1.5-4.5 mm.
37-91 cm.
scaberulous to scabrous
scabrous
membranous, lacerate
4-8 mm., sometimes more
0.2—4.5 em.
30-58 cm.
usually 3.5-5 em.
scaberulous to scabrous
(mostly scabrous)
greenish tan

6-12 cm.
1.6 cm.

0.2 cm.
1.5-2.5 mm.
glabrous
1.4-2 mm,

glabrous except for hairs toward the base of the margins

(in both)
4.5-16 mm.

Contr. U.S. Nat. Herb. 29: 412.

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

M. angustifolia holotype

111-139 cm.
glabrous
glabrous to puberulent

below
glabrous
scaberulous or scabrous
glabrous

1.5-2 mm.

48 cm.

glabrous or scaberulous
secaberulous to scabrous
membranous, lacerate
6-14 mm.

1.7-3 cm.

30-48 em.

2-2.5 em.

scabrous

light green

4-6.5 cm.
1.5 cm.
lacking
2.2-2.5 mm.
glabrous
2.1-2.3 mm.

8-9 mm.

1950.

Muehlenbergia distichophylla var. mutica Scribn. ex Urbina. Cat. Pl. Mex.
393. 1897. Nomennudum. Of the two collections listed, Pringle, 2360 =
M. macrotis, and Pringle 5577 = M. versicolor.

Caespitose perennial; culms glabrous or somewhat puberulent,
82-199 (mostly 100-150) em. tall; nodes 3 or 4, glabrous or pubescent
below; basal sheaths compressed, but not strongly keeled, the old
sheaths brown and somewhat fibrillose; ligule membranous throughout,
5-22 (usually more than 10) mm. long, usually flanked on either side
by auricles that become membranous and twisted above, the firm,
brown base of such auricles usually 4-8 mm. long; blades of the
middle of the culm shorter than those of the lower part, 12-27 cm.
long, usually folded, 2-5 mm. wide when flat; upper surface of the
blade scaberulous to scabrous, the lower surface scabrous (lower
surface usually scabrous and the upper surface scaberulous) ; upper
throat scaberulous to scabrous; collar glabrous to puberulent.

SODERSTROM—PODOSEMUM AND EPICAMPES 135

Panicle erect, narrow, the branches ascending, lead green to purplish
green, 17-60 (usually 20-40) em. long, 1.5-6 em. wide; axis scaberulous
to scabrous; primary branches 4-11.5 cm., secondary branches
1.5-3.5 cm. long, tertiary branches lacking or to 0.5 em. long, when
present; pedicels of the lateral spikelets 1-2.5 mm. long, those of the
terminal spikelets relatively long, to 3-5 mm., the pedicels glabrous
on the lower part, becoming scaberulous on the thickened part below
the spikelet; spikelets 2.5-3.5 (usually 3 or more, rarely to 4) mm.
long; glumes usually a little shorter than the floret, the second glume
Jonger than the first, the tips erose (under high power) or awn-tipped,
the awns to 1.2 mm. long, frequently only the second glime awned
and the first acute, usually scaberulous (rarely glabrous), hyaline,
nerveless or faintly 1-nerved; lemmas acute, 2.5-3.5 (mostly about 3)
mm. long, 3-nerved, villous over the lower half or two-thirds of the
back on and between the nerves, and more densely so on the margins,
awn 17-30 mm. long, flexuous; callus of lemma long-pilose; palea
equaling or a little longer than the lemma, moderately to densely
villous between and, sometimes, on the keels.

Holotype in the U.S. National Herbarium, no. 1961991, collected
about 170 kilometers north of Oaxaca City, Oaxaca, Mexico, Decem-
ber 13, 1945, by Efraim Herndndez Xolocotzi and J. A. Jenkins
(no. X-810).

DistrisuTIon: Rocky mountainsides and ledges, and slopes of
barrancas. Rather uncommon, ranging in Mexico from Michoacdén
and the state of Mexico southeastward to Honduras.

MEXICO: Guanasvato: NE. of Cd. Guanajuato, Reeder & Reeder 3095
(YU. Pursua: Zapotitldn de las Salinas, SE. of Techuacdn, Herndndez,
Oct. 27, 1960 (CHAP, US). Zapotitldn, near Tehuacdn, Ross, Oct. 1906 (US).
Between Apala and the top of Cerro Chichiltepec, Smith, Peterson, & Tejeda
3879 (US). More os: Sierra de Ocuila rumbo Mexicapa, Lyonnet 2873 (US).
Valle de Tepeite, Lyonnet & Elcoro 1838 (US). Veracruz: Maltrata, Kerber 226
(US). Mexico: Rineén, Temascaltepec, Hinton 2081 (MO, NY, US). Socabén,
Temascaltepec, Hinton 2324 (GH, LL, MICH, NY, US). Tejupilco, Temascal-
tepec, Hinton 2719 (GH). Sultepec, Matuda 26615 (US). Ixtaccihuatl, Purpus
1611 (ILL, MO, US). Salto de Agua, Purpus 1629 (US). 4 mi. S. of Tenan-
cingo, Reeder & Reeder 4168 (US, YU). Distrito Feperau: Lomas de Tacubaya,
Gdéndara, in 1938 (US). Villa Guerrero, Tateoka 1119 (US). Micuoacin:
Lomas de la Huerta, Arsene 2465 (ILL, MO), 2465!4 (F, US), 2825 (MO, NY, US),
5438 (NY, US), 5446 (ILL, MO, NY, UC, US), 5851 (MO, US), May 30, 1909
(US). Punguato, Arséne 2674 (US). Cerros de San Miguel, Arséne 5304 (NY,
US), 6803 (US). Loma Santa Maria, Arséne 8585 (MO, US). Along road from
Tsitzio-Tiquicheo-Huetamo-Altamira, Sohns 880 (US). GurrreEro: Teotepec,
Mina, Hinton 14803 (ARIZ, ILL, LL, NY, TEX, US). Oaxaca: Ca. 170 km.
N. of Oaxaco City, Hernéndez & Jenkins X-810 (US holotype). Ixtlan de Judrez,
Krueger & Gillespie 25 (CAS, LL, MO). Mina de Dolores, near Talea, Liebmann
673 (US fragment), 984, in 1841-43 (US). Mecoatl4n, S. Andrés, Liebmann
736 (US fragment). Dolores, Liebmann 737 (MO, US). Reyes, Nelson 1780 (US).

221-352—67_5

136 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

(?)Sierra de San Felipe, Pringle 5576 (GH, MEXU, US), 5577 (US); (2) Smith 927
(in part) (MO, NY, US). Temascalapa to San Ildefonso de Villa Alta proper,
Santos 3530 (NY, US). 45 mi. N. of Oaxaca, Soderstrom 417 (US). Cutapas:
Between San Cristébal Las Casas and San Gregorio, Hernandez & Sharp 534 (US).
Barranca Honda, Siltepec, Matuda 4124 (GH, LL, MICH, NY).

GUATEMALA: HvenvetTensanco: Aguacatin road, 10 km. E. of Hue-
huetenango, Standley 82073 (F, US). Between San Sebastiin H. and large
pefiasco above town, Steyermark 50504 (F, US). Barranco “Palo Negro’’ about
10 km. W. of Aguacatén, Williams, Molina, & Williams 21850 (US). San
Marcos: San Marcos, Lehmann 1578 (US fragment, ex G). Soros: Lago
Atitlin, de Koninck 146 (US).

HONDURAS: Morazin: Suyapa, Swallen 11277 (US).

Muhlenbergia versicolor is distinguished by the long ligule and usually
short sheath auricles, the narrow, densely flowered dark green or
purplish-green panicles, the spikelets over 3 mm. long, the lemmas
long-awned and villous over most of the surface on and between the
nerves. It appears to be most closely related to Af. emersleyi, and
most herbarium specimens have been determined as that species.
Although the two are similar in having long ligules and pubes-
cence on the long-awned lemmas, they are easily separated. In
M. versicolor almost the whole lemma is sparsely to densely villous
on and between the nerves; in M. emersleyi the pubescence is confined
to the midnerve and margins. The sheaths in M. versicolor are
characterized usually by short auricles but these are lacking in
M. emersleyi. The spikelet is dark green or purplish green in JM.
versicolor, but lighter and most often purplish or tan in AZ. emersleyt.

Specimens of Hpicampes (Smith 927; Pringle 5576, 5577) collected
on the summit ledges of Cerro de San Felipe (above the city of
Oaxaca, at an elevation of about 10,000 ft.) are without doubt allied
to M. versicolor. The base of the plant, the lead-green panicles,
long spikelets, and villous lemmas are characteristics of M. versicolor.
These specimens, however, exhibit some fundamental differences
from the typical plants: The glumes are blunt rather than acute or
dentate and awn-tipped as in AZ. versicolor; the auricles of the sheaths
are very long, 1-3.5 em., those of M. versicolor being less than 1 cm.
long; the panicles are narrower, shorter, and the branches more
appressed. In one specimen the lemmas are awnless, but long-awned
in others, as in typical M. versicolor. When more information
becomes available from additional field studies on Cerro de San
Felipe, this curious group of specimens may prove to represent a
distinct species.

10. Muhlenbergia gigantea (Fourn.) Iliteche. N. Amer. Fl. 17(6):461. 1935.
Epicampes expansa Fourn. Mex. Pl. 2:88. 1886. (Botteri 104 in part
[Botteri & Sumichrast 104], Orizaba [Veracruz], Mexico. Botteri 104 in

part is also cited by Fournier under Epicampes mutica Rupr. ex Fourn.)

SODERSTROM—PODOSEMUM AND EPICAMPES 137

Epicampes bourgaet Fourn. Mex. Pl. 2:88. 1886. (Bourgeau 2973,
Escamela pr. Orizaba [Veracruz], Mexico.)

Epicampes bourgaet var. mutica Fourn. Mex. Pl. 2:88. 1886. (Liebmann
676, 678 Mirador [Veracruz], Mexico, and Liebmann 677, Tlaltengo
[Veracruz], cited.)

Epicampes laxiuscula Fourn. Mex. Pl. 2:88. 1886. (Botteri 155, Orizaba
[Veracruz], Mexico.)

Epicampes gigantea Fourn. Mex. Pl. 2:88. 1886. Basis of Muhlenbergia
gigantea (Fourn.) Hitche. (Bourgeaw 3137, Rfo Blanco near Orizaba
[Veracruz], Mexico.)

Epicampes ehrenbergit Mez. Repert. Sp. Nov. Fedde 17:212. 1921. (Type
from Cuesta de Pinoleo [Hidalgo?], Mexico.) This species was questionably
referred to Muhlenbergia robusta (Fourn.) Hitche. in N. Amer. Fl. 17(6) :462.
Although the type has not been seen, the description is that of M. gigantea
for it states, in part: “ligulis magnis .. .” (the ligule of M. robusta is very
short) and “‘Inflorescentia myriantha, bene thyrsoidea, 3 pinnatim panni-
culata [sic] . . .” (which must refer to the typically large and widely
spreading panicle of M. gigantea).

Muhlenbergia alta Hitche. N. Amer. FI. 17(6):461. 1935. (Hitchcock
7180, Jalisco, Mexico.)

Muhlenbergia magna Hitche. N. Amer. Fl. 17(6):460. 19385. (Pringle
3335, Jalisco, Mexico.)

Strongly caespitose perennial; culms stout and thick, robust,
glabrous or somewhat scaberulous, 130-300 cm. (mostly 1.5-2.5 m.)
tall; nodes 3, glabrous below; basal sheaths strongly compressed-
keeled, glabrous or somewhat scaberulous; ligule membranous through-
out, generally undivided at the base, becoming lacerate above, 1-3.5
(usually 1-1.5) em. long; sheath auricles lacking, rarely rudimentary
ones present; blades of the culm 35-115 (usually more than 65) em.
long, flat, the edges saw-toothed, 2.5-12 (usually 5-8) mm. wide,
upper surface of the blade usually scaberulous, the lower surface
scaberulous to scabrous, scabrous-ciliate on the margins near the
ligule; upper throat scaberulous to scabrous; collar glabrous (some-
times scaberulous).

Panicle very large and open, the branches weak and lax and some-
what drooping or more rigidly flexuous, purplish or purplish brown,
45-110 (usually 50-100) cm. long, 20-30 cm. or wider, the branches
opening wide, long-naked below, the aggregates of spikelets borne on
the tertiary and quaternary branches, the groups of branches borne
distant from each other on the axis; primary branches 11-27 (usually
15-20) em. long, the secondary branches 1.5-12 (usually 4-10) cm.
long, the tertiary branches 0.5-5.5 (usually 1-4) cm. long, quaternary
branches, when present, to 2.5 em. long; pedicels shorter than the
spikelets, usually glabrous (sometimes scaberulous) ; spikelets 1.4—2.5
(mostly 1.5-2.2) mm. long; glumes rounded or acutish at the tip, about
equal, a little longer than the floret, glabrous or scaberulous, awnless,
1-nerved; lemmas rounded or acutish at the tip, 1.3-2.2 (mostly 1.6-2)

138 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

mm. long, glabrous, 3-nerved, awnless (the midnerve sometimes be-
coming darker and thickened toward the tip and extending beyond it
as a tiny point); palea equaling the lemma, glabrous.

Fragment of type in the U.S. National Herbarium, collected in re-
gion d’Orizaba, Rio Blanco, Veracruz, Mexico, Sept. 30, 1866, by
Bourgeau (no. 3137).

Distripution: Open rocky slopes and barrancas, at elevations of
2,000-7,500 ft., most commonly found at ca. 3,500-5,000 ft. Known
only from Mexico, ranging from Sinaloa and Durango southeastward
to Chiapas. Apparently infrequent.

MEXICO: Sinatoa: E. of Santa Luefa, Reeder & Reeder 2488 (YU). Dv-
ranao: W. of La Ciudad (Mazatlin-Durango road), Reeder & Reeder 2522 (YU).
Jauisco: SW. Autlin, Herndndez 4643 (CHAP). Zapotlin, Hitchcock 7180 (US,
holotype of M. alta), 7247 (ILL, US), Puente San Pedro, SW. of Tecalitlan
McVaugh 18096 (MICH). Guadalajara, Pringle 3335 (F, GH, NY, UC, US,
isotypes of M. magna), 11738 (CAS, F, GH, MO, US). Sierra Madre, W. of
Bolafios, Rose 3002 (US). Guadalajara, Soderstrom 607 (US, YU). VERACRUZ:
Orizaba, Botteri 103 (US fragment, ex CGE), 104 (US fragment of type of E.
expansa, ex CGE), 159 (US fragment, ex CGE). Escamela, region d’Orizaba,
Bourgeau 2973 (US fragment of type of E. bourgaei, ex P). Orizaba, Rfo Blanco,
Bourgeau 3137 (US fragment of type of E. gigantea, ex LE, P). Orizaba, Cerro
del Borrego, “Curso de ecologia vegetal,” Dec. 14, 1959 (CHAP). Mango de Dn.
Bartolo carretera-Veracruz-Cérdoba, Hernandez, Tapia, & Malthus SE-1482
(CHAP). Mirador, Liebmann 676 (US), 678 (US). Orizaba, ‘“C. Mohr herbar-
ium” (US). Orizaba, Muiller, in 1850 (NY). Veracruz a 22 km. de Catemaco
sobre la carretera Acayucan, Tapia & Martinez SE-644 (CHAP). Loc.? von
Rozynski, in 1932 (1933?) (F, NY, YU). Mexico: Cajones, Temascaltepec, Hin-
ton 2386 (MO, NY, US). Vigas, Temascaltepeec, Hinton 2626 (MO, NY, US).
Micuoackn: Pto. Cruces, Coaleoman, Hinton 12471 (ARIZ, GH, LL, MO, NY,
TEX, US). 22 km. S. of Uruapan, King & Soderstrom 4860 (MEXU, MICH,
SMU, TEX, US). Between Charapengo and El Guaco on road from Uruapan to
Apatzing’n, Moore, Hernandez & Porras 5750 (UC, US). GUERRERO: Machén,
Mina, Hinton 9635 (MO, NY, US). 138 mi. N. of Tierra Colorada, Reeder &
Reeder 4157 (US, YU). Oaxaca: Cerro de San Felipe, Conzattt 710 (GH),
Conzatti 2534 (US). Tepenixtlahuaca, distrito de Tuquila, Conzatti 4376 (GH,
US). Reyes, Nelson 1779 (GH, US). Temascalapa to San Ildefonso de Villa
Alta, district of Villa Alta, Santos 3531 (US). Curapas: between San Geronimo
and El Pozo via Abasolo and El Retiro, Hernandez & Sharp X-597 (US). Tuxtla
Gutiérrez, Tateoka 1103 (US).

Muhlenbergia gigantea is distinguished by its large size, robust
culms, long ligule, large, wide-spreading panicles, and relatively small
glabrous spikelets. It is most closely related to M. mutica and the
type specimens of both are from Orizaba, Veracruz, Mexico. The
spikelets differ in the ratio of glume length to floret length: In MM.
gigantea the glumes are about equal to the floret; in M7. mutica the
clumes exceed the floret by several millimeters. In the latter species

SODERSTROM—PODOSEMUM AND EPICAMPES 139

the glumes are very acute, but more rounded or acutish in the former.
This is possibly a variation within the same species, for the terminal
spikelets of M. gigantea rarely have elongated glumes extending
beyond the floret for several millimeters. An example is found in the
type material of M. gigantea (Bourgeau 3137). The spikelets average
2 mm. in length, but in one case the terminal spikelet is 2.5 mm.,
enclosing a 1.5 mm. floret. As relatively few collections of M. mutica
have been made, I do not feel safe in generalizing on the variability
of the species at this time, and for this reason am not combining them.
Additional field work, particularly in the region of Orizaba, Veracruz,
will be necessary before this problem can be solved.

Herbarium specimens of M. robusta are often confused with M.
gigantea. When only a panicle is present, the wide-spreading habit
is often not obvious in dried condition. If vegetative parts are
present, the short ligule and rudimentary sheath auricles of MZ. robusta
readily separate it from AZ. gigantea.

Hitchcock (1935) described two new species from Jalisco, Mexico:
Muhlenbergia magna (Pringle 3335) and M. alta (Hitchcock 7180).
These specimens, like M. gigantea, are large robust plants with long
lacerate ligules, wide-spreading panicles, and glabrous spikelets.
Muhlenbergia alta is described as differing from M. gigantea in its
smaller spikelets, 1.5 mm., as opposed to 2-2.5 mm. <A study of the
holotype of M. alta reveals that the spikelets actually range from
1.5-2 mm. in length but such variability also occurs in specimens
referred to M. gigantea! The holotype of M. magna exhibits no basic
differences from MM]. gigantea except for the more rigid and erect
panicle branches. Color of the panicle is the only character used in
the key to separate the two species (purplish in M. gigantea, brownish
in MZ. magna). Many of the specimens available for study could not
be placed satisfactorily on the basis of erectness versus laxness of
panicle branches or on the color of the inflorescence. A striking
example of color difference was found in specimens collected in
Durango, Mexico (J. & C. Reeder 2522, yellowish brown) and Sinaloa,
Mexico (J. & C. Reeder 2488, dark purple). Both plants are similar
except for color, The label of the former reads: “On a steep rocky
slope. Both yellow and purple forms here.”

Within limits, characters such as spikelet size and panicle color
and habit are variable in species of Hpicampes and are not of specific
magnitude. As M. magna and M. alta appear to represent nothing
more than normal variation within A/. gigantea, they are here com-
bined. The following comparison of characters of these three sup-
posed species illustrates their similarity:

140

height

culm surface
node surface
basal sheaths

upper throat

collar

blade width

upper surface of
blade

ligule texture

ligule length
sheath auricles
panicle length
panicle width
axis surface
pedicel length
pedicel surface

panicle color

panicle branches:

primary
secondary
tertiary
quaternary
glume Ist/2nd
glume surface

spikelet length
lemma length
lemma surface
awn length of
lemma

M. gigantea specimens

160-300 em.
glabrous
glabrous
glabrous,
compressed-
keeled
scaberulous to
scabrous
mostly glabrous
(sometimes
scaberulous)
4-12 (usually 7—9)
mm,
glabrous to
scaberulous
(mostly
scaberulous)
membranous,
becoming
lacerate
5-35 mm. (usually
1 cm. or more)
lacking
45-110 em.
20-35 cm. or
more, widely
spreading
scabcrulous

0.2-0.8 mm., less
than spikelet

glabrous to
scaberulous

purplish (some-
times brownish)

12-27 cm.

4-12 cm.

1-5.5 cm.

to 2.5 cm.

about equal

glabrous to
scaberulous

1.4-2.5 mm.

1.3-2.2 mm.

glabrous

awnless or extend-
ing from mid-
nerve as a tiny
point

M. alta holotype

193 em.
glabrous
glabrous
glabrous
eompressed-
keeled
scaberulous

glabrous

6-9 mm.

seaberulous

membranous,
becoming
lacerate

6 mm.

lacking
100 cm.
over 30 cm.

scaberulous
less than spikelet
glabrous

dark purple

25 em.
11 cm.
2 cm.

equal
seaberulous

1.5-2 mm.
1.8 mm.
glabrous
awnless

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

M. magna holotype
172 cm.

glabrous

glabrous

glabrous
compressed-keeled

scaberulous

glabrous

3.5 (in other speci-
mens 2,.5-7) mm.

glabrous to
scaberulous

membranous,
lacerate

ca. 10 mm.

lacking
46 cm.
25 em.

glabrous to
seaberulous

0.4-0.7 mm., less
than spikelet

scaberulous

brownish

14 cm.
5.5 em.
1.5 em.
equal
glabrous

1.7-2 mm.

1.6 mm.

glabrous

awnless or extend-
ing from mid-
nerve as a tiny
point

SODERSTROM—PODOSEMUM AND EPICAMPES 141

11. Muhlenbergia mutica (Rupr. ex Fourn.) Hiteche. N. Amer. Fl. 17(6):459.
1935.

Epicampes mutica Ruprecht. Bull. Acad. Roy. Sci. Bruxelles, Belg. IX(2):
230. 1842. Nomen nudum. (Galeotti 5797, savanes de Mirador, Zacuapan,
et Cantaranas prés du Puente Nacional, Veracruz, Mexico.)

Epicampes mutica Rupr. ex Fourn. Mex. Pl. 2:87. 1886.

Strongly caespitose, robust perennial; culms stout, thick, glabrous,
2 meters or more tall; nodes 3, glabrous below; basal sheaths strongly
compressed-keeled, glabrous; ligule membranous, undivided at the
base, lacerate above, about 1 cm. long; sheath auricles lacking; blades
of the culm 35 cm. long (in specimen examined), 4-5 mm. wide, the
upper and lower surfaces of the blades scaberulous to scabrous; upper
throat scaberulous; collar glabrous.

Panicle very large and open, the branches weak and lax, brownish or
purplish brown, about 1 m. long and 30 cm. or wider; the primary
branches as much as 30 cm. long, the secondary branches 6-12 cm.
long, the tertiary branches 1.5—2 cm. long, quaternary branches lack-
ing; axis scaberulous; pedicels shorter than the spikelet, those of the
terminal spikelets longer than those of the lateral ones, scaberulous,
becoming more scabrous just beneath the spikelet; spikelets 2.1-3.1
(mostly 2.5-2.8) mm. long; glumes about equal, much longer than the
floret, scaberulous, acute, awnless, nerveless or indistinctly 1-nerved;
lemmas 1.9-2.2 mm. long, glabrous, roundish at the tip, obscurely 3-
nerved, awnless; palea a little shorter than the lemma, glabrous.

Fragment of type (ex P) in the U.S. National Herbarium, collected
in “savanes de Mirador, Zacuapan, et Cantaranas prés du Puente
Nacional, Veracruz, México,” by Galeotti (no. 5797). This specimen
was cited by Ruprecht in 1842 without description. In his descrip-
tion of the species, Fournier (Mex. Pl. 2:87. 1886) cited this speci-
men, along with several other syntypes. Galeotti 5797 (P) is here
designated as the lectotype.

Distrisution: Open rocky hillsides, at elevations of about 4,000-
6,300 ft. Known only from Mexico in the states of Veracruz and
Chiapas. Apparently rare.

MEXICO: Veracruz: Orizaba, Botteri 111 (US fragment, ex P), 729 (US frag-
ment); Bolteri & Sumichrast 104 (US fragment, ex P); Hitchcock 6348 (US).
Mirador, Zacuapan, and Cantaranas, Galeotti 5797 (US fragment of lectotype, ex
P). Cutapas: El Pozo to Oxchuc, Hernandez & Sharp X-616 (US).

Muhlenbergia mutica is distinguished by its nonauriculate sheaths,
large, wide-spreading panicle, and acute glumes much longer than the
floret. It is most closely related to and doubtfully distinct from M.
gigantea. The only character to separate the two is the length of the
glumes in relation to the floret: In M. gigantea the glumes equal the
floret, but in M. mutica the floret is exceeded by the glumes. Until

142 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

more material is available to study this character critically, I prefer not
to combine the two species.

12. Muhlenbergia inaequalis Soderstrom, sp. nov.

Perennis caespitosa, culmis angustis erectis glabris vel scaberulis
50-70 cm. altis, nodis 3-4 glabris, vaginis inferioribus compressis et
carinatis; ligula membranacea-firma, 0.5-1.5 mm. longa; laminis culmi
angustis attenuatisque conduplicatis vel planis utrinque scaberulis, ca.
15-26 cm. longis, 2-3 mm. latis; collis glabris; panicula laxa, 25-30 cm.
longa, usque ad 10 cm. lata, axe scaberulo, ramis primariis usque ad
8 cm. longis, secundarilis usque ad 3 cm. longis, pedicellis scaberulis;
spiculis 2.5-3.1 (plerumque 3) mm. longis, glumis acutis, inaequalibus,
quam flosculo brevioribus, glabris, nervis 0 vel 1, indistinctis, lemmati-
bus acutis, 2.5-3(3) mm. longis, inferne brevi-pilosis, infra apicem
aristatis, aristis 1-2 em. longis.

A M. emersleyi Vasey ligulis brevioribus, glumis inaequalibus quam
flosculo brevioribus, et lemmatibus inferne brevi-pilosis differt.

Caespitose perennial; culms erect, narrow, glabrous to scaberulous
about %-% m. tall; nodes 3-4, glabrous; basal sheaths compressed-
keeled, glabrous to scaberulous; ligule a short, brown, firm mem-
branous rim 0.5-1.5 mm. long; sheath auricles lacking; blades of the
culm 15-26 em. long, 2-3 mm. wide, flat or folded, the upper and
lower surfaces scaberulous; upper throat scaberulous; collar glabrous.

Panicle pyramidal, lax above, the branches loosely ascending or
spreading, purplish to purplish tan, 25-30 cm. long, 5-10 em. wide;
axis scaberulous; branches naked below, the lower primary branches
of the panicle 5-8 cm. long, the secondary branches 2-3 em. long;
pedicels scaberulous, those of the lateral spikelets 0.5-1.8 (1-1.2)
mm. long, those of the terminal spikelets longer; spikelets 2.5-3.1
(mostly 3) mm. long; glumes acute and some awn-tipped, unequal and
shorter than the floret, the first about three-fourths as long as, and
narrower than, the second, 1.6-2.5 (1.9) mm. long, the second 2-3
(2.4-2.5) mm. long, glabrous or scaberulous, nerveless or indistinctly
1-nerved; lemmas acute, 2.5-3(3) mm. long, short-pilose over the
lower part and especially on the midnerve one-half to three-fourths
its length, the lemma 38-nerved, the middle nerve excurrent from
below the tip as a flexuous awn 1-2 cm. long; palea about equaling
the lemma, short-pilose between the keels for one-half to three-fourths
its length; anthers yellow, 1.1-1.2 mm. long.

Holotype in the U.S. National Herbarium, no. 2380325, collected
in Quebrada de Duri, Estado de Trujillo, Venezuela, November 11,
1939, by A. Miller (no. 973). Tsotypes at VEN.

Distrisution: Mountains to the south and southwest of Lago de
Maracaibo, in Venezuela and Colombia.

SODERSTROM—PODOSEMUM AND EPICAMPES 143

VENEZUELA: Trusii.0o: Quebrada de Durf, Miller 973 (US holotype, VEN
isotypes). Mé&ripa: Funck & Schlim 1017 (US fragment ex Herb. Boissier).

COLOMBIA: (?) Mutist 1760-1808 (US fragment ex MA). Norre pE San-
TANDER: Ocafia Schlim 190 (US fragments ex Herb. Boissier and K).

Muhlenbergia inaequalis has the habit of MZ. emersley: and possesses
features of other species closely related to that species. The pyramidal
panicle of long-awned spikelets looks much like typical M. emersleyi
but the spikelets differ in important respects. The glumes are unequal
and shorter than the floret, as in MZ. scoparia. The lemma is short-
pilose over the lower part, on and between the nerves, somewhat as
in M. versicolor. The ligule is a short, membranous rim as in M.
breviligula. The short ligule, unequal glumes shorter than the lemma,
and pubescence on and between the nerves of the lemma readily
distinguish this species.

The collection of José Celestino Mutis, 1760-1808, is represented
in the U.S. National Herbarium by only a panicle. Tentatively it
is assigned to this species, as the glumes are unequal and shorter
than the floret. However, the glumes are broader and more scabrous
and less acute than those of the type and the panicle is more compact.
The lemmas are short-pilose like those of the type, but shorter.
The spikelets of the Schlim, and Funck and Schlim collections match
closely those of the type specimen.

The name “inaequalis” refers to the very unequal glumes, an
unusual feature in E’picampes,

13. Muhlenbergia scoparia Vasey. Contr. U.S. Nat. Herb. 1:283. 1893.
Muhlenbergia carinata Mez. Repert. Sp. Nov. Fedde 17:213. 1921. (Based
on Pringle 2350, which is also the lectotype of Muhlenbergia scoparia Vasey
[1893].)

Caespitose perennial; culms 55-88 (usually about 75) cm. tall,
glabrous to scaberulous; nodes glabrous, covered by the sheaths;
basal sheaths compressed-keeled, glabrous, becoming scaberulous
toward the collar, the old sheaths and blades at the base curled,
generally reddish-brown, fibrillose; ligule membranous throughout,
usually hidden between the folded blade above, 8-16 (usually 10 or
more) mm. long; sheath auricles lacking; blades of the culm 25-50
(mostly about 50) cm. long, flat or folded, 1-4.5 mm. wide when
flat (mostly 2.5-3 mm. wide), the edges of the blade slightly saw-
toothed, upper and lower surfaces of the blade scaberulous to scabrous;
upper throat scabrous; collar glabrous to scaberulous (mostly scab-
erulous).

Panicle loose, nodding, the branches narrow, borne in groups
at intervals along the axis, light green to purplish or brownish green,
22-30 cm. long, 1.5-5 cm. wide; axis scaberulous to very scabrous,
strongly ribbed; primary branches 3.5-14 (mostly 4-9) cm. long,

144 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

naked below, the secondary branches 1-5.5 cm. long, tertiary branches
lacking or, when present, 1-2 cm. long; pedicels of the lateral spikelets
0.6-1.2 mm. long, those of the terminal spikelets to 3 mm. long,
scaberulous scabrous, especially below the spikelet; spikelets 1.5-2.2
(averaging 2) mm. long, excluding the awn tips; glumes very acute,
first glume shorter than the second, awn-tipped (especially the
second), the awn sometimes as much as 1 mm. long, glumes glabrous
to scabrous, hyaline, nerveless or indistinctly 1-nerved; lemmas
1-2 (usually 1.5-2) mm. long, exceeding the glumes by a few mm.,
glabrous or scaberulous dorsally, the lower margins of the lemma
conspicuously short-pilose, bearing short straight hairs to one-third
the length of the lemma; lemmas acutish at the tip, 3-nerved, awned
from just below the tip, the awn 6-15 mm. long; palea a little shorter
than the lemma, glabrous.

Lectotype in the U.S. National Herbarium, no. 999089, collected on
rocky hills near Guadalajara, Jalisco, Mexico, Nov. 10, 1889, by
C. G. Pringle (no. 2350).

Vasey cited two specimens in the original description and remarked,
“collected at Batopilas by Dr. Edward Palmer in 1885, and at
Guadalajara (No. 2350) by C. G. Pringle in 1889.” Of these syn-
types, Hitchcock (1935b, p. 457) cited the latter as the type, thus
effectively choosing it as lectotype. A note by Dr. Agnes Chase on the
sheet of Pringle 2350 reads “‘second of the 2 specimens cited, but first
is incomplete, description of habit, height, leaves, etc. must have
been drawn from this.”

DistrisuTion: Exposed rocky mountainsides and open areas in
the oak zone, at elevations of 2,500-6,000 ft. Mexico, from Chi-
huahua, Sonora, and Sinaloa, south to Michoacén. Apparently
infrequent.

MEXICO: Palmer, in 1886 (DS, ILL, NY). Sonora: Canyon Sapopa, Rfo
Mayo, Gentry 1289 (GH, MO). Sierra Tecurahui, Barclay & Arguelles 19387
(US). Crimvanva: Batopilas, Palmer, in 1885 (ARIZ, US, isosyntypes).
StnaLoa: Mesa Malqueson, Cerro Colorado, Gentry 5178 (ARIZ, F, GH, MO,
NY, UC). Loc.? Ortega 1120 (F, US). 10 mi. E. of Santa Luefa (road from Villa
Union to Durango), Reeder & Reeder 2451 (YU). Nayarit: La Barranca, Jones
23446 (CAS, GH, MO, UC), Feb. 21, 1927 (NY, POM). Mina Esperanza, Rosa
Morada, Ortega 6672 (CAS, US), Acaponeta, Rose, Aug. 2, 1897 (US). Jatisco:
Km. 211 Guadalajara to Barra de Navidad Aquilera in 1958 (CHAP). 38 mi.
N. of Pihaumo, McVaugh & Koelz 1394 (MICH, US). San Sebastidn, trail to
Monte Oscuro, Sierra Madre Mountains, Mexia 1644 (CAS, DS, F, GH, MICH,
MO, NY, UC, US). Guadalajara, Pringle 2350 (F, LL, MEXU, US, isolecto-
types). Micnoackn: Coalcomin, Hinton 12699 (ARIZ, GH, LL, MO, NY,
TEX, US).

Muhlenbergia scoparia is distinguished by its unequal, awn-tipped
elumes that are shorter than the floret. It is probably most closely
related to AM. grandis, sharing the features of reddish-brown basal

SODERSTROM—PODOSEMUM AND EPICAMPES 145

sheaths, short spikelets, and long-awned, comparatively glabrous
lemmas with a short pilose callus. In M. grandis, however, the
glumes are about equal, not awn-tipped, and the ligule is shorter,
usually no more than 6 mm. Both species have a range farther west
in Mexico than the majority of species of Epicampes, occurring in the
states of Sinaloa, Nayarit, and Jalisco. Muhlenbergia scoparia also
has a more extensive north-south range, from Chihuahua and Sonora
to Michoacdn.

14. Muhlenbergia iridifolia Soderstrom, sp. nov.

Perennis caespitosa, culmis erectis glabris 14% m. altis, nodis 4
glabris, vaginis inferioribus forte compressis et carinatis; ligula mem-
branacea-firma, 5-7 mm. longa; laminis culmi latis acutisque condu-
plicatis vel planis utrinque glabris, 30-70 cm. longis, 4-7 mm. latis;
callis glabris; panicula supra laxa, 45 cm. longa, 3 cm. lata, axe glabro,
ramis primariis 7-8 cm. longis, secundariis ca. 3 cm. longis, pedicellis
elabris; spiculis 4-4.2 mm. longis, glumis angustis minute aristatis,
quam flosculo paulo longioribus, glabris, nervis 0 vel 1, indistinctis,
lemmatibus angustis, 4 mm. longis, 3-nervis, glabris, inferne brevi-
pilosis, infra apicem aristatis, aristis 2-2.5 cm. longis.

Caespitose perennial; culms erect, glabrous, about 14 m. tall; nodes
4, glabrous; basal sheaths strongly compressed-keeled, glabrous,
spreading outward in a distinct flabellate fashion; ligule brown, firm-
membranous, lacerate but rather blunt or truncate, conspicuous from
side view, 5-7 mm. long; sheath auricles lacking; blades of the culm
30-70 cm. long, 4-7 mm. wide, the older blades folded but younger
blades mostly flat, the upper and lower surfaces glabrous except for the
scabrous, saw-toothed margins and midnerve on the undersurface;
upper throat glabrous; collar glabrous.

Panicle lax above, the branches loosely ascending or spreading,
purple, 45 cm. long, 3 cm. wide; axis completely glabrous; branches
naked below, the lower primary branches of the panicle 7-8 cm. long,
the secondary branches about 3 cm. long; pedicels glabrous, the swollen
portion just beneath the spikelet puberulent, mostly 1-2 mm. long,
those of the terminal spikelets longer and more variable in length;
spikelets 4—4.2 mm. long; glumes purple, narrow, acute, minutely awn-
tipped, the second a little longer than the first, just exceeding the
floret, glabrous, nerveless or indistinctly 1-nerved; lemmas narrow,
acute, about 4 mm. long, glabrous, short-pilose at juncture of lemma
and callus at the base on palea side, the hairs about one-fourth the
length of lemma, 3-nerved, the middle nerve excurrent from just below
the tip as a slightly flexuous purple awn 2-2.5 cm. long; palea a little
shorter than lemma, completely glabrous.

146 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Holotype in the University of Michigan herbarium, collected in
Jalisco, Mexico, in “Steep mountains 11-12 miles south of Talpa de
Allende, in the headwaters of a west branch of Rio de Talpa; barranca
above a rapid clear stream, in dense forest of Quercus, Carpinus,
Distylium, Magnolia, Podocarpus, with pine forest on the ridges above;
abundant in the pine forest; in clumps, inflorescence purple; alt.
1200-1700 m., 23-25 November 1960,” by Rogers McVaugh (no.
21313). Isotype at US. Known only from the type collection.

Although I would prefer to have additional collections at hand in
describing this new species, there is little doubt that it is distinct.
The habit of the inflorescence and spikelets with awn-tipped glumes
and lemmas glabrous except for the short-pilose base, suggest a
relationship with M/. scoparia. The distinct fan-shaped appearance
of sheaths and the stiff, rather wide, flat blades ascending erectly
much like an iris suggested the name “‘iridifolia.”” The firm brown
ligules are conspicuous and appear to be chopped off, leaving a firm
brown base about 5-7 mm. long. The combination of long spike-
lets (4 mm.), awn-tipped glumes, and lemmas glabrous except at
the base where the callus is short-pilose, readily distinguishes this spe-
cies from all others in Epicampes.

15. Muhlenbergia speciosa Vasey. Bull. Torrey Club 18 (12):231. 1886.
Bealia speciosa (Vasey) Beal. Grasses N. Amer, 2:268. 1896. Based on
Muhlenbergia speciosa Vasey.
Epicampes speciosa (Vasey) Jones. Contr. West. Bot. 14:7.1912. Based on
Muhlenbergia speciosa Vasey.

Caespitose perennial; culms pubescent or villous, the pubescence
present as broad bands on the culm, glabrous between the bands,
95-129 em. tall; nodes 5, densely pubescent to villous below and
usually above; basal sheaths compressed-keeled, glabrous below and
becoming villous toward the collar, or villous throughout, especially
along the edges; ligule a membranous rim, 0.3-1.1 mm. long; sheath
auricles lacking; blades of culm 36-52 cm. long, 2.5-4 mm. wide when
flat, the blades usually folded with the edges rolled backward on the
lower (abaxial) surface, upper surface of the blade villous (or scabrous)
at the base near the ligule, the pubescence becoming sparse toward the
distal end, here the surface scaberulous, or scaberulous with scattered
long hairs, the lower surface of blade generally more villous than
the upper surface and for the greater part of its length, especially
villous near the ligule, becoming less so distally, here the surface
scabrous; upper throat densely villous.

Panicle large and plumose, the branches ascending or opening out-
ward, whitish or purplish white, 41-62 cm. long, 5-11 em. wide; axis
elabrous to scabrous with scattered long hairs; lower primary branches
9-13 em. long, naked below; secondary branches 3.5-5 ecm. long,

SODERSTROM—PODOSEMUM AND EPICAMPES 147

naked below; tertiary branches 1.3-3 cm. long; quaternary branches,
when present, to 0.8 cm. long; pedicels of the lateral spikelets 0.5-2
mm. long, of the terminal spikelets relatively long, 1.5-5.5 mm.,
glabrous to scaberulous, often with scattered long hairs on the surface;
spikelets 1.2-2.2 (averaging less than 2) mm. long; glumes equal or
the second a little longer than the first, 1.4-2.3 mm. long, usually
a little longer than the floret, mostly villous, rounded or lobed at the
tip or rarely with an awn tip, hyaline, nerveless, white; lemmas
1.2-2.2 mm. long, villous throughout, especially on the lateral margins,
3-nerved, the nerves purple and become thickened toward the tip,
awn of the lemma to 17 mm. long; callus of the lemma short and
densely pilose; palea obtuse, a little shorter than the lemma, villous,
especially between the keels.

Type in the U.S. National Herbarium, no. 999062, collected in
“southwestern Chihuahua, Mexico,’ August to November 1885, by
Dr. Edward Palmer (no. 30). Isotypes at GH and MEXU.

DistrisuTion: Found only in Mexico, where it is confined to rocky
slopes in the pine-oak zone and in arroyos, at elevations of about
1,000-6,600 ft. From southwestern Chihuahua through western
Durango, Sinaloa, Nayarit, and into Jalisco. Apparently infrequent.

MEXICO: Palmer, in 1886 (TAES). Cutnuanva: Palmer 30, Aug. to Nov.
1885 (US holotype, GH, MEXU, isotypes). Sinatoa: Culiacdn, Cofradfa,
Brandegee, Oct. 26, 1904 (POM, UC, US). Palmito and vicinity along Durango-
Sinaloa highway, Gentry & Arguelles 18201 (LL, US). Santa Lucia, Mexia 490
(CAS, MO, UC, US). Duraneo: La Bajada, Tamazula, Ortega 4335 (US).
21 mi. E. of Santa Lucia, Reeder & Reeder 2476, aff. M. speciosa (YU). Nayarit:
Tepic, Jones 23438 (CAS, GH), Feb. 10, 1927 (NY). 9 mi. N. of Compostela,
McVaugh & Koelz 575 (MICH). Ca. 5.5 mi. SW. of Jalisco, road to El Malinal,
McVaugh & Koelz 668 (MICH). ‘Along banks of arroyos,’’ Palmer 1919, in
1892 (GH, MO, NY, US). 7 mi. NW. of Tepic, Reeder & Reeder 4210 (US, YU),
4211 (US, YU). Jaxisco: SW. Autlin, Herndndez 4638 (CHAP, US). 4 mi.
N.-NE. of Talpa de Allende, McVaugh 20102 (MICH, US). Ca. 8 mi. SW. of
La Resolana, McVaugh 21105 (MICH, US). 3-5 road mi. above (N. of) La
Cuesta, road to Talpa de Allende, McVaugh 21226 (MICH, US).

Muhlenbergia speciosa is distinguished by the more or less villous
sheaths, blades, and culms, the very short ligule (about 1 mm. or less),
a large plumose panicle, and small spikelets (mostly less than 2 mm.
long), the lemmas long-awned. It appears to be most closely related
to M. pubescens. Both species are generally villous throughout, the
ligule is relatively short, and the glumes, lemmas, and paleas are more
or less villous. Muhlenbergia pubescens differs in the longer ligule
(2-3 mm.), a denser panicle with larger spikelets (mostly 2.4-2.6 mm.),
and the shorter awns of the lemmas. The ranges of both overlap
in southwestern Chihuahua and western Durango, Mexico, and some
hybridization may possibly occur. One specimen, Reeder & Reeder
2476, from western Durango, seems to be somewhat intermediate

148 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

between these two species and perhaps represents such a hybrid. The
long spreading branches of the panicle in this specimen and the very
short ligule are characters of AZ. speciosa. The 2.5 mm. long spikelets
and the short-awned lemmas are features of MZ. pubescens. The
habit and overall appearance are those of M. speciosa, to which it
appears to be more closely allied. ‘The extent of such putative hybrids
cannot be determined until many more collections are made in this
region. Comparison may also be made with MZ. grandis which is
similar in habit and occupies a similar range. Unlike M. speciosa
it has glabrous culms, foliage, and glumes, and longer ligule (1.5-6 mm.)
16. Muhlenbergia pubescens (II1.B.K.) Hitche. N. Amer. Fl. 17(6):460. 1935,
as to type, not as to description (see M. distans Swallen).

Agrostis pubescens H.B.K. Nov. Gen. & Sp. J:186. 1815. Basis of
Muhlenbergia pubescens (H1.B.K.) Hitche. (Type collected in “‘scopulosis
juxta Villalpando, Santa Rosa, et Cerro del Cubilete,’? 7800-8760 ft.
[mining areas around the city of Guanajuato, Guanajuato, Mexico].)

Agrostis lanata H.B.K. Nov. Gen. & Sp. 1:186. 1815. (Type from same
localities as preceding.)

Cinna? pubescens (H.B.K.) Kunth. Rev. Gram, 1:67. 1829. Based on
Agrostis pubescens H.B.K.

Cinna? lanata (H.B.K.) Kunth. Rev. Gram. 1:67. 1829. Based on
Agrostis lanata H.B.K.

Epicampes pubescens (H1.B.K.) Presl ex Kunth. Enum. Pl. 1:209. 1833.
Based on Agrostis pubescens H.B.K.

Epicampes lanata (H.B.K.) Presl ex Kunth. Enum. Pl. 1:209. 1833,
Based on Agrostis lanata H.B.K.

Epicampes pubescens (1.B.K.) Jones. Contr. West. Bot. 14:7. 1912.
Based on Agrostis pubescens H.B.K.

Muhlenbergia lanata (H.B.K.) Hitehe. N. Amer. Fl. 17(6):459. 1935.
Based on Agrostis lanata II.B.K.

Caespitose perennial; culms erect, 57-154 (usually about 100-125)
cm. tall, sparsely to densely villous, especially on the lower part;
nodes 3-4, sparsely to densely villous, especially below; basal sheaths
compressed-keeled, villous (rarely scabrous with long sparse hairs) ;
collar villous or scabrous with sparse long hairs; ligule membranous,
usually finely lacerated, 1.5—6 (usually 2-4) mm. long; sheath auricles
lacking; blades 12-61 (usually 25-50) em. long, usually short on the
upper part of the culm, 2.5-6.5 (usually 3-4) mm. wide, flat or folded,
upper surface villous at base, becoming less villous toward the tip
(rarely only scabrous with long scattered hairs), upper surface villous
throughout or sometimes scabrous with long scattered hairs; blades
of the culm constricted and villous at point of attachment to sheath;
upper throat villous.

Panicle erect, columnar, densely flowered, gray or purplish gray,
12-49 (usually 20-40) cm. long, 2-7 (usually 3-7) em. wide, the
branches of the panicle ascending, floriferous to base; axis villous,
rarely sparsely so; primary branches 3-11.5 (usually 5-8) em. long,

SODERSTROM—PODOSEMUM AND EPICAMPES 149

secondary branches 1-5 (usually 1-2) cm. long, tertiary branches
rarely to 1.5 em. long; pedicels shorter than the spikelets, villous
(sometimes sparsely so), 0.3-1 mm. long; spikelets 2.5-3 mm. long;
elumes broadly acute at the tip, longer than the floret, almost equal
or the first 1-2 mm. shorter than the second, sparsely to densely
villous, nerveless, gray, often purplish at the base; lemma 2.2-3
(usually 2.4-2.6) mm. long, sparsely to densely villous throughout
except at tip, 3-nerved, the median nerve often extending as an awn
from just below the tip, 1-10 (usually 6-7) mm. long, straight or
somewhat flexuous; palea villous throughout, rarely sparsely so.

Fragment of panicle of possible type marked “Agrostis pubescens”
(in Kunth’s script) in the U.S. National Herbarium, ex Herb. Hum-
boldt & Bonpland (P), collected near Guanajuato, Guanajuato,
Mexico, “in scopulosis juxta Villalpando, Santa Rosa, et Cerro del
Cubilete,” 7800-8760 ft. A photograph of the specimen is with the
fragment.

DisTRisuTION: Common on rocky mountain slopes and ledges and
in rocky ravines, canyons, and barrancas, at elevations of about 5,600-
8,000 ft. Known only from Mexico, from Chihuahua and Durango
south to Puebla. A collection from Oaxaca is doubtfuly referred to
this species.

MEXICO: Curmvuauva: Hills near Chihuahua, Pringle 391 (F, GH, MO,
US), 983 (MEXU, NY). 37 mi. S. of Villa Matamoros, Reeder, Reeder, & Soder-
strom 3451 (US, YU). Duranao: Canyon Cantero, Sierra de Gamén, Gentry 8406
(GH, MEXU, MICH, UC, US). 40 km. 8. of Durango, Hernéndez, Ruppert, &
Guevara X-2561 (US). 20 km. 8. of Durango, Herndndez, Ruppert, & Guevara
X-2576 (US). Km. 1395, Carretera Judrez, Parral-La Zarca, Hernandez &
Tapia N-589 (CHAP). Durango, Hitchcock 7587 (ILL, US), 7652 (NY, US).
SW. of Durango on Durango-Mazatlin road, Soderstrom 763 (US, YU), 768
(US, YU), 769 (US, YU), 773 (US, YU), 774 (US, YU). N. of Durango on
Durango-Judrez road, Soderstrom 788 (US, YU). 23 mi. W. of Durango, Reeder &
Reeder 2548 (YU), 2592 (YU). 31 mi. W. of Cd. Durango, Reeder & Reeder
2578 (YU), 2590 (YU). 6 mi. W. of Navios, Reeder, Reeder, & Soderstrom 3377
(US, YU), 3411 (US, YU), 3412 (US, YU), 3413 (US, YU), 3414 (US, YU).
Sierra Madre Occidental, W. of Cd. Durango, Reeder, Reeder, & Soderstrom
3428 (US, YU). 3 mi. E. of Navios, Reeder, Reeder, & Soderstrom 3429 (US, YU),
3430 (US, YU). 5 mi. E. of Navios, Reeder, Reeder, & Soderstrom 3433 (US,
YU), 3434 (US, YU), 3435 (US, YU). 11 mi. W. of Durango, Reeder, Reeder, &
Soderstrom 3444 (US, YU). Zacatecas: 2 mi, W. of Sombrerete, Soderstrom
733 (US). San Luts Porosf: 22° N. Lat., Parry & Palmer 933 (F, GH, MO, US).
20 mi. SW. of San Luis Potosf, Reeder, Reeder, & Soderstrom 3304 (US, YU).
Schaffner 190 (US). “Ex convalli 5. L. P.,” Schaffner, in 1876 (GH). San
Francisco in the Sierra de San Miguelito, Sohns 1049 (US). Canyon del Lobo,
Sierra de San Miguelito, Sohns 1158 (US). Canyon del Muerto, Sierra de San
Miguelito, Sohns 1319 (US), 1327 (US). AGUASCALIENTES: 7 mi. E. of Ojo
Caliente, Reeder & Reeder 4193 (US, YU). Mountains above Presa Calles,
Shreve 9258 (ARIZ, US). Jaxisco: San Juan de los Lagos, Herndndez X—4608
(CHAP, US). Ojuelos-Aquascalientes, Hernandez, Ruppert, & Guevara X-2513

150 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

(US), X-2515 (US). Road to Leén, Gto., 5 mi. SE. of junction at Lagos de
Moreno, McVaugh 17800 (MICH). 7 mi. SE. of Lagos de Moreno, Reeder &
Reeder 2306 (YU). 15 mi. SE. from Encarnacién de Dfaz on highway 45 to
Lagos de Moreno, Soderstrom 692 (US), 693 (US), 694 (US), 696 (US), 698 (US).
GuaNnasuato: About 9 mi. from Guanajuato, Moore 1355 (GH, MICH, UC, US).
695 (US), 12 mi. E. of Guanajuato, Reeder & Reeder 3088 (YU). 13 mi. NE. of
Guanajuato, Reeder & Reeder 3093 (YU). 2-4km. EF. of Guanajuato, Sohns 298
(US). About 6 km. E. of Guanajuato, Sohns 302 (US), 318 (US). About 8 km.
W.ofSan Felipe, Sohns 433. Puppia: 5 mi. NE. of Zacatepec, Soderstrom 483 (US,
YU). Tuaxcaua: Santa Ana Chiautempan, Arséne 1715 (GH, MO, NY, US).
Mexico: About 45 mi. 8. of San Juan del Rfo, Reeder & Reeder 2235 (YU). EI
Pedregal, just S. of Mexico, City Reeder & Reeder 3047 (YU). Oaxaca: (?)27 mi.
NW. of Cd. Oaxaca, Reeder & Reeder 2184, aff. M. pubescens (YU).

Muhlenbergia pubescens appears to be most closely related to M.
speciosa and like that species has moderately to densely villous vege-
tative parts, a relatively short ligule, and blades usually constricted
at the base. It is distinguished from M. speciosa by the denser
panicles with the branches more appressed, longer spikelets (usually
2.4-2.6 mm. in contrast to 2 mm.), and shorter awns on the lemmas
(7 mm. or less as opposed to 10 mm. or more).

In 1815 Kunth described two species of Agrostis, A. pubescens and
A. lanata, and indicated that the latter might be a variety of the
former. Agrostis pubescens was described as 3-4 ft. tall, generally
glabrous throughout, pubescent under the nodes, having a short,
lacerate ligule, white, nerveless, punctate-scabrous glumes, and
lemmas pubescent at the base. Agrostis lanata was described as
3 ft. or taller, generally lanate throughout, having pubescent nodes,
a short, lacerate ligule, and pilose glumes, lemmas, and paleas.

Presl (1830), along with his description of the original species of
Epicampes, mentioned that these two species of Agrostis probably
belonged with his new genus and added that A. lanata might be a
variety of A. pubescens, but he made no transfers. ICunth (1833),
following the opinion of Presl, formally transferred them to Epicampes.
Jones (1912), commenting on EL. pubescens, stated: “This is Agrostis
pubescens H.B.K. . .. A. lanata is a synonym.” He is the only
author who stated, without reservation, that they are clearly synony-
mous and are not worthy of recognition as varieties within one species.
Hitchcock (1935) transferred L. pubescens and E. lanatus to Muhlen-
bergia and, following the lead of Kunth and Presl, indicated that
M. lanata migk*, be a variety of M. pubescens.

Muhlenbergia lanata, as based upon Agrostis lanata, refers to a
widespread plant characterized by villous lemmas, a very short
ligule (about 3 mm. long), and moderately to densely villous glumes,
culms, and sheaths. Muhlenbergia pubescens, as circumscribed by
Hitchcock, is characterized by lemmas pubescent over the lower half,

SODERSTROM—PODOSEMUM AND EPICAMPES 151

a ligule 5-10 mm. long (or a little longer), and relatively glabrous
glumes, culms, and sheaths.

The type collections of ‘A. pubescens’’ and ‘A. lanata” are from
the same locality, near the city of Guanajuato. The majority of
specimens from this area of Mexico are quite villous throughout,
although occasionally the pubescence is confined mostly to the lemma,
appearing more sparsely on the glumes and remainder of the plant.
According to the original description these plants with sparse pubes-
cence would be referred /to “A. pubescens.” The very short ligule,
short blades, and at least some pubescence on the other vegetative
parts seem to indicate a less pubescent form than “‘A. lanata,” but
certainly not a distinct species.

A fragment of a specimen marked Agrostis pubescens, in Kunth’s
handwriting (Mn. 5225, from the H.B.K. herbarium, P), and accom-
panying photograph in the U.S. National Herbarium are those of
“A. lanata” of the original description. An annotation on the same
sheet by Dr. Agnes Chase, dated 1935,” reads: ‘‘No specimen named
‘A. lanata’ by Kunth has been found and no other named ‘A.
pubescens.’ ”’ This fragment is mentioned above as the possible type
of Agrostis pubescens, but the spikelets have villous rather than punc-
tate-scabrous glumes as A. pubescens should have from the description,
and so it is likely that this specimen is actually the type of A. lanata,
and that the type of A. pubescens has not been found.

From the descriptions of Kunth, the amount of pubescence on
the plant separated ‘Agrostis pubescens” from ‘A. lanata.’”? The
former was described as mostly glabrous except below the nodes and
the lemmas as pubescent toward the base; the latter as having pubes-
cent basal sheaths, nodes, and spikelets.

A large number of specimens of both types were collected by the
author and Prof. and Mrs. J. R. Reeder in the autumn of 1960 in
central Mexico. Specimens referable to both “species” were col-
lected near each other, in addition to specimens intermediate in
amount of pubescence between the two. This leaves little doubt
that the amount of pubescence is a variable character in these species
and that both types are representatives of one species, M. pubescens.
Special reference is made to the following comparison of specimens,
showing extreme and intermediate types of pubescence. All speci-
mens were collected in the same locality, 6 miles west of Navfos,
Durango, Mexico.

12Jn 1935, Dr. Agnes Chase, of the United States National Herbarium, criti-
cally studied and made notes on type specimens of grasses in many European
herbaria.

221-352—67——8

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

152

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SODERSTROM—-PODOSEMUM AND EPICAMPES 153

When correctly understood, M. pubescens embraces short to
medium-sized plants, the ligule is short (about 3 mm.), the usually
short blades of the culm are constricted and villous at the base, and
the lemma is pubescent on and between the nerves. The basal
sheaths, blades, and other parts are often densely villous, but the
amount varies and is often restricted to the lower part of the basal
sheaths or as sparse hairs over the various parts. Hitchcock (1935)
misinterpreted M/. pubescens; the plants he described have relatively
long ligules, glabrous vegetative parts, relatively long brownish
spikelets, and awnless or short-awned lemmas pubescent over the
lower part, on and between the nerves. Such specimens appear to
belong properly to M. distans. The two species, as treated in this
work, may be distinguished as follows:

spikelets
glumes
lemma

ligule

blades of culm
nodes

basal sheaths

M. pubescens

about 2-8 mm. long, white
to purplish

sparsely to densely villous,
obtusish

usually densely pubescent
throughout

usually 2-3 mm. long,
membranous throughout

constricted and villous at
at base

pubescent below

villous, or at least slightly

pubescent, with scattered

long hairs

17. Muhlenbergia robusta (Fourn.) Hitche.

N. Amer. Fl. 17(6):462.
Epicampes strictus J. S. Presl. Rel. Haenk. 1:235. 1830. Basis of Muh-

M. distans
averaging 3 mm, long

glaucous, with roughed
surface, acute
pubescent over lower part

5-10 mm. long, firmer and

browner at base than
above

not constricted and villous
at base

glabrous to scaberulous
below; rarely pubescent

glabrous to scaberulous

1935.

lenbergia presliana Hitche. (Mexico: without precise locality, Haenke).
Not M. stricta (Presl) Kunth. 1833.

E picampes berlandieri Fourn. Mex. Pl. 2:89. 1886. Basis of Muhlenbergia
fournieriana Hitche. (Mexico: without precise locality, Berlandier 670).
Not M. berlandierit Trin. 1841.

Epicampes robusta Fourn. Mex. Pl. 2:89. 1886. Basis of Muhlenbergia
robusta (Fourn.) Hitche. Fournier cites the following specimens in the
original description:

Bourgeau 683, supra Tacubaya (state of Mexico). He also cites this collec-
tion under Epicampes stricta Presl (p. 89). Hemsley (1885) likewise cites it
under E. stricta Presl. The lemmas are pubescent over the lower half and
from a study of a fragment it appears to be Muhlenbergia distans.

Bourgeau 1153, Santa Fée, vallée de Mexico. A complete specimen ex P
is in the U.S. National Herbarium. The lemmas are glabrous except for a
few minute hairs at the base. Fragments in an envelope labeled Bourgeau
1153 ex LE consist of two elements. In one, probably M. distans, the
lemmas are pubescent on the lower half and are awned. The other has

154 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

awnless lemmas but is pubescent on the lower midnerve and margins.
Neither of these can be Bourgeau 1153.
Hahn 420, Cuernavaca, [Morelos], Mexico. Not seen.

Muhlenbergia fourntertana Hitehe. Journ. Washington Acad. Sci. 23:453,
1933. Based on Epicampes berlandiert Fourn.

Muhlenbergia presltiana Hitche, N. Amer. Fl. 17(6):462. 1935. Based on
Epicampes strictus Presl.

Strongly caespitose perennial; culms stout, thick, erect, glabrous,
1-3 meters tall; nodes usually 4, glabrous below, the nodes covered by
the sheaths; basal sheaths strongly compressed-keeled, glabrous
(rarely scaberulous); ligule short, membranous, finely lacerated into
narrow strands and hairs, 1.5—5 (usually 2-4) mm. long, longer on the
innovations; rudimentary sheath auricles present, 1.5-5 (mostly 2-4.5)
mm. long, rarely lacking or more than 5 mm. long; blades about 25-50
(mostly 30-50) cm. long, 3.5-6 (usually 4-5) mm. wide, the upper and
lower surface scabrous, usually strongly so, the edges of the blades
sharply saw-toothed; upper throat scabrous; collar glabrous, rarely
scaberulous.

Panicle large, densely flowered, erect, purplish or greenish gray,
columnar before anthesis, the branches spreading at flowering and
becoming appressed at seed-set, 30-60 (usually 40-60) cm. long,
rarely as much as 70-80 cm. long, 2-4 em. wide (on herbarium speci-
men, much wider in field), the branches of the panicle naked below
or floriferous to the base; axis thick, angled, scaberulous or scabrous;
primary branches 4.5-17 (usually 5-8) cm. long, the secondary
branches 1.3-6 (usually 1.5-3) cm. long, the tertiary branches 0.7-2.5
(usually 1 or less) cm. long; pedicels scaberulous or scabrous, those
of the lateral spikelets very short, less than 1 mm. long, of the terminal
spikelets a little longer; spikelets about 1.8-3.3 (mostly 2.1-2.8) mm.
long, rarely longer; glumes acute to obtuse at the tip, equaling each
other or the second a little longer than the first, glabrous to even
scaberulous, purplish or greenish gray, longer than the floret, 1-nerved,
often indistinctly so; lemma acutish, glabrous or evenly scaberulous,
at most a few hairs at the base of the midnerve and margins, 3-nerved,
the median nerve often extending into a rudimentary awn from just
below the tip, usually less than 1 mm. long; callus of lemma glabrous
or with only a few hairs; palea a little shorter than the lemma, glabrous
or, at most, a few hairs between the keels toward the base.

Isolectotype in the U.S. National Herbarium, no. 999036, collected
in “Barranca prés Santa Fée, vallée de Mexico,” Oct. 3, 1865, by E.
Bourgeau (no. 1153).

Fournier cited three specimens in the original description. Of
these Hitchcock (1935b, p. 462) cited Bourgeau 1153 as the type,
and thus this is the lectotype.

SODERSTROM—PODOSEMUM AND EPICAMPES 155

Distripution: Steep rocky mountainsides, slopes of canyons and
barrancas, and eroded areas, at elevations of 3,000-7,000 ft., occurring
most frequently at 5,000-6,000 ft. The most common species of
Epicampes in southern Mexico, extending from Nayarit and Jalisco
to Chiapas and into Guatemala and Nicaragua.

MEXICO: Berlandier 670 (US fragments of type of EF. berlandieri, ex LE, P, W).
Haenke (US fragment of type of E. strictus). Palmer, in 1884 (DS, NY, US).
“Near river bank,” Schumann 1754 (US). Nayarit: SE. of Tepic, McVaugh
13393 (MEXU, SMU, US). Jauisco: Entre Pequeros y Tepatitla4n, Herndndez
4609 (CHAP). Zapotldn, Hitchcock 7172 (US), 7173 (US). Barranca de Oblatos,
Hitchcock 7335 (ILL, NY). La Barranca, Jones 27658 (MO). N. of Autldén,
McVaugh 19698 (MICH, US). SW. of Autlin, McVaugh 19899 (MICH, US).
Rio Blanco, Palmer 518, in 1886 (GH, MEXU, MO, NY, US). Guadalajara,
Pringle 2321 (F, GH, MO, NY, UC, US), 11746 (CAS, F, GH, MO, US). 18 mi.
S. of Guadalajara, Reeder & Reeder 2343 (YU). 3 mi W. of Tequila, Reeder &
Reeder 4206 (US, YU). Sierra Madre, W. of Bolafios, Rose 2997 (GH, US).
Ciudad Granja, W. of Guadalajara, Soderstrom 643 (US), 644 (US). SE. of
Guadalajara, on road to Aquascalientes, Soderstrom 653 (US), 654 (US). 6 mi.
E. of Santa Cruz, Soderstrom 566 (US), 567 (US), 568 (US), 569 (US), 570 (US),
571 (US), 572 (US). W. of Guadalajara on road to Tepic, Soderstrom 603 (US),
606 (US), 634 (US), 635 (US). 8 mi. E. of Guadalajara, near Barranca de
Oblatos, Soderstrom 678 (US), 679 (US), 680 (US). GuanasuaTo: Guanajuato,
Kenoyer 1765 (M. robusta X M. emersleyi?) (GH). 11 mi. NE. of Guanajuato,
Reeder & Reeder 3081, aff. M. robusta (YU). 2-4 km. E. of Guanajuato, Sohns
297 (M. robusta?) (MICH). Veracruz: Orizaba, Miller 2130 (US). Pursua:
Cerro Chiquihuite, Arséne 115 (US). Pursia: Cerro et fort de Guadalupe,
Arséne 1186 (US). Mayorazgo, Arséne 1275 (MO, US). Cholula, Arstne &
Nicolas 5441 (US); Nicolas, Sept. 15, 1910 (F). Rancho Posada near Puebla,
Nicolas 303 (US). Esperanza, Nicolas, Sept. 8, 1909 (NY). Acatzingo, Nicolas,
in 1909 (US). La Venta, Sharp 441664 (US). 6 mi. E. of Puebla, Soderstrom
387 (US), 388 (US), 389 (US), 390 (US). Zacatepec, Soderstrom 484 (US), 485
(US). Moretos: Tautepec, Gandara, in 1935 (US). Atlatlahucan, Gandara
(US). Teposteco, Lyonnet 2553 (US). Sierra de Ocuila rumbo Mexicapa,
Lyonnet 2872 (US). Casa de Tepoztero, Seler 4320 (GH). Mexico: Santa Fe
(as Santa Fée), Bourgeau 1153 (US lectotype, fragments ex LE, P). Carboneras,
Temascaltepec, Hinton 1853 (GH, NY). Rincén del Carmen, Temascaltepec,
Hinton 1959 (MO, NY, US). Temascaltepec, Temascaltepec, Hinton 2053 (LL,
MO, NY, US). Anonas, Temascaltepec, Hinton 2591 (CAS, LL, MICH, MO,
NY, US). Sultepec, Sultepec, Hinton 3219 (NY). Luvianos, Temascaltepec,
Hinton 5302 (NY). Tenango, Matuda 18846 (US). Guajimalpa to Rfo Hondo,
Matuda 21811 (US). Valle de Bravo, Matuda 27232 (MO). Carretera México-
Cuautla, Ortega & Vézquez, Oct. 12, 1960 (CHAP). 5 mi. N. of Tenancingo,
Reeder & Reeder 4173 (US, YU). Santa Fe, Rose & Painter 8009 (US). 3 mi.
NW. of Tlamanalco, Soderstrom 380 (US), 382 (US). Distrito FEDERAL: Chalco
region, Collins, Oct. 4, 1921 (US). Pedregal, Hitchcock 5939 (SMU, US). Near
Chapultepec Castle, Hitchcock 7839 (US). % mi. 8. of La Cima R.R. Station,
Iltis, Koeppen, & Iitis 146 (US, WIS). Pedregal de San Angel, Juzepczuk 216
(US); Lyonnet 1670 (US); Matuda 19479 (US). Churubusco, Orcutt 4317 (GH,
US). Coyoacdn, Ortega & Vazquez, Oct. 9,1960 (CHAP). Lava fields above the
Valley of México, Pringle 9077 (GH, MO, US). El Pedregal, Reeder & Reeder
3053 (YU). Huipulco, Rzedowski 1350 (IPN). Between Ixtle and El Conejo,

156 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Rzedowski 1929 (IPN). Eslava, Rzedowski 1999 (US). Sobre el Ixtle, Rzedowskz,
Dec. 21,1952 (IPN). About 5km. W. of San Andrés, Sohns 183 (US). Pedregal,
St. Pierre 804 (MICH, US), 805 (MICH, US). Dolores, St. Pierre 869 (US).
Ciudad Universitaria, Tateoka 1106 (US). Mricnoachn: Lomas de la Huerta,
Arsene 2628 (ILL, MO, NY, US), Oct 15, 1909 (GH). Barranca NW. de Pun-
guato, Arsene 2673 (ILL, MO, NY, US), 3145 (ILL, NY, TEX, US), Oct. 18, 1909
(GH). Loma Santa Marfa, Arsene, Sept. 2, 1909 (US). Zitdcuaro-Coyota,
Hinton 13409 (DS, GH, NY, US). 5 mi. N. of Cotija and 22 mi. S. of Jiquilpan,
King & Soderstrom 4648 (MICH, SMU, TEX, US). 22 mi. 8. of Uruapan,
King & Soderstrom 4859 (MICH, US). Below Tecambaro on road to Chipio,
Moore, Hernandez, & Porras 5616 (UC, US). Between Mil Cumbres and Ciudad
Hidalgo, Moore, Hernandez, & Porras 5774 (US). Road from Lorelia to Toluca,
Reeder, Reeder, & Goodding 1476 (YU). Zacapt, Ruppert & Guevara X-2830
(US). 3 mi. E. of Morelia, Soderstrom 544 (US), 545 (US), 545a (US). Quiroga,
Soderstrom 552 (US), 553 (US). Uruapan to Nueva Italia, Sehns 865 (US).
Oaxaca: Oaxaea, ITitchcock 6141 (US), 6154 (US). Reyes, Nelson 1778 (GH, US).
27 mi. NW. of Cd. Oaxaca, Reeder & Reeder 2183 (YU). Huajuapan de Leén,
Soderstrom 412 (US). Loc.?, Tateoka 1107 (US). Cutapas: Km, 1134, Baldwin
14345 (US). 15 mi. S. of Comitin, Reeder & Reeder 2096 (YU). 20 mi. E. of
Chiapa de Corzo, Reeder & Reeder 2128 (YU).

GUATEMALA: Huraverenanco: Huehuetenango, Melhus & Goodman 3562
(F). 10 km. W. of Aguacatdn, Williams, Molina, & Williams 21848 (US).
QUEZALTENANGO: Aguascalientes, Seler 3236 (GH). Guatemaua: Aguilar 311
(F). Guatemala City, Hitchcock 9035 (CAS, US), 9063 44 (US). Eureka, Hitch-
cock 9081 (MICH, US). Between Antigua and Guatemala City, Hitchcock 9146
(US). N. of Guatemala City, Popenoe 738 (US). SacaTerfiquEz: Santa Marfa,
Hitchcock 9129 (US). Antigua, Standley 61698 (F, US). CHIMALTENANGO:
Alameda, Johnston 976 (F). Finca La Alameda, near Chimaltenango, Standley
59136 (F, US). SE. of Patzim, Standley 61596 (F, US). Finca La Almeda,
near Chimaltenango, Standley 79806 (F), 79729 (F). Sono.Ld: Cerea del Lago
Atitlin, de Koninck 145 (US). Santa Rosa: Naranjo, Heyde & Lux 3932 (GH,
MO, NY, US).

NICARAGUA: Garnier 1953 (GH, US). Manaava: Sierra de Managua,
Garnier, “about 1930-1940” (F).

Muhlenbergia robusta is distinguished by its robust habit, very
short fimbriate ligule, rudimentary sheath auricles, and awnless or
short-awned, mostly glabrous, lemmas. Its closest relative appears
to be M. virletiz, which has a longer ligule (4-8 mm. rather than 2-4
mm.), rudimentary sheath auricles lacking, longer spikelets (mostly
3.5-3.6 mm. as opposed to 2.1-2.8 mm.), and the lemmas pubescent
toward the base of the margins. The sheath auricles, short ligule,
glabrous lemmas, and robust habit of MZ. macrotis also suggest a close
relationship with M. robusta.

Muhlenbergia robusta is the dominant species of Epicampes found
on rocky mountainsides in southern Mexico and might be thought of
as the counterpart of AZ. emersleyi Vasey which is dominant in north-
ern Mexico and southwestern United States. Asin the latter species,
M. robusta varies widely and some of the extreme forms have been
described as species. Hitchcock (1935b) recognized two species, J.

SODERSTROM—PODOSEMUM AND EPICAMPES 157

robusta and M. presliana, the latter based on the type of Epicampes,
EL. strictus Presl.

The present study indicates that M/. presliana and M. robusta are
members of one polymorphic species. A comparison of characters
from Hitchcock’s descriptions discloses that the major distinction
between the two species is spikelet size; in M. robusta it is 2.5-3 mm.
long and only 2 mm. in M. presliana. Spikelet size is a variable
character in all species of Epicampes. Within the M. robusta complex
the spikelets vary in size from 1.8-3.3 mm., although usually 2.1-2.8
mm. long. The spikelets of a fragment of the type of M. presliana
are mostly 2 mm. long, a few are 2.3mm. Specimens with spikelets
2 mm. long are similar to specimens of M. robusta with longer spikelets
and exhibit no other characters to delimit them as a distinct species.

In addition to variations of spikelet size and glume surface, the
branches of the panicle are naked at the base in some specimens and
in others floriferous to the base. Specimens exhibiting various com-
binations of these characters and intergrading forms have been found.
Nevertheless, all possess the robust habit, relatively glabrous lemmas,
short fimbriate ligule, and characteristic rudimentary sheath auricles
generally about 3 mm. in length. In view of the lack of correlation of
characters, it seems more realistic to regard MM. robusta and M.
presliana as members of a single variable species.

A rather unusual specimen, Kenoyer 1765 (Guanajuato, Mexico),
may possibly be a hybrid between M. robusta and M. emersleyi. The
Gray Herbarium specimen, the only one I have seen, has an incom-
plete culm and complete inflorescence. The latter is about 45 cm.
long with ascending branches. The ligule is short-lacerate and small
auricles are present, as in M. robusta, and the panicle is typical of
that species. Although the shape and appearance of the spikelets
are of that species, the lemmas are awned and slightly pubescent,
in the pattern of MZ. emersleyi.

18. Muhlenbergia virletii (Fourn.) Soderstrom, comb. nov.
Epicampes virletii Fourn. Mex. Pl. 2:88. 1886. (Virlet 1415, San Luis de
Potosi, Mexico)

Caespitose perennial; culms stout, erect, glabrous to scaberulous,
about 1 m. tall; basal sheaths strongly compressed-keeled, glabrous
to finely scaberulous; ligule membranous, lacerate above, 4-12
(usually 4-8) mm. long; sheath auricles lacking; blades of the culm
folded, about 30-40 cm. long, 2.5-6 (usually 2.5-3.5) mm. wide, the
upper and lower surfaces scabrous, the margins sharply saw-toothed;
upper throat scabrous; collar scaberulous.

Panicle erect, dark green, 29-55 (usually 30-50) em. long, 1.5-4
(usually 2-4) cm. wide, the branches ascending or spreading, naked

158 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

below, the spikelets clustered above; axis scabrous, primary branches
6-16.5 cm. long, secondary branches 2.5-5.5 cm. long; tertiary branches
1-1.5 em. long; pedicels shorter than the spikelets, 0.7-2 mm. long,
scabrous; spikelets 8-4 (mostly 3.5-3.6) mm. long; glumes equal to or
longer than the floret, usually equal to each other, or the second
glume 1-2 mm. longer than the first, green-translucent, often punctate-
roughened, acute at tip, indistinctly 1-nerved; lemmas acuminate,
2.7-3.8 mm. long, sparsely to densely pubescent on the lower margins,
glabrous dorsally (rarely a few sparse hairs on the midnerve below),
3-nerved, awnless or the awns rudimentary, no more than 0.5 mm.
long; callus of lemma pilose; palea shorter than the lemma, glabrous
or with a few hairs between the nerves.

Fragments of panicle of type, ex P-CO and Fournier herbarium (P)
in the U.S. National Herbarium, no. 73225, collected in ‘Mexique,
Prov. de San Luis,” in 1851, by Virlet d’Aoust (no. 1415).

Distrisution: Arroyos and rocky hillsides in the pine-oak zone,
at elevations of about 6,600-8,400 ft. Infrequent from San Luis
Potosf in central Mexico, south to Puebla. A single collection from
Durango is doubtfully referred to this species.

Mexico: Durango: Km. 143 Durango-Mazatldn, Hernandez & Tapia N-849
(M., virletii vel aff.) (CHAP). San Luts Porosf: Virlet 1415 (US fragment of type,
ex P). Pursua: Manzanilla, Arséne 1693 (GH, ILL, MO, NY, US). Km. 47,
camino de Puebla, Matuda 18611 (US). Moregtos: Lagunas Zempoala, Lyonnet
2515 (US). Mexico: Comunidad, Temascaltepec, Hinton 2439 (MICH, NY,
US). Sultepec, Sultepec, Hinton 3219 (ILL, MO, NY, US). 23 mi. SE. of
México on highway 190, Pratt 751 (TEX). Near Santa Fe, Rose & Painter
8010 (NY, US). E. of Ixtlahuaca, Soderstrom 512 (US). Distrito FEDERAL:
Desierto de Leones, Matuda 25930 (US). Cerca del Xitle, Rzedowski 2167 (IPN).
Western part of the Pedregal, Rzedowski 2242 (US). Eslava, St. Pierre 813 (US).
Micnoacdn: Quinceo, Arséne 3201 (ILL, MO, NY, US). Morelia, Arséne 6999
(US). Ca. 18 mi. S. of Pdtzcuaro, King & Soderstrom 5200 (US).

Muhlenbergia virletit is distinguished by the very long green spikelets,
the lemmas glabrous except toward the base of the margins, the
medium ligule, and’ the absence’ of sheath auricles. In its nearest
ally, MM. robusta, rudimentary sheath auricles are usually present, the
spikelets are shorter, the lemmas are mostly glabrous throughout,
and the ligule is shorter (mostly 2-4 mm. as opposed to 4-8 mm.
long). The glumes of M. robusta are generally purplish or ashy gray;
those of M. virletii are dark green and appear translucent.

Glume surface is more or less glabrous in some specimens, in others
scaberulous or scabrous. Specimens collected in the same locality
exhibit such differences. In spikelets of specimens previously referred
to M. presliana the glumes are glabrous, although Hitchcock described
them as scabrous. Specimens have been cited, however, with spikelets
about 2 mm. long and the glumes scabrous, e.g., Hitchcock 9129

SODERSTROM—PODOSEMUM AND EPICAMPES 159

(Guatemala), Géndara (Morelos, Mexico), and Moore, Herndndez, &
Porras 5616 (Michoacén, Mexico). Furthermore, specimens were
found with the glumes glabrous and the spikelets similar to those of
the type of LE. strictus, except larger, 2.5-3 mm. long, e.g., Rzedowski
1999 (México, D. F.). Sohns 183 (México, D. F.) has spikelets similar
to those of Rzedowski 1999 but the glumes are scabrous.

19. Muhlenbergia breviligula Hitchc. N. Amer. Fl. 17(6) :458. 1935.

Strongly caespitose perennial; culms stout, glabrous to scaberulous,
75-141 cm. tall; nodes 3, glabrous or slightly puberulent below, some-
times puberulent above the nodes; basal sheaths compressed-keeled,
the old basal sheaths persistent, becoming brown, curled and fibrillose;
ligule of the culm leaves a membranous rim of hairs 0.1-2 mm. long
(usually negligible, less than 1 mm. long) those of the innovations
sometimes over 1 cm. long; sheath auricles lacking or rudimentary,
whitish, 1.5-6 mm. long when present; blades of the culm flat or
folded 24-47 cm. long 2-3 (mostly 2.5) mm. wide when flat, the
upper surface glabrous to scaberulous, the lower surface scaberulous
to scabrous; upper throat scabrous, often pubescent; collar scabrous
or puberulent.

Panicle nodding, rather dense, yellowish to purplish brown or
purplish green, the branches ascending or opening outward, 30-51
(mostly 30-40) cm. long, 3-9 (mostly 3-5) em. wide; axis scabrous,
the primary branches 6-12 cm. long, the secondary branches 1.5-7 cm.
long, the tertiary branches 0.5-3.5 cm. long; pedicels of the lateral
spikelets 0.5-1.5 mm. long, those of the terminal spikelets to 3 mm.
in length, scaberulous on the lower part, becoming scabrous just
below the spikelet; spikelets 2-3 mm. long or sometimes longer (mostly
2.5-3 mm.); glumes about equal, equaling or a little longer than the
floret, glabrous (scaberulous at the tips under high power), acute,
often slightly awned (awns to 0.2 mm. long), hyaline, faintly
1-nerved; lemmas 2.5-2.6 (usually about 2.5) mm. long, 3-nerved,
pubescent on the lower two-thirds of the midvein and lower margins
(rarely with hairs between the nerves below), bearing an awn from
below the tip, 0.7-1.7 (this usually more than 1) cm. long, flexuous;
callus of lemma sparsely pilose; palea a little shorter than the lemma,
sparsely pubescent on the lower two-thirds between the keels.

Holotype in the U.S. National Herbarium, no. 995888, collected
on clay hill, Guatemala City, Guatemala, alt. 1,500 m., December 2,
1911, by A. S. Hitchcock (no. 9063).

Distrispution: Rocky hillsides and exposed pine-oak areas, at
elevations of 2,000-4,500 ft. Apparently common in Guatemala and
Honduras.

GUATEMALA: Hurnvuerenanco: W. of Aguacatdn, on the road to Hue-
huetenango, Standley 81190 (F). Aura Verapaz: Sequanquim, Cook & Doyle

160 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

60 (US). Zacapa: Volein Gemelos, Sleyermark 43251 (I). CHIquimuLa:
Between Chiquimula and Montafia Barriol, 3-15 miles NW. of Chiquimula,
Steyermark 30649 (F, US). Januapa: Vicinity of Jalapa, Standley 76731 (F,
US). Guaremana: Guatemala City, Hitchcock 9063 (US holotype); Hitchcock
9064 (US); Hitchcock 9109. Juriapa: Between Jutiapa and Plan de Urrutia,
Standley 75582 (I).

HONDURAS: Comayacua: Between Comayagua and Villa San Antonio,
Archer 3842 (CAS). Francisco MorazAn: 30 km. W. of Tegucigalpa, Archer
3846 (US). Rfo Yeguare at about longitude 87° W. and latitude 14° N., Molina
1593 (US). Rfo Yeguare a Danii, Rodriguez 1564 (F, GH). Above El Zamorano,
Standley 14630 (F); Standley 24850 (F). San Antonio del Oriente, above San
Antonio, Swallen 10968 (US). Mt. Uyuea, beyond Las Floras, Swallen 11323
(US). En Parafso: Mt. Yuscardin, Molina 605 (F, GH, US). Giiinope,
Rodriguez 1848 (F). Las Casitas, road to Giiinope, Swallen 11081 (US). Road to
Danlf, beyond Rio San Francisco, Swallen 11216 (US). Road to Yuscardn, Swallen
11359 (US); Swallen 11367 (US). 5 km. 5S. of Ojo de Agua, Williams & Molina
11220 (F, GH, US). CnoLureca: San Marcos, Williams & Molina 10921 (FP,
GH, MICH, MO, US).

Muhlenbergia breviligula is distinguished by the very short ligule,
long-awned lemmas pubescent on the lower two-thirds of the midnerve
and margins, and spikelets usually 2.56 mm. or longer. In overall
aspect it resembles most closely MM. emersleyi, but in that species the
ligule is usually 1 cm. or longer, the blades are considerably broader
at the base, and the basal sheaths are not so persistent and fibrillose
when old. Muhlenbergia aurea is closely allied by the very short
ligule and long-awned lemmas; however, its spikelets are small,
2-2.2 mm., and the lemmas are only pubescent toward the base.

20. Muhlenbergia aurea Swallen. Contr. U.S. Nat. Herb. 29:411. 1950.

Strongly caespitose perennial; culms stout, glabrous, becoming
scaberulous to scabrous toward the panicle, about 34 m. tall, glabrous
beneath the nodes; basal sheaths compressed-keeled, glabrous at
the base, becoming scabrous toward the collar; ligule a short mem-
branous rim appressed against the blade and hardly noticeable,
0.3-1.5 mm. long; sheath auricles rudimentary, 0.8-3 mm. long;
blades of the culm to 42 cm. long, flat or folded, 2.5-5 mm. wide
when unfolded, the upper surface scaberulous, the lower surface
scabrous, the edges of the blade saw-toothed; upper throat scabrous;
collar scabrous.

Panicle nodding, dense, golden yellow, about 36 em. long, 7-9 em.
wide, the floriferous branches of the panicle appressed; axis scabrous;
primary branches of the panicle 15-17 cm. long, secondary branches
4—6 cm. long, tertiary branches 1.5-2 cm. long; pedicels 1.5-2 mm.
long, scabrous; spikelets 1.7-2.2 (mostly 2-2.2) mm. long; glumes
about equaling the lemma, the second a little longer than the first,
scabrous, very acute or slightly awn-tipped, especially the second
glume, hyaline, 1-nerved; lemmas 1.9 mm. long, pubescent on the

SODERSTROM—PODOSEMUM AND EPICAMPES 161

lower third of the midvein and margins, acute, indistinctly 3-nerved,
bearing an awn from just below the tip, mostly 1-1.5 cm. long; callus
of the lemma with short hairs; palea a little shorter than the lemma,
elabrous.

Holotype in the Field Museum of Natural History, no. 1057948,
collected on lower south-facing slopes of Voledn Santa Maria de
Jestis and Calahuache in thickets at base of vertical slopes along
r.r., Dept. of Quezaltenango, Guatemala, alt. 1,800-1,500 m., Dee. 31,
1939, by J. A. Steyermark (no. 33175). A fragment of the holotype
and photo are in the U.S. National Herbarium.

Distripution: Known only from the type collection.

Muhlenbergia aurea is distinguished by the very short ligule,
slightly awn-tipped glumes that equal the floret, and long-awned
lemmas pubescent on the lower third of the midnerve and margins.
In the original description the author remarked, “MJuhlenbergia
aurea is probably most closely related to M. scoparia Vasey, which
differs in having much narrower sheaths and blades, an elongate
ligule, a narrow, less densely flowered, purple panicle, and longer
awns.” Although related to M. scoparia, its closest relative appears
to be M. brevligula, with which it shares the characteristics of a very
short ligule and subequal slightly awn-tipped glumes (in M. breviligula
the glumes are acute or occasionally somewhat awn-tipped). In
M. scoparia the ligule is usually 1 cm. long or more and the glumes
are unequal, a little shorter than the floret, and longer awn-tipped.
Muhlenbergia aurea and M. breviligula occur much farther south
than M. scoparia. Muhlenbergia aurea is known only from the type
collection in Guatemala, M. breviligula from Guatemala and Honduras,
and M. scoparia from northern and central Mexico.

21. Muhlenbergia grandis Vasey. Contr. U.S. Nat. Herb. 1:283. 1893.

Epicampes grandis (Vasey) Beal. Grasses N. Amer. 2:309. 1896. Based on
Muhlenbergia grandis Vasey.

Epicampes bourgaet var. grandis (Vasey) Jones. Contr. West. Bot. 14:7,
1912. Based on Muhlenbergia grandis Vasey.

Caespitose perennial; culms glabrous, 88-164 (mostly 100-150)
cm. tall; nodes 4, pubescent below (rarely above); basal sheaths
compressed-keeled, glabrous, the old ones inflated, papery, brown or
usually reddish brown, becoming fibrillose; ligule membranous,
lacerate, 1.5-6 mm. long; sheath auricles lacking; blades of the culm
15-34 (mostly about 30) cm. long, flat or folded, 3-8 mm. wide when
flat (usually 3-5 mm.), the upper surface of the blade scaberulous to
scabrous, the lower surface scaberulous to scabrous, the upper and
lower surfaces of the same blade alike; upper throat pubescent; collar
a fringe of short white appressed hairs, rarely glabrous.

162 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Panicle erect or somewhat nodding, usually very dense, the panicle
branches mostly appressed, the groups of branches borne at intervals
along the axis, yellowish or yellowish brown, 30-63 (usually 30-50)
em. long, 2-8 (usually 4-5) em. wide; axis scaberulous, sometimes
with a few hairs, ribbed, the primary branches to 9 (usually 7-9) em.
long, the secondary branches 1.5-4.5 cm. long, the tertiary branches
1-2 cm. long; pedicels of the lateral spikelets 0.5—-1.5 mm. long, those
of the terminal spikelets longer, to 2.5 cm., glabrous to scabrous, at
least scaberulous under the spikelet; spikelets 1.3-2.5 (usually 1.5-2)
mm. long; glumes usually about equal to the floret, equal or subequal
(the second a little longer than the first), glabrous to scaberulous,
roundish at the tips, somewhat erose at the tips (under high power),
awnless, hyaline, nerveless; lemmas 1.5-2.2 (usually 1.8-2.1) mm.
long, glabrous except for hairs on the lower part of the midnerve,
acutish at the tip, 3-nerved, bearing a flexuous awn from below the
tip, 6-19 (usually more than 10) mm. long; callus of lemma con-
spicuously short-pilose; palea a little shorter than the lemma,
glabrous or with a few hairs at the base between the keels.

Type in the U.S. National Herbarium, no. 73222, collected in Rio
Blanco, Jalisco, Mexico, 1886, by Edward Palmer (no. 515).

Distrisution: Rocky mountainsides and rocky slopes of barrancas
and canyons, at elevations of 3,000-5,000 ft. Of limited distribution
in western Mexico, occurring from Sonora, south into Nayarit and
Jalisco.

MEXICO: Palmer 170, in 1885 (TAES), in 1885 (GH, NY). Sonora: Cafion
Estrella, Dist. Alamos, Geniry 332 M (MICII). Nayanir: 10 mi, SE. of Tepic,
McVaugh 13377 (MEXU, SMU, US). 10 mi. SE. of Ahuacatlin, McVaugh &
Koelz 743 (MICH). Jatisco: SW. Authin, Hernandez 4637 (CHAP, US).
1-2 mi. N. of Ahuacapin, McVaugh 19531 (MICH, US). Ca. 8 mi. 5. of La
Resolana, McVaugh 21086 (MICH, US). Rio Blanco, Palmer 415, in 1886 (F),
515, in 1886 (US type), 515a, in 1886 (GH, MO, US), 682, in 1886 (GH, NY, US).
Guadalajara, Pringle 1760 (F, UC, US, paratypes), 2362 (ARIZ, CAS, GH,
MICH, SMU, UC, US), 2765 (F, MO, UC, US fragment), 11751 (CAS, F, GI,
MICH, MO, US). 2 mi. W. of Tequila, Reeder & Reeder 4205 (US, YU).

Muhlenbergia grandis is distinguished by the reddish-brown basal
sheaths, collar a dense fringe of short hairs, the short spikelets, and
long-awned lemmas short-pilose on the callus. It appears to be most
closely related to M. scoparia with the reddish-brown sheaths unique
to both species; the short spikelet, long-awned lemma, and short-
pilose callus are features of both. The glumes of M. grandis are about
equal and shorter than the floret, but are unequal in M. scoparia,
long-awned, and about equal to the floret. The ligule of MZ. grandis
is usually no more than 6 mm., as opposed to 8-16 mm. long in AZ.
scoparia.

SODERSTROM—PODOSEMUM AND EPICAMPES 163

22. Muhlenbergia distans Swallen. N. Amer. Fl. 17(6) :461. 1935
Muhlenbergia pubescens (II. B. K.) Hitche. N. Amer. Fl. 17(6):460. 1935 (as
to description only, not as to name).

Caespitose perennial; culms erect, 66-154 cm. tall, glabrous below,
becoming scaberulous or scabrous below the panicle; nodes 2-4
on the lower part of the culm, puberulent to pubescent below, especially
of the lower nodes; basal sheaths compressed-keeled, glabrous, the
keels scaberulous; ligule membranous, firmer below but not decurrent,
lacerate above, 0.5-1.5 cm. (usually 5-10 mm.) long; sheath auricles
lacking; blades of the culm 11-62 (usually 25-40) cm. long, folded,
narrow, 1-5 (mostly 2-3) em. wide when flat, the edges saw-toothed;
upper surface of blade mostly scaberulous, the lower surface mostly
scabrous; upper throat scaberulous to scabrous; collar scaberulous
to scabrous (mostly scaberulous).

Panicle erect, narrow, greenish brown, 24-40 cm. long, 1.5-4
(usually 2-4) cm. wide, the densely flowered branches ascending,
floriferous almost to the base; axis scabrous; primary branches 5.5—9
em. long, the secondary branches 1-5 (mostly 2-3) cm. long, the
tertiary branches mostly 0.5 cm. long; pedicels scabrous, especially
just below the spikelet, shorter than the spikelet, those of the lateral
spikelets about 1 mm. long, those of the terminal spikelets 1.5-2
mm. long; spikelets 2.7-3.6 (usually 3-3.3) mm. long; glumes broadly
acute at the tip, rarely a bit dentate, glabous-appearing with a rough-
ened surface, the second glume 1-4 mm. longer than the first, in-
distinctly 1-nerved; lemmas acutish, 2.7-3.4 mm. long, shorter
than the glumes, lightly pubescent on and between the nerves over the
lower part, the upper part scabrous, the lateral margins densely
pubescent toward the base, 3-nerved, the median nerve in many or all
of the florets of the panicle extending from below the tip into a short
awn, 1.5-5 (at least less than 5) mm. long; palea pubescent on the
lower part.

Holotype in the U.S. National Herbarium, no. 746297, collected
on dry hills, Las Sedas, Oaxaca, Mexico, alt. 6,000 ft., Sept. 15, 1894,
by C. G. Pringle (no. 5575). Isotype at F.

DistrisuTion: Rocky slopes and barrancas, in the pine-oak zone,
at elevations of 5,000-8,500 ft. Ranging in Mexico from Durango
and San Luis Potosi southeastward into Guatemala. Apparently
infrequent.

MEXICO: Duranao: (?) Navfos, Reeder, Reeder, & Soderstrom 3379 (US, YU).
(?) Llano Grande, Reeder, Reeder, & Soderstrom 3396 (US, YU), 3397 (US, YU),
3398 (US, YU). Zacatecas: W. of Zacatecas on highway to Villanueva,
Soderstrom 724 (US). San Luis Potosf: 8 mi. W. of Santa Catarina, Reeder &
Reeder 4103 (US, YU). SW. of San Luis Potos{ on road to Aguascalientes,

Reeder, Reeder, & Goodding 1387 (YU). SW. of San Luis Potosf, Reeder, Reeder,
& Soderstrom 3327 (US, YU), 3331 (US, YU). Puerta Huerta in the Sierra de

164 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Alvarez, Sohns 1044 (US). Jatisco: Paso de Troje, SW. of Ojuclos on road to
Aguascalientes, McVaugh 16801 (MICH, US). Hrpateo: Pachuca, Hitchcock
6708 (US); Purpus, Sept. 1905 (UC). Ixmiquilpan, Purpus 1466 (UC). Puebla:
Cerro Tepoxuchil, Arstne 1901 (GH, MO, US). Acatzinco, Distrito de Tepeaca,
Arsene (Bro. Amable) 2265 (MO, US). Fort de la Guadalupe, Arsene (Bro.
Nicolas) 5436 (US). Between haciendas Santa Barbara and Cristo, Arséne, Aug.
1, 1907 (MO, US). Esperanza, Hitchcock 6485 (US). NE. of Zacatecas,
Soderstrom 486 (US), Tuaxcata: Km. 70 de la carretera Apizaco-México,
Guerrero, Sept. 25, 1962 (IPN). Mmxtico: “Pédregal prés Mexico,” Bourgeau
683(?), 1865-1866 (US fragment ex P). Tenancingo, Matuda 28985 (US).
Distrito FEepERAL: Xochitepec, Garcia 86 (IPN). Lomas de Santa Fe, Lyonnet
55 (GH, MO, NY, US). Barrancas de Mixcoac, Lyonnet 2369 (US). Zacatepec,
pedregal de Tlalpan, Matuda 21358 (US). Near Tlalpan, Rose, Painter, & Rose
8255 (US). On pedregal, Rose, Painter, & Rose 9456 (US). Tacubaya, St.
Pierre 313 (MICH, US). Tacubaya, Dolores, St. Pierre 862 (MICH, US), 866
(MICH, US), 869 (US). Jajalpa, St. Pierre, 929 (CAS, MICH, US). Micnoa-
ckn: N. of Loma del Zapote, Arsene, Sept. 26, 1909 (MO, US). Oaxaca: De
Las Sedas a Felix Mahuaca, Conzatti 4870 (CAS). Las Sedas, Pringle 5574 (F),
5575 (US holotype, F isotype); Smith 916 (MO).

GUATEMALA: Zacapa: Sierra de Jas Minas, below Finca Alejandria,
Steyermark 29773 (EF).

Muhlenbergia distans is distinguished by the greenish-brown
spikelets, usually 3 mm. long or more, lemmas pubescent on the lower
part, on and between the nerves, and ligules mostly 5-10 mm. long.
The panicles are narrow, densely flowered, and have a distinctive
brownish color. The basal sheaths are characteristically dark brown
and firm when old. Apparently it is most closely related to M.
versicolor, having in common the relatively greenish spikelets consist-
ing of subequal glumes, lemmas pubescent on and between the
nerves, and nodes pubescent below. In M. versicolor the glumes are
often awn-tipped (especially the longer second glume) but only
slightly dentate at the tip and not awned in AM. distans. The lemmas
of MM. versicolor bear long awns on all the spikelets of the panicle;
in typical AZ. distans short-awned and awnless spikelets occur in the
same panicle. Moreover, Af. versicolor is generally characterized by
sheath auricles, which are lacking in AZ. distans.

Muhlenbergia distans was described from a specimen collected on dry
hills, Las Sedas, Oaxaca, Mexico, in 1894 and has been known only
from the type collection. Tt is a small tufted perennial with narrow
culms and panicles with loosely ascending branches. The brownish
spikelets range in length from 2.7~3.5 mm., the somewhat scaberulous
glumes are acutish, and the short-awned lemmas are lightly villous
on and between the nerves for a greater part of the length, with
densely pubescent margins. Specimens previously referred to M.
pubescens (H.B.K.) Hitehe. appear to belong to this species. As the
name M. pubescens (H.B.K.) Hitche. is untenable for these speci-
mens, AM. distans Swallen is the proper epithet. The majority of

SODERSTROM—PODOSEMUM AND EPICAMPES 165

cited specimens differ from the type of WM. distans in being more robust
and having thicker culms and denser panicles. However, the basic
characters are similar, for the brownish spikelets are about 3 mm.
Jong, the short-awned lemmas are pubescent on and between the
nerves, although the pubescence is confined mostly to the lower
half, and the ligules are about 5-10 mm. long. The holotype of
M. distans appears to represent a smaller than typical and less robust
representative of the species. Conzatti 4870, collected near the type
locality, is more robust and has larger panicles than the type and is
in all respects more like the majority of specimens referred to M.
distans. Such differences as size and looseness versus denseness of
panicles are variable characters within species of Epicampes. The
spikelet and ligule are the more stable characters and are of ereater
importance in referring these specimens to M. distans.

Three collections (Reeder, Reeder, & Soderstrom 3379, 3396, and
3398) are doubtfully referred to M. distans. These specimens have
long spikelets with the lemmas pubescent over the lower part and
ligules about 1 cm. long. The habit of the panicles and plants, how-
ever, is more like that of Af. emersleyi than of M. distans.

23. Muhlenbergia longiglumis Vasey. Contr. U.S. Nat. Herb. 1:283. 1893.

Caespitose perennial; culms glabrous, 3-1 m. tall; nodes 2 (or 3?),
pubescent below; basal sheaths compressed-keeled, glabrous, the old
ones becoming brown and fibrous; ligule membranous, brownish and
firmer on the lower part, thinner and lacerate above, 4-10 mm. or
longer; sheath auricles lacking; blades of the culm 20-30 em. long,
folded, 1.5-2 mm. wide when flat; upper surface of the blade scab-
erulous to scabrous, the lower surface mostly scabrous (rarely glab-
rous); upper throat scabrous; collar scabrous.

Panicle erect or somewhat nodding, narrow, 20-40 (mostly about
30) cm. long, 1-2.5 cm. wide (excluding awns), the branches of the
panicle about 3-3.5 cm. long, ascending, borne in verticils of ap-
proximate clusters of 2-6 at intervals along the axis, the body of the
panicle light green, sharply contrasting with the dark purple awns;
axis scaberulous, ribbed; pedicels of the lateral spikelets 2-3 mm.
long, those of the terminal longer, scabrous on the narrow lower
portion, pubescent on the swollen upper portion subtending the
spikelet; spikelets 5-7 (mostly 6-7) mm. long; glumes about equal,
a little longer than the floret, the surface with sparse short hairs,
attenuate, awn-tipped (the awn 1 mm. long or less), grayish green,
hyaline, 1-nerved; lemmas 4~5 mm. long, the margins densely pu-
bescent on the lower 4-4, internerves elabrous or slightly pubescent
near the base, remainder of lemma glabrous to scaberulous, lanceolate,
3-nerved, bearing a flexuous awn from below the tip, 2-6.5 em. long;

166 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

callus of lemma bearing a dense ring of white hairs up to 1.5 mm.
long; palea about equaling the lemma, glabrous or sparsely pubescent
between the keels; anthers 3, yellow or greenish yellow, 2~3 mm. long.

Holotype in the U.S. National Herbarium, no. 2383429, collected
near Guadalajara, Jalisco, Mexico, July-October 1886, by Edward
Palmer (no. 766).

Disrripution: Rocky mountainsides in the pine-oak zone, at eleva-
tions of 5,000-7,000 ft. Apparently of limited distribution, all collec-
tions except one being from Jalisco; the one from Aguascalientes is
from near the Jalisco border, and possibly on the Jalisco side.

MEXICO: Palmer, in 1886 (MO, may be part of Palmer 766 in 1886). AGuaAs-
CALIENTES: Sierra del Laurel, ca. 10 mi. SE. of Calvillo, McVaugh & Koelz 171
(MICH). Jaxisco: 4-4.5 mi. NW. of Cuautla, road to Los Voleanes, McVaugh

20462 (MICH). Guadalajara, Palmer 766, in 1886 (US, holotype) ; Pringle 2365
(US), 11752 (GH, US).

Muhlenbergia longiglhumis is distinguished by its awn-tipped, long
elumes, light grayish green in contrast to the long dark purple awns of
the lemmas, and lemmas glabrous except at the base. The purplish
ereen panicle is not unlike that found in AZ. versicolor, although the
species is probably most closely related to M. emersleya with its long
ligule, pyramidal panicle, and generally similar habit.

24, Muhlenbergia emersleyi Vasey. Contr. U.S. Nat. Herb. 3:66. 1892.

Muhlenbergia distichophylla sensu Vasey in Rothrock, Bot. Wheeler Surv.
283. 1878. Not Podosaemum distichophyllum J. S. Presi in K. B. Presl,
Rel. Waenk. 1:231. 1830=Muhlenbergia distichophylla (J. S. Presi)
Kunth. Description based on Rothrock 282 and 283.

Muhlenbergia vaseyana Scribn. Rep. Missouri Bot. Gard. 10:52. 1899. Based
on Muhlenbergia distichophylla sensu Vasey, not Pedosaemum distichopyllum
J. S. Presl, 1830. Syntypes (by inference) Rothrock 282 and 283, from
Rocky Canyon, Arizona, the basis of Vasey’s description as cited above.
Rothrock 282 is chosen as lectotype.

Epicampes stricta var. distichophylla sensu Jones, Contr. West. Bot. 14:6.
1912, in part as to the synonyms cited, M. vaseyana and M. emersleyi, not
as to the basionym, Podosaemum distichophyllum J. 8. Presl.

Epicampes emersleyi (Vasey) Hitche. Bull. U.S. Dept. Agr. 772:144, 1920.
Based on Muhlenbergia emersleyi Vasey.

Epicampes subpatens Hitche. Bull. U.S. Dept. Agr. 772:144. 1920. (Hutch-
cock 13541, Guadelupe Mountains near Queen, New Mexico.)

Caespitose perennial; culms erect, narrow to thick, glabrous (rarely
roughened), 1-1.5 meters, occasionally to 2 meters tall; nodes 3-4,
clabrous or somewhat puberulent beneath; basal sheaths compressed -
keeled, glabrous; ligule long, membranous, usually a little firmer at the
base than above where thin and usually lacerated into narrow strands,
ligules of the innovations long, narrow, and membranous throughout,
conspicuous; sheath auricles lacking; blades of the culm 23-47 (mostly

SODERSTROM—PODOSEMUM AND EPICAMPES 167

20-40) cm. long, 2-6 mm. wide, flat or folded, the upper surface gla-
brous to scaberulous, the lower surface scabrous, at least the lower
surface more scabrous than the upper surface of the same blade, the
edges saw-toothed; upper throat scaberulous to scabrous (mostly
scabrous); collar scaberulous.

Panicle typically lax above, the branches loosely ascending or
spreading, the panicle pyramidal, light purplish to purplish tan,
20-40 (mostly 30-40) cm. long, 4-7 cm. wide (often much wider in
the field) ; axis glabrous to scaberulous (rarely scabrous), the branches
of all the orders naked below, the primary branches 9-17 (usually
9-12) em. long, the secondary branches 3-9 (mostly 3-5) cm. long, the
tertiary branches 0.5-4.5 (mostly 0.5-1.5) em. long, quaternary
branches sometimes present, as much as 2 cm. long; pedicels glabrous
to seaberulous (sometimes scabrous), more scabrous just below the
spikelet, pedicels of the lateral spikelets 0.5-1.5 mm. long, those of
the terminal spikelets 1.5-3 mm. long; spikelets 2.2-3.2 mm. long,
occasionally shorter or longer (usually 2.5-3, but most frequently
2.5-2.7 mm. long); glumes broadly acute, about equal to each other
or the second glume 1-2 mm. longer than the first, mostly scaberulous
to evenly scabrous, indistinctly 1-nerved; lemmas acutish, a little
shorter than the glumes, pubescent on the midnerve and margins
about one-half to three-quarters of the length, the pubescence varying
from sparse to rather dense, that of the margins generally denser
than that of the midnerve, the lemma 3-nerved, the middle nerve
often excurrent from below the tip into an awn as long as 15 mm.,
generally purplish and flexuous, the awn persistent or deciduous,
breaking off easily at the point of attachment to the lemma; palea
a little shorter than the lemma, more or less pubescent between the
keels for one-half to three-quarters of the length.

Type in the U.S. National Herbarium, no. 73223, collected in
southern Arizona in 1890, ‘‘a coarse range grass that stock do not like,”
by J. D. Emersley (no. 46).

DisrriButTion: Rocky mountain slopes, barrancas, canyons, ar-
royos, and rocky ledges, at elevations of 4,500-7,500 ft., occurring
most frequently at about 6,000-7,000 ft. ‘The most common species
of Epicampes, ranging from Texas to Arizona into northern Mexico,
becoming less frequent south of Durango to Oaxaca.

UNITED STATES: Texas: Hueco, Mulford 80 (ILL,MO, NY). Nealley, in
1889 (DS, MICH, NY, UG, US), in 1890 (NY, US). “Common all over mts.,”
Swallen 1101 (US). Pine Spring, Whitehouse 8533 (US). ‘Expedition from west-
ern Texas to El Paso,” Wright 729 (UC, US), 730 (GH, UC, US), 736 (US).
BREWSTER county: Chisos Mountains, Bailey 392 (US). Green Gulch, Berkman
Sept. 10, 1925 (TEX). Highway 223, Brown 30 (TIX). 6 mi. 8. of Alpine, Brown
47 (TEX). N. of Alpine, Carter 19 (US). Blue Creek Trail, Cory 7272 (POM).
Davis Mountains, 6 mi. W. of Alpine, Cory 40248 (TEX), 40413 (TAES).

221-352-677

168 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Chisos Mountains, Havard, August 1833 (US). Alpine, Hitchcock 13594 (US).
5 mi. SW. of Marathon, Johnston 3336 (CAS, TEX). Elephant Mesa, Johnston
6427 (LL). Green Gulch, Lundell 13250 (LL, UC). Chisos Mountains, Marsh 56
(F). Lower Pine Canyon, Marsh 282 (GH, TAES, TEX), 322 (GH, TAES, TEX,
US). Upper Pine Canyon, Marsh 351 (GH, TEX). Big Bend National Park,
Marsh 51-1153 (TEX). Boot Spring, Moore & Steyermark 3181 (GH, MICH,
MO, NY, UC, US). Emory Peak, Mueller 7850 (GH, MICH, NY, SMU, TAES,
TEX, UC, US). Chisos Mountains, Muller, July 25, 1932 (DS, GH, UC). Blue
Creek Canyon, Parks & Cory 7270 (TAES). Upper Blue Creek, Parks & Cory
7271 (TAKES). Boot Ravine, Parks & Cory 30350 (TAKES). Iron Mountain,
Parks & Cory 30567 (TAES). Big Bend, Reed 90 (SMU), 99 (ARIZ). Iron
Mountain, Reed 109 (UC), 112 (CAS). Hidden Valley, Rose-Innes & Moon 1266
(TEX). Big Bend National Park, Shinners 8799 (SMU). SR Hill, Sperry T487
(ARIZ, TAES). Green Gulch, Sperry 1805 (TAES). Casa Grande, Warnock
806 (DS, SMU). Mt. Emory, Warnock 12378 (TAES). MelIntire Hill, Warnock
20270 (TEX). Split Tanks, Warnock 20766 (TEX). Baldy Peak, Warnock 20805
(TEX). Goat Mountain, Warnock 21470 (TEX). Old Marfa highway 2 mi. 8.
of Alpine, Warnock 21731 (TEX). S. of Alpine, Warnock, May 5, 1938 (ARIZ,
GH). Blue Creek Canyon, Wolff 4693 (TAES), 4698 (F, TAES). Nail Place,
Young, Aug. 7, 1915 (TEX). CULBERSON couNTY: Guadalupe Mountains, Bailey
739 (US); Havard, in 1881 (US); Silveus 7674 (CAS, DS, SMU, TAES, TEX);
Young, Sept. 5, 1916 (TEX). NE. of Pine Springs, Correll & Johnston 24231 (LL).
McKittrick Canyon, Moore & Steyermark 3636 (CAS, DS, GH, MICH, MO, NY,
UC, US); Warnock 10930 (ILL, LL, SMU), 12089 (LL, SMU). Mountains to-
ward the Rio Grande, Wright 729 (GH). mL paso county: Hueco Tanks, Parks
& Cory 30973 (TAES) ; Whitehouse, Nov. 11, 1931 (TEX). Franklin Mountains,
Warnock 14271 (LL, TEX). El Paso, Whitehouse 8531 (TEX). GILLESPIE
county: feed, January 1890 (MICH). HarpDIN counry: Camp Mountain,
Parks & Cory 7093 (TAES). uupsperuy country: Eagle Mountain, Waterfall
6341 (MO). srr Davis county: Davis Mountains, Brown 4213 (TEX); Young
20a (TEX), 40 (MO, US), Aug. 6, 1914 (TEX), Aug. 10,1914 (GH, TEX). Fort
Davis, Carter 61 (US); Silveus 7675 (CAS, SMU, TAES, TEX). Sawtooth Moun-
tain, Correll 15008 (LL). Halloran, Nov. 29, 1940 (TAES). Mt. Livermore,
Hinckley 101(MICH), July 1986 (ARIZ, GH, NY), Aug. 27,1939 (TEX), Limpia
Canyon, Nealley 133 (DS, GH, ILL, MO, NY, POM, UC, US), in 1890 (US);
Reeder & Reeder 4234 (YU); Silveus 294 (TAES, TEX). Castle Mountain, Reeves
1646 (TAES). Alpine-Ft. Davis, Silveus 824 (US). 15 mi. N. of Alpine, Silveus
825 (TAES, TEX). Mosquiz Canyon-Alpine-Ft. Davis road, Silveus 826 (CAS,
TEX, US). McDonald Observatory, Turner 2970 (SMU). Scenic Loop of Davis
Mountains, Warnock 9285 (LL, SMU). Mosquiz Canyon, Warnock 46557
(TEX). Prestpio county: Chinati Peak, Correll & Johnston 24378 (LL). 26
mi, SE. of Marfa, Cory 2852 (GH). Knox Canyon, Hinckley 2039 (SMU),
July 20, 1941 (TEX). Nevapa: Wheeler, in 1872 (US). cLARK county: Las
Vegas, Arstne, October 1925 (NY). New Mexico: Gray, Goodding M-251.1
(US). Vasey (NY). Wright 1990 (CAS, GH, MO, NY, TAES, US). BERNa-
LILLO COUNTY: Sandia Mountains, Rominger 16 (UNM). caTRoN couNTyY:
Mangas Canyon, Metcalfe, Sept. 15, 1896 (US), November 1896 (US). Tuloros
Creek, Olson 439 (UC). Mogollon Mountains, Rusby 463 (ILL, MICH, MO, NY,
UC, US), 597 (US). pona ana county: Dripping Springs, Archer 498 (MICH),
Dunn 7284 (UNM). Organ Mountains, Hitchcock 3769 (US), 3795 (MICH, US);
Vasey, August 1881 (US); Wooton 461 (DS, MO, NY, US), May 1891 (UNM),
Sept. 17, 1893 (D8, UC, US), August 1894 (UNM), Sept. 17, 1903 (DS); Wooton
& Standley, Sept. 23,1906 (US). Mesilla Valley, Fosberg S-3674 (A, POM). Las

SODERSTROM—PODOSEMUM AND EPICAMPES 169

Cruces, Griffiths 7427 (US). Pefia Blanca, Standley, Oct. 21, 1906 (US), Oct. 28,
1906 (US). Van Pattens Camp, Wooten, Sept. 10, 1899 (ILL). GRANT couNTY:
Mule Creek road, Anderson & Rhinehart 503 (TAES). Fort Bayard Watershed,
Blumer, Nov. 18, 1905 (GH, NY, US). Black Range, Goodding & Goodding 381
(ARIZ). Silver City, Greene 439 (F, US), 440 (US). Rusby, in 1880 (US).
Greenwood Cajion, Smith, Sept. 24, 1896 (US). ‘‘Copper-mines,” Wright 1994
(GH, MO, NY, UC). utpauao county: Clanton Canyon, Castetter 10235
(UNM), 10244 (UNM). Little Hatchet Mountains, Castetter 10387 (UNM).
Skeleton Canyon, Castetter 11256 (UNM). Coronado Forest S. of Animas,
Goodding CG-14 (US). 20 mi. N. of Rodeo, Goodding M-248 (ARIZ, US).
Below Animas Peak, Goodding & Goodding A-9775 (ARIZ, SMU), M-385
(ARIZ, LL). White Water, Mearns 352 (US). Dog Mountains, Mearns 2373
(US). Animas Valley, Mearns 2507 (US). LINCOLN county: 8. of Fort Stanton,
Goodding & Goodding M-183 (MO). oTERO counTy: Guadalupe ridge, Chapline
690 (US). Guadalupe Mountains, Goodding & Goodding M-201 (ARIZ, NY);
Hitchcock 13541 (US holotype of HE. subpatens). Gray’s Ranch, Goodding &
Hardies M-135 (MICH), M-136 (US), M-137 (ARIZ). Queen, Hitchcock 13500
(US), 18579 (US) ; Amer. Gr. Nat. Herb. 13882 (CAS, GH, LL, MO, UC), 1383 (CAS,
GH, LL, MO, UC, US). san miaguet county: Las Vegas, Anect 198 (GH, NY,
US); Arséne 17968 (UC); Drouet & Richards 3224 (F). sIERRA couNTy: Lake
Valley, Beals, in 1914 (MICH, US). Black Range, Metcalfe 1366 (CAS, F, GH,
MO, NY, UC, US). socorro county: Socorro, Plank 53 (US). 12 mi. 8. of
Magdalena, Potter 12 (UNM). Arizona: 3 mi. W. of Palominas, Goodding, Oct.
6, 1935 (ARIZ). Celero Mountains, Griffiths 6122 (MO, US). Stein’s Pass,
Jones 4219 (ARIZ, CAS, F, POM, UC, US). Wood’s Ranch, McDougal 490 (US).
Niggerhead Mountains, Mearns 1871 (US). Mera 1 (UNM). Devils Canyon,
Peebles, Harrison, & Kearney 3200 (ARIZ). Pringle, June 17, 1884 (NY). Mon-
tezuma Wells, Purpus 8289 (MO, UC, US). ‘‘Explorations and surveys W. of the
100th. meridian,’”’ Rothrock 282 (GH syntype of M. vaseyana). “In all canyons,”
Rusby G(MO, NY). White Mountains, Schroeder, summer 1938 (ARIZ). Vasey,
in 1889 (US). cocHIsre county: Chiricahua Mountains, Blumer 68 (ARIZ).
Paradise, Blumer 1698 (ARIZ, DS, F, GH, MO, NY, US). Upper Hunt Canyon,
Clark 8457 (ARIZ). Upper Rhyolite Canyon, Clark 8510 (ARIZ). 8 mi. W. of
Wilcox, Darrow, Sept. 10, 1942 (ARIZ, UC). Carr Canyon, Darrow, Gould, Phil-
lips, & Pultz 1402 (DS), 2436 (ARIZ, CAS, UC, US). Black Canyon, Goodding
429-49 (ARIZ). Coyote Canyon, Goodding 1123-49 (ARIZ). Golden Eagle Box
Canyon, Goodding 88-54 (ARIZ, UC), 30-55 (ARIZ). Huachuca Canyon, Good-
ding 709-58 (ARIZ). Huachuca Mountains, Goodding 251-62 (ARIZ); Jones,
Sept. 3, 1903 (DS, POM). Guadalupe Canyon, Gould & Haskell 4540 (ARIZ, GH).
Dragoon Mountains, Griffiths 1876 (ARIZ, US). Pearce, Griffiths 1957 (ARIZ,
US). Ramsey Canyon, Jones 24736 (CAS, GH, MO, POM, UC), Sept. 30, 1929
(NY). Dos Cabezas, McDougal 778 (US). Pedregosa Mountains, Mearns 838
(US). San Simon, Thornber 2225 (ARIZ). Pine Canyon, Touwmey 14 (US, YU),
Sept. 10, 1896 (NY, US). Blue Canyon, Toumey 15a (US). Fort Huachuca,
Wilcox 410 (US), 446 (TAEHS), in 1894 (NY, US). coconino county: Kaibab
Forest, Pase 343 (US). GILA county: on road from Globe to Coolidge Dam,
Eastward 16646 (CAS). Pocket Creek Canyon, Gould 3741 (ARIZ, GH, NY, UC,
US). Parker Creek Canyon, Gould & Hudson 3867 (ARIZ). Pinal Mountains,
Shreve 7454a (ARIZ). Pine, Towmey 67 (US), July 23, 1892 (ARIZ, DS, UC).
GRAHAM COUNTY: 7 mi. S. of Stockton Pass road, Anderson & Rhinehart 603
(ARIZ). Clifton road, Anderson & Rhinehart 622 (ARIZ, TAKES). Porter
Springs, Kessler 485 (ARIZ, TAES). Graham Mountains, Moeller 186 (ARIZ,
TAES). San Simon Valley, Thornber, Aug. 22, 1905 (ARIZ). MOHAVE COUNTY:

170 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Hualpi Mountains, Nichol, Sept. 14, 19387 (ARIZ). prima county: Redingon
Pass, Benson 9216 (ARIZ, DS, POM). Sabino Canyon, Benson 10559 (POM).
Happy Valley Trail, Blumer 3453 (F). Sabino Trail, Carter, Nov. 20, 1932
(ARIZ). Soldier Canyon, Darrow, Sept. 27, 1936 (GH). Fresnal Canyon, Gilman
11 (ARIZ). Baboquivari Canyon, Gilman 17 (DS, NY, US). Prison Road,
Ginter, Oct. 6, 1941 (UC). Coyote Mountains, Goodding & Lusher 97-45 (ARIZ,
NY). About 25 mi. E. of Tueson, Gould 2565 (ARIZ, MO, UC, US). Santa Rita
Forest Reserve, Griffiths 3437 (US), 5986 (ILL, US). Santa Catalina Mountains,
Griffiths 7043 (US), 7047 (US), 7048 (US), 7149 (ILL, US), 7164 (US); Pringle,
May 14, 1883 (MO, YU); Thornber 9217 (ARIZ). Baboquivari Mountains, Jones
24737 (CAS, GH, MO, POM, TEX, UC), Sept. 19, 1929 (NY). Molina Basin,
Leader & Leader 228 (ARIZ). Redding Pass Road, Leader & Leader 371 (ARIZ).
Baboquivari Canyon, Peebles 532 (ARIZ). Mt. Lemmon, Peebles & Harrison 2293
(MO). Rineon Mountains, Pringle, July 19, 1884 (CAS, F, GH, LL, NY, US).
Soldier Canyon, Shreve 5420 (ARIZ, GH). White House Canyon, Stewart, Sept.
23, 1942 (NY). Soldiers’ Trail, Sfewart, Oct. 10, 1942 (NY). Floritas Canyon,
Streik 8-32 (ARIZ). Rosemont, Thornber, Sept. 30, 1901 (ARIZ, DS, UC).
Tucson, Towmey, in 1892 (NY, UC, US). Reddington Pass, Young, Oct. 2, 1939
(ILL). pinaL county: Oracle, Thornber, Sept. 9, 19(?) (ARIZ). santa cruz
county: Santa Rita Mountains, Arnold & Arnold, Aug. 23, 1939 (UC); Griffiths
7257 (US); Griffiths & Thornber 46 (US); Jones, Aug. 24, 1903 (POM, UC);
Thornber & Griffiths 46 (ARIZ). Box Canyon, Arnold, Sept. 16, 1986 (GH).
White House (Madera) Canyon, Benson 9732 (POM); Graham 3635 (DS); Parker
7079 (ARIZ). Stone Cabin Canyon, Culley 36 (ARIZ); Thornber 188 (ARIZ, DS,
MO, NY, POM, UC), 5245 (ARIZ), Sept. 14, 1903 (UC). Sycamore Canyon,
Goodding M-258 (LL, SMU); Goodding & Goodding 431 (ARIZ); Parker 7381
(ARIZ, ILL). Nogales, Harris, Sept. 19, 1927 (MO, POM); Peebles & Harrison
4695 (ARIZ). Patagonia, Hitchcock 3689 (US), 3719 (US), 3720 (US); Kearney &
Peebles 10050 (MICH, US). Coronado National Forest, Killip 40854 (US).
YAVAPAI coUNTY: Granite Dells, Benham M-95 (ARIZ); Darrow, Sept. 3, 1939
(ARIZ, UC). Mingus Mountain, Benham M-109 (MICH, YU); Goodding 174-46
(ARIZ, NY). Fort Whipple, Cowes & Palmer 424 (MO). 10 mi. 8. of Prescott,
Featherly, July 18, 1929 (SMU). Prescott, Kearney & Peebles 9735 (ARIZ, US);
Purchase 527 (MICH); Thornber 8644 (ARIZ); Williams 3037 (US).

MEXICO: Palmer, in 1886 (ARIZ). Wilkinson 9601 (F, NY, US). Basa
Cauirornia: El Taste, Brandegee, Sept. 13, 1898 (NY, UC, US), Nov. 1902
(UC). Sierra de San Francisquito, Brandegee, Sept. 24, 1899 (US), Sept. 29,
1899 (UC). Sierra de la Laguna, Brandegee, Oct. 4, 1899 (UC,US). Cerro de la
Giganta, Carter, Alexander, & Kellogg 2025 (DS, GH, LL, MO, UC, US). Sierra
de la Laguna, E. of Todos Santos, Carter, Alerander, & Kellogg 2882 (DS, GH,
UC, US). Sonora: Cajion Estrella, Gentry 394 M (DS, MICH), Oct. 1, 1933
(US). Cerro del Capulin, NW. of Aribabi, Harvey 1688 (CAS, GH, MICH, MO,
US). Estaci6n de Tajo, Herndndez & Tapia N-1569 (CHAP). San José
Mountains, Mearns 962 (US). Sierra de los Alamos, Palmer 409, in 1890 (CAS,
GH, US). Colonia Morelos, Santos 30 (US) 174 (US). 20 mi. W. of La
Angostura, Santos 1798 (GH). El Tigre to El Rancho de Robles, Santos 1925
(ARIZ, GH, MICH, MO, NY, SMU, TEX, US). El Rancho de Robles to
Colonia Morelos, Santos 1974 (ARIZ). El Puerto del Cumarito, Santos 2090
(GH, SMU, TEX). El Cafion de la Bellota, Santos 2109 (DS, SMU, TEX).
El Picacho del Pilar, Santos 2166 (GH, SMU, TEX). Canon del Agua Amarga,
White 3631 (MICH). 8 mi. §. of Nogales, Sonora, Wiggins 7535 (DS, MICH,
US). Curauanua: Arroyo Mestefio, Sierra del Nido, Christman 170 (UC).
15 mi. S. of Encinillas, Correll 23294 (LL). Minas Nuevas, Correll & Gentry

SODERSTROM—PODOSEMUM AND EPICAMPES 171

22736 (LL). Between Guadalupe y Calvo and Tecolote, Correll & Gentry 23237
(LL). Guasaremos, Gentry 1874 (ARIZ, F, GH, MEXU, MO, UC). Rancho
Carretas, Harvey 1627 (CAS, GH, MICH, MO, US), 1641 (MICH). Km. 1773
carretera Cd. Judrez, Hernandez & Mathus N-1837 (GH). Sanchez, Hitchcock
7669 (US), 7669 % (NY, US), 7714 (ILL, US). Mifiaca, Hitchcock 7742 (US).
Pirdmide, Johnston 8118 (GH). Guayanopa Canyon, Jones, Sept. 24, 1903
(NY, POM). Rfo Gavildén, Leopold 225 (US). Majalca, Le Sueur Mex-021 (GH,
MO, SMU, TEX, US). Chuhuichupa, Le Sueur Mex-087 (TEX). NW. of
Chihuahua, Le Sueur 0149 (CAS, GH, MO, SMU, UC, US). 20 mi. SW. of
Chuhuichupa, Muller 3594 (GH, LL, MICH, UC). Sierra Gazachic, SW. of
Mifiaca, Pennell 18957 (US). 15 mi. S. of Madera, Reeder & Reeder 2640 (YU).
23 mi. SE. of Chuhuichupa, Reeder & Reeder 2670 (YU), 2671 (YU). 37 mi. 5.
of Villa Matamoros, Reeder, Reeder, & Soderstrom 3449 (US, YU). 4 mi. E. of
Parral, Reeder, Reeder, & Soderstrom 3457 (US, YU). 1 mi. E. of Cuauhtémoc,
Reeder, Reeder, & Soderstrom 3467 (US, YU). 16 mi. W. of Cuauhtémoc, Reeder,
Reeder, & Soderstrom 3469 (US, YU), 3470 (US, YU), 3471 (US, YU), 3472 (US,
YU). 12 mi. N. of Chihuahua, Reeder, Reeder, & Soderstrom 3482 (US, YU),
3485 (US, YU). 15 mi. W. of Flores Magon, Reeder, Reeder, & Soderstrom 3499
(US, YU). 12 mi. W. of Colonia Judrez, Reeder, Reeder, & Soderstrom 3502
(US, YU), 3503 (US, YU). S. of Hernandez, Reeder, Reeder, & Soderstrom 3524
(US, YU), 3533 (US, YU) NE. of Cuesta Blanca, Reeder, Reeder, & Soderstrom
3542 (US, YU), 3543 (US, YU), 3548 (US, YU), 3549 (US, YU). Parral,
Soderstrom 828 (US, YU). San Francisco de Oro, Soderstrom 870 (US, YU),
871 (US, YU). 13 mi. SW. of Cuauhtémoe, Soderstrom 917 (US, YU), 918 (US,
YU). 12 mi. N. of Chihuahua, Soderstrom 930 (US, YU). Sierra Hechiceros
near the Coahuilan border, Stewart 202 (GH). Organos, Stewart & Johnston
2066 (GH). Casas Grandes, Townsend & Barber 357 (CAS, F, GH, ILL, MO,
NY, POM, UC, US). Santa Eulalia Tills, Wilkinson, Aug. 25, 1885 (F, US),
Sept. 21, 1885 (UC). Coanvuita: Corte Blanco fork of Charretera Canyon,
Johnston 8950 (GH, MO). 2 mi. NE. of Nava, Johnston & Graham 4166 (TEX).
La Noria, Johnston & Muller 462 (GH, LL), 587 (GH, LL). Del Carmen
Mountains, Marsh 624 (GH, TEX, US), 655 (GH, TEX, US). Santa Rosa
Mountains, Marsh 1511 (F, GH, TEX). Sierra de la Gloria, Marsh 1948 (F,
GH, TEX). 1 mi. S. of Cuesta Blanca, Reeder & Reeder 3227 (YU). 26 mi. E.
of Saltillo, Reeder, Reeder, & Soderstrom 3289 (US, YU). Jaral, Schumann 1753
(US). La Noria, Stewart 1208 (GH). Sierra del Carmen, Villa Acufia, Wynd &
Mueller 594 (US). Nurvo Leon: Dulees Nombres, Meyer & Rogers 3092 (MO,
US). Monterrey, Mueller & Mueller 385 (GH). Ca. 10 mi. E. of Galeana,
Reeder & Reeder 3973 (US, YU). Mt. Mirador, Roybal 60 (US). 8 mi. N. of
Escondido, Shreve & Tinkham 9714a (ARIZ). TaMAvLipas: (?)El Rosario,
Bartlett 10882 (CAS, GH, MICH, NY, POM). 11 mi. W. of Victoria toward
Jaumave, Johnston & Graham 4141 K (TEX). (?)Sierra de Tamaulipas, at Santa
Marfa de los Nogales, Martinez & Borja F-1939, aff. M. emersleyi (TEX), F-2083,
aff. M. emersleyi (TEX). Srnatoa: Cerro Babuyo, Gentry 11420 (US). Ortega
3118 (US). 18 mi. E. of Santa Lucfa, Reeder & Reeder 2469 (YU). DURANGO:
40 mi. S. of Durango, Herndndez, Ruppert, & Guevara X-2558 (US). Durango,
Hitchcock 7621 (US); Palmer 542, in 1896 (CAS, F, GH, MO, UC, US). Otinapa,
Palmer 344, in 1906 (F, GH, NY, US), 553, in 1906 (US). 25 mi. W. of Durango,
Reeder, Reeder, & Soderstrom 3440 (US, YU), 3441 (US, YU). 27 mi. SE. of
Durango, Soderstrom 756 (US), 758 (US). SW. of Durango on Durango- Mazatlan
road, Soderstrom 767 (US), 772 (US), 775 (US). 6 mi. N. of Durango, Soderstrom
787 (US), 789 (US). Zacatecas: 6 mi. SW. of Carboneras, Genéry 8522. 15 mi.
W. of Zacatecas, Soderstrom 725 (US). 25 mi. NW. of Fresnillo, Soderstrom 728

172 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

(US), 729 (US). Sombrerete, Soderstrom 734 (US), 735 (US), 746 (US), 753
(US). San Luis Porosf: 8 mi. W. of Santa Catarina, Reeder & Reeder 4104,
SW. of San Luis Potosi, Reeder, Reeder, & Soderstrom 3303 (US, YU), 3324 (US,
YU), 3325 (US, YU), 3326 (US, YU), 3330 (US, YU). Sierra de San Miguelito,
Sohns 1140 (US), 1151 (US), 1157 (US), 1820 (US), 1465 (US). Sierra de Alvarez,
Sohns 1385 (US). A@uascaLIENTES: Rineédn de Romos, Reeder, Reeder, &
Soderstrom 3335 (US). Jatisco: Bolafios, Rose 3003 (US). 15 mi. SH. of
Inearnacién de Diaz, Soderstrom 694 (US), 697 (US). GuaNnasuato: Ojo de
Agua, Sohns 252 (MICH, US). 6 km. E. of Guanajuato, Sohns 315 (MICH,
US). Hipatco: 18 mi. NE. of Zimapdn, Reeder & Reeder 4121 (US, YU).
Pursta: Cerro de Soluche, Purpus 4081 (F, GH, MO, NY, UC, US). Moretos:
Km. 53 Autopista México-Cuernavaca, Espinosa 66 (IPN). Mexico: Naranjo,
Temasealtepec, Hinton 1987 (NY, UC). Distriro FrpErAu: Mixcoae, Sté.
Pierre 905 (MICH, US). Micnoacdn: La Huerta, Arséne, Oct. 30, 1909 (NY).
San Miguel, Arsene, Dec. 4, 1910 (NY). Oaxaca: Huahuapan de Leén,
Soderstrom 413 (US), 414 (US), 447 (US).

Muhlenbergia emersleyt is distinguished by the typically long mem-
branous ligule, lemmas pubescent on the midnerve and margins,
palea pubescent between the keels, and the purplish, flexuous, more
or less pyramid-shaped panicle. It appears to be most closely re-
lated to M. lekmanniana with their similar overall habit, long ligules,
and lemmas pubescent on the lateral margins. Muhlenbergia leh-
manniana differs in that the midnerve of the lemma is usually gla-
brous or if pubescent the hairs are sparse and rather inconspicuous;
the palea is glabrous between the keels, rarely a few hairs are present.
The panicles of M. lehmanniana are golden yellow but usually pur-
plish in M. emersleyi. The callus is blunt or truncate at the base
in M. emersleyi but more or less acute in M. lehmanniana. Muhlen-
bergia emersleyt is most common in southwestern United States and
northern Mexico and has not been collected south of the Isthmus of
Tehuantepec. South of the Isthmus, in Chiapas through Central
America, the specimens with yellowish panicles are referable to MM.
lehmanniana. Comparison may also be made with M. breviligula,
its ligule only a short rim or almost lacking. However, long ligules
are sometimes found on the leaves of the innovations in the same
plants that have the typically short ligule on the culm leaves. This
suggests, perhaps, the ancestral nature of a long ligule in this species
and points to a close relationship with M. emersleyi.

The awned condition of the lemmas requires further study. Awned
and awnless specimens, alike in all other respects, were collected side
by side (e.g., Pennell 18957, Greene 439 and 440, awnless and awned
forms; and Nealley 133, slightly awned and very long-awned speci-
mens). In this species the condition of the awn is not correlated
with other characters to separate the awned from the awnless plants.
The length of the awn is also quite variable. The author has collected
plants side by side; in one the awns were short (/eeder, Reeder, &

SODERSTROM—PODOSEMUM AND EPICAMPES 173

Soderstrom 3548) and in the other very long (Reeder, Reeder, & Soder-
strom 3543). All of the specimens I have seen from Baja California
and Sonora, Mexico, are long-awned.

An awnless form of this species from New Mexico (Hitchcock
13541) served as the type for Epicampes subpatens Hitche. In the
type specimen the spikelets are about 3 mm. long, the glumes whitish
and scabrous, and the lemmas densely pubescent on the midnerve
and margins. Although most of the spikelets are awnless, some
bear a rudimentary awn from just below the tip. The branches of
the panicle are stiffer than those of the type of M. emersleyi (Emersley
46) and the panicle is not lax. With this exception no basic differ-
ence exists between the two specimens. Hitchcock, who described
E. subpatens, later placed it in synonymy with M. emersleyi.

Three specimens deserve special comment. Bartlett 10882 (Sierra
de San Carlos, Tamaulipas, Mexico) has an inflorescence much like
that of M. emersleyi with long-awned spikelets in which the lemmas
have the typical pubescence pattern of the species. However, the
basal sheaths in many specimens are not strongly compressed-keeled,
the ligule is firm below and membranous above, and the blades are
narrow and folded, reminiscent in these latter features of M. gooddingii.
Although it looks distinctive, no basic characters separate this
specimen from the M/Z. emersleyt complex. Two additional specimens,
Martinez & Borja F-1939 and F-2083 (Sierra de Tamaulipas),
are quite distinctive, but I can find no good characters to separate
them from the M. emersleyi complex. Further collections of Epicampes
from the state of Tamaulipas may clarify the position of these unusual
specimens. I have seen only four specimens from the whole state.

Muhlenbergia emersleyi is the most common species of Epicampes
and the most variable. Attempts to segregate the forms into more
than one species on the basis of morphology are unrewarding because
of a thorough intergradation of characters among the specimens.
An intensive cytological investigation of this species complex will be
necessary before we can be certain of the precise nature and extent
of the various races present.

25. Muhlenbergia xanthodas Soderstrom, sp. nov.

Perennis caespitosa, culmis robustis erectis glabris 60-90 cm.
altis, nodis 3 glabris, vaginis inferioribus compressis et robuste
carinatis basin versus glabris adversum collum scaberulis; ligula
membranacea (ad basin firma), margine versus apicem lacerata, usque
ad 30 mm. longa; laminis culmi angustis attenuatisque conduplicatis
utrinque scabris, ca. 70 cm. longis; collis scabris; panicula erecta
angusta aurea, 23-27 cm. longa, usque ad 3 em. lata, ramis appressis
ad basin florescentibus, axe glabro, ramis primariis usque ad 4.5 cm.

174 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

longis, secundariis usque ad 1.5 cm. longis, tertiariis usque ad 5 mm.
longis, pedicellis glabris infra spiculam scaberulis quam_ spiculis
brevioribus; spiculis 2-3 (plerumque 2-2.5) mm. longis, glumis
acutissimis fere aequalibus, quam flosculo paullo longioribus, glabris
lucidis paene translucentibus enerviis, lemmatibus acutis, 2-2.7 mm.
longis, glabris nisi quod puberulentis lateribus ad basin, nervis 3,
infra apicem aristas, 6-10 mm. longas, aureas ferentibus, callo brevi-
piloso,

A M. lehmanniana Henr. ramis paniculae brevioribus et magis
appressis, et glumis glabris lucidis (paene translucentibus) sine
nervis differt.

Caespitose perennial; culms stout, erect, glabrous, 60-90 cm. tall;
nodes 3, glabrous beneath; basal sheaths strongly compressed-keeled,
elabrous below, becoming scaberulous toward the collar; ligule mem-
branous (somewhat firm at the base), lacerate above, as short as
0.5 mm. on the lower culm leaves, to as much as 30 mm. on the higher
culm leaves; sheath auricles lacking; blades of the culm to 70 em.
long, narrow and long-attenuate, tightly folded, 1.5-2 mm. wide
when unfolded, the upper and lower surfaces scabrous; upper throat
scabrous; collar scaberulous.

Panicle erect, narrow, golden yellow, 23-27 cm. long, to 3 cm.
wide, the branches of the panicle strongly appressed, floriferous
almost to the base; axis glabrous; primary branches to 4.5 cm. long,
the secondary branches to 1.5 cm. long, the tertiary branches to 5
mm. long; pedicels shorter than the spikelet, glabrous, scaberulous
beneath the spikelet; spikelets 2-3 (mostly 2-2.5) mm. long; glumes
acute, about equal to each other, a little longer than the floret, gla-
brous, shining, almost translucent, nerveless; lemmas acute, 2~-2.7
mm. long, glabrous except for short hairs toward the base of the
lateral margins, 3-nerved, the middle nerve extending into a golden
awn from below the tip of the lemma, 6-10 mm. long; callus of lemma
short-pilose; palea equal to or a little longer than the lemma, glabrous.

Holotype in the U.S. National Herbarium, no. 1817864, collected
on rock on Mt. Ovando, Chiapas, Mexico, at 2,300 meters altitude,
November 14-18, 1939, by Eizi Matuda (no. 4003). Isotypes at F,
MICH.

DistRIBUTION: Known only from Chiapas, Mexico, from the type
and from the following additional collection:

MEXICO: Cutapas: Between Escuipulas and Cafiada Honda, Hernandez &
Sharp X-311 (US).

Muhlenbergia zanthodas appears to be most closely related to A.
lehkmanniana which ranges from Costa Rica to Colombia. Both species
are like AZ. emersleyi in general habit and both also have long-
awned lemmas, pubescence on the margins of the lemma at the base,

SODERSTROM—PODOSEMUM AND EPICAMPES 175

and long membranous ligules. In M. emersleyi the lemma is pubescent
on the midnerve and the margins, the panicles and awns of the lemma
are generally purplish. In M. zanthodas and M. lehmanniana the
lemma is short-pilose at the base, but the midnerve of the lemma is
glabrous or bears relatively few inconspicuous hairs, and the panicles
are golden, the rather straight awns of the lemmas are golden, and the
strongly compressed-keeled sheaths are golden. In typical M.
emersleyi the paleas are pubescent between the keels, but in M. zan-
thodas and M. lehmanniana the paleas are glabrous. Muhlenbergia
zanthodas is distinct from M. lehmanniana in its much shorter, closely
appressed panicle branches, producing a very dense panicle; in M.
lehmanniana the branches are longer and spreading. The glumes of
M. zanthodas are glabrous, shining (almost translucent), and nerveless;
the glumes of M. lehmanniana are scabrous and 1-nerved.

The specific epithet is from the Greek words tav@és, yellow or
golden, and éés, torch, in allusion to the dense golden panicles.

26. Muhlenbergia lehmanniana Henr. Med. Rijksherb. Leiden 40:59. 1921.

M. multinodis Asplund. Bot. Not. 1939: 796. f. 1. 1939.
M. attenuata Swallen. Ann. Missouri Bot. Gard. 30:138. 1943.

Caespitose perennial; culms glabrous to scaberulous, 72-167 cm.
tall; nodes 3-4, glabrous beneath; basal sheaths compressed-keeled,
glabrous, tightly imbricate, flat, golden yellow, not becoming brown
and shredded; ligule somewhat firm at the base, becoming long and
membranous, conspicuous from side view, slightly decurrent, 1 cm.
long, to as long as 2.5 cm., enclosed in the folded blade above, entire
below and for the greater part of its length; sheath auricles lacking
or rarely rudimentary ones present, no more than 0.1 mm. long;
blades of the culm folded, long and narrow, becoming threadlike
toward the tip, 1.5-4 mm. wide when flat, the upper surface of the
blade slightly scaberulous to scabrous, the lower surface scaberulous
to scabrous; upper throat scabrous; collar scaberulous to scabrous.

Panicle erect, usually dense, the branches ascending 19-40 cm.
long, 3-6 cm. wide, generally golden, golden brown, or purplish
ereen; axis scaberulous to scabrous; the primary branches 3.5-12
(rarely to 19) cm. long, the secondary branches 1-5 (mostly 1-2) cm.
long, tertiary branches usually lacking (rarely to 1.5 cm. long);
pedicels of the lateral spikelets 0.7-2 mm. long, those of the terminal
spikelets relatively long, 2-4 mm., scaberulous to scabrous, especially
just beneath the spikelet; spikelets 2-3.5 (usually about 2.5) mm.
long; glumes longer than the floret, the second glume usually a little
longer than the first, mostly scabrous, blunt at the tip or slightly
dentate (under high power), usually slightly awn-tipped, the awns to
0.3 mm., hyaline, l-nerved (1 or 2 additional faint nerves sometimes

176 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

present) ; lemmas 1.8-3 (mostly about 2.5) mm. long, glabrous dorsally
except at the base of the midvein, the lower part of the margins
conspicuously short-pilose, acutish at tip, 3-nerved, the awns straight,
or somewhat flexuous, over 1 (to 3.5) cm. long; callus of the lemma
short-pilose; palea a little shorter than the lemma, glabrous or at
most with a few hairs at the base between the keels.

Holotype in the Rijksherbarium, Leiden, collected near Popaydn,
Cauca, Colombia, by F. C. Lehmann, no. B. T. 1267 (in Herb. Lugd.
Bat. under no. 908. 350-519). Fragments of type ex K in the U:S.
National Herbarium, no. 72978, ex L, no. 72977, and ex. Herb. Lugd.
Bat. (L), no. 72979. Isotype at NY.

Disrripution: Hillsides and barrancas at elevations of mostly
between 2,500 and 6,000 ft. One label states “from 100 m.,” another
“10,000 ft.,”’ but I doubt if the species is found at these extremes.
Rather common from Costa Rica, south and eastward into Colombia.

COSTA RICA: Tonduz 4630 (US). Las Céncavas, Lankester 676 (US)
Oersted 14065 (F, US), 14171 (US). Bord du chemin de Mano de Tigre, Pitlier
4630 (US). Buenos Aires, Canton de Osa, Valerio 873 (F). PuNTARENAS:
Savanes de Cafias Gordas, Pittier 7355 (US), 7358 (US), 11019 (US). AtasuEwa:
San Pedro de San Ramén (cerea del Rio Barranca), Brenes 21415 (F, NY). San
Joss: Tablazo hasta 1800 m., Valerio 1052 (F). Carraco: Tejar, Solis 618
(44) (F). Mount Irazu, Oersted 14012 (US).

PANAMA: CurriQqvuf: El Boquete, Hitchcock 8174 (US holotype of M. attenuata).
Grassy hillsides, Hitchcock 8241 (US). Foothills, Hitchcock 8242 (US). Barranca,
road toward Dolega, Hitchcock 8340 (US). Cerro Vaca, Pittier 5360 (US), 5362
(US). Cocni: 8. of El Valle de Antén, Allen 2812 (GH, MO, US). Vicinity of
Old, Pittter 5042 (US), Pittier 5062 (US).

COLOMBIA: Antioquta: Medellin, Bro. Daniel 246 (US). Barbosa, vicinity
of Medellin, Toro 697 (US). Vatue: Cordillera Occidental, Hoya del Rio Cali,
Cuatrecasas 18737 (F, US). Cauca: Popaydn, Lehmann B.T. 1267 (US fragments
of type ex K and L, NY isotype). Loma de Bichiquf near Toribié, Pittier 1468
(US). El Tambo, von Sneidern 1323 (S holotype of M. multinodis), A. 231 (A).

Muhlenbergia lehmanniana is distinguished by the long ligule,
pyramidal panicles of spikelets with scabrous glumes, long-awned
lemmas pubescent only at the base, paleas glabrous between the
keels, and calluses acute rather than truncate.

This species looks much like M. emersleyi and has been confused
with it. The panicles of long-awned spikelets resemble some forms of
M. emersleyx. However, a number of characters differentiate the
two species. Muhlenbergia lehmanniana occurs in the mountains
of Costa Rica through Panama and into the Colombian Andes;
M. emersleyi is most abundant in southwestern United States and
northern Mexico, becoming less frequent southward into Mexico
until, beyond the Isthmus of Tehuantepec, it disappears. The basal
sheaths of MZ. lehmanniana are distinctly flabellate and golden,

SODERSTROM—PODOSEMUM AND EPICAMPES 177

more so than M. emersleyi, and the blades are long, narrow, and
attenuate, rather threadlike at the ends. The acute callus of the
floret distinguishes this species from MM. emersleyt. The hairs on
the callus and lower part of the margins of the lemma and midnerve
(where confined) are longer individually than in M. emersleyi. In
that species the hairs usually extend about two-thirds up the lemma.
In most specimens of M/. lehmanniana the palea is glabrous but
pubescent on a half to two-thirds of its length in M. emersleyi.

Swallen’s M. attenuata, although distinctive because of the un-
usually long panicle branches long-naked at the base, is evidently
an extreme form of this species, and the type of M4. multinodis unques-
tionably fits within the morphological range of M. lekmanniana.

Muhlenbergia lehmanniana is closely related to M. xanthodas which
differs in the nerveless, glabrous, almost translucent glumes, and
short, appressed panicle branches. The ranges of both species are
well separated, Muhlenbergia xanthodas being known only from
Chiapas.

Species Described as Epicampes but Excluded from Section Epicampes

Epicampes argentea (Vasey) Jones. Contr. West. Bot. 14:7. 1912.=Muhlen-
bergia argentea Vasey. Bull. Torrey Club. 13:232. 1886. A weak, some-
what decumbent perennial with rounded basal sheaths [SW. Chihuahua, Mexico].

Epicampes arundinacea (Trin. [error for Griseb.]) Hack. Anal. Mus. Nac. Buenos
Aires 11:107. 1904.=Sporobolus rigens (Trin.) Desv. in C. Gay. Fl. Chile
6:295. 1853. [Chile.]

Epicampes buchingert Fourn. Mex. Pl. 2:88. 1886.=An annual Muhlenbergia
(from description; type not seen). [Mexico.]

Epicampes clomena (Trin. [error for Kunth]) Jones. Contr. West. Bot. 14:7.
1912.=Muhlenbergia peruviana (Beauv.) Steud. Nom. Bot. ed. 2. I[:41.
1840. An annual [Peru].

Epicampes coerulea Griseb. Abh. Ges. Wiss. Gottingen 19:256. 1874=?Muhlen-
bergia angustata (Presl) Kunth. Rev. Gram. I. (Supp.): xvi. 1830. [Colombia
to Argentina.]

Epicampes coerulea var. submutica Hack. ex Stuck. Ann. Mus. Nac. Buenos Aires
13:471. 1906.=?Muhlenbergia angustata (Presl) Kunth. 1830. Rev.
Gram. I (Supp.): xvi. [Colombia to Argentina.]

Epicampes crassiculmis Piper. Proc. Biol. Soc. Wash. 18:144. 1905.=Sporo-
bolus wrightii Munro ex Scribn. Bull. Torrey Club 9:103. 1882. [USA,
Mexico. ]

Epicampes kunthiana Griseb. Abh. Ges. Wiss. Géttingen 19:256. 1874.=Muh-
lenbergia angustata (Presl) Kunth. Rev. Gram. I (Supp.): xvi. 1830.
[Colombia to Argentina.]

Epicampes macroura (H.B.K.) Benth. Journ. Linn. Soc. Bot. 87. 1881.
= Muhlenbergia macroura (I].B.K.) Hitche. A member of section Podose-
mum. [Mexico and Guatemala. ]

Epicampes mexicana (Seribn.) Jones. Contr. West. Bot. 14:7. 1912.
= Muhlenbergia biloba Hitche. Contr. U.S. Nat. Herb. 17:294. 1913. An
annual [Mexico].

178 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Epicampes phleoides (H.B.K.) Griseb. Abh. Ges. Wiss. Gottingen 19:256.
1874.=Muhlenbergia angustata (Presl) Kunth. 1830. Rev. Gram. I
(Supp.): xvi. [Colombia to Argentina.]

Epicampes rigens Benth. in Journ. Linn. Soc. Bot. 19:87. 1881.=Muhlen-
bergia rigens (Benth.) Hitche. A member of section Podosemum. [SW.
USA and Mexico.]

Epicampes rigens (Trin.) Phil. Anal. Mus. Nac. Chile sect. II(8):82. 1891.
Based on Vilfa rigens Trin. which=Sporobolus rigens (Trin.) Desv. in C.
Gay. Pl. Chile 6:295. 1853. [Chile.] Not Epicampes rigens Benth.
Journ. Linn. Soc. 19:87, 1881. [USA, Mexico.]

New Species and New Combination

Muhlenbergia gooddingiit Soderstrom, sp. nov.
Muhlenbergia inaequalis Soderstrom, sp. nov.
Muhlenbergia tridifolia Soderstrom, sp. nov.
Muhlenbergia reederorum Soderstrom, sp. nov.
Muhlenbergia virletit (Fourn.) Soderstrom, comb. nov.
Muhlenbergia xanthodas Soderstrom, sp. nov.

Voucher Specimens Examined

Specimens used in the study of leaf anatomy, embryos, and lodi-
cules are listed below. The letters in the herbarium citation in-
dicate the study for which the specimen was used (A=leaf anatomy:
cross section and/or epidermis, E=embryo, and L=lodicule). The
author was fortunate to have access to the John R. Reeder collection,
indicated by (R), of slides and drawings of many grasses.

MUBLENBERGIA

M. andina (Nutt.) Hitehe. USA: Arizona: Grand Canyon Natl. Park, North
Rim, along Point Sublime Road, Reeder 1808 (YU): L.

M. angustata (Presl) Kunth. ECUADOR: ‘anar, near the village of San
Marcos, Camp E-2519 (NY): A, L.

M. arentcola Buckl. MEXICO: Curinuanua: Plains near Chihuahua, Pringle
479 (NY): A, L.

M. argentea Vasey. MEXICO: Cutnuanua, Palmer 160, in 1885 (US holo-
type): A.

M. arizonica Scribn. MEXICO: Duranao: N. of Cd. Durango, Reeder 2602
(YU): A(R).

M. articulata Scribn. MEXICO: San Luis Porosf: Cardenas, Pringle 3913
(NY): A, L.

M. asperifolia (Nees & Mey.) Parodi. USA: New Mexico: Albuquerque,
Hardies M-103 (YU): A(R).

M. biloba Hitche. MEXICO: Cuinuanua: Near Chihuahua, Pringle 819
(NY): IL.

M. breviliqula Hiteche. GUATEMALA: Janapa: Vicinity of Jalapa, Standley
76731 (F): A.

M. brevis C. O. Goodding. MEXICO: Cutnuanvua: 12 miles W. of Cuauhté-
moc, Reeder 2620 (YU): L(R).

SODERSTROM—-PODOSEMUM AND EPICAMPES 179

M. brevivaginata Swallen. MEXICO: Jauisco: 6 mi. NW. of Cuautla, Me Vaugh
13671 (US holotype): A.

M. capillaris (Lam.) Trin. USA: Fuoripa: Near Highland View, Kral &
Godfrey 54211(NY): L. Texas: Bastrop State Park, Silveus 2396 (YU): A(R).

M. confusa (Fourn) Swallen. MEXICO: Cuinuanua: 10 miles N. of Largo,
Reeder 2684 (YU): A(R).

M. crispiseta Hitche. MEXICO: Duranao: 23 miles W. of Cd. Durango,
Reeder 2545 (YU): A(R).

M. curvula Swallen. MEXICO: Guawnasuato: 12 mi. from Guanajuato on
road to Santa Rosa, Moore 1353 (US holotype): A.

M. depauperata Scribn. MEXICO: Guanasvaro: 11 miles 8. of San Miguel
Allende, Reeder 2279 (YU): L(R).

M. distans Swallen. MEXICO: Disrrrro FepEerAt: Lomas, Lyonnet 55 (NY):
A. Oaxaca: Las Sedas, Pringle 5574 (NY): A, L.

M. distichophylla (Presl) Kunth. MEXICO: Jauisco: Near Guadalajara,
Pringle 2346 (NY): A, L. Gurrrero: Vallecitos, Montes de Oca, Hinton
11635 (NY): A.

M. dubia Fourn. MEXICO: Mexico: 12 miles E. of Ixtlahuaca, Soderstrom
509 (YU): A, L.

M. dubioides C. O. Goodding. MEXICO: Cuinvanua: 12 miles N. of Chi-
huahua, Soderstrom 931 (YU): A.

M. elongata Scribn. MEXICO: Cutnuanva: Near Chihuahua, Pringle 938
(NY): A.

M. emersleyi Vasey. MEXICO: Zacatecas: 13 miles W. of Sombrerete,
Soderstrom 746 (YU): A, L.

M. erectifolia Swallen. COLOMBIA: Santanper: Péramo de Santurbdn,
near Vetas, Killip & Smith 17470 (US holotype): A.

M. eriophylla Swallen. MEXICO: Mexico: Luvianos, Temascaltepec, Hinton
5307 (US holotype): A.

M. expansa (DC.) Trin. USA: SourH Caroutna: 5 miles 8. of Georgetown,
Godfrey 8151 (NY): A, L. Norra Carouina: Wilmington, Amer. Gr. Nat.
Herb, 1374 (NY): A, E.

M. filiculmis Vasey. USA: Cotorapo: Green Mountain Falls, Sheldon 321
(US holotype): A.

M. firma Beal. MEXICO: Hinauao: Pachuca, Amer. Gr. Nat. Herb. 1351 (NY):
A, L.

M. flexuosa Witche. PERU: Huacachi, estacién near Muna, Macbride 3874
(US holotype): A.

M. frondosa (Poir.) Fernald. USA: Connecticut: New Haven (no voucher): L.

M. gigantea (Fourn.) Hitche. MEXICO: Duranao: 22 miles E. of Santa Lucfa,
Reeder 2488 (YU): A, L. Janisco: Near Guadalajara, Pringle 3335 (NY):
A, E; 8 miles W. of Guadalajara, Soderstrom 607 (YU): A.

M. glabrata (H. B. K.) Kunth. MEXICO: Pursia: San Marcos, Amer. Gr. Nat.
Herb. 1356 (NY): A, L.

M. glomerata (Willd.) Trin. USA: Connecticut: 6 miles W. of Tolland, Reeder
944 (YU): L.

M. grandis Vasey. MEXICO: Jauisco: Near Guadalajara, Pringle 2765 (NY):
A, E, L.

M. hintonit Swallen, MEXICO: Mexico: Crucero, Temascaltepec, Hinton 3059
(US holotype): A.

M. holwayorum Hitche. BOLIVIA: Sorata, Rusby 201 (NY): A.

M. involuta Swallen. USA: Texas: Boerne-Bandera Road, N. of San Antonio,
Silveus 780 (NY): A(R), L.

221-352—67——-8

180 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

M. jaliscana Swallen. MEXICO: Jaxisco: Sierra de Manantlin (15-20 mi. SE.
of Autldn), McVaugh 13991 (US holotype): A.

M. lindheimeri Hiteche. USA: Texas: 15 miles 8. of San Antonio, Burr 513 (NY):
A, L.

M. longiglumis Vasey. MEXICO: Jarisco: Pringle 2365 (NY): A, L.

M. longiligula Hitche. USA: Arizona: Chiricahua Mts., Barfoot Park, Blumer
1424 (NY): A. MEXICO: Cumuanua: 38 miles 8. of Casas Grandes, Reeder
2699 (YU): A, E; 31 miles W. of Colonia Judrez, Reeder 3222 (YU): L.

M. lucida Swallen. MEXICO: Curmvanva: Sierra Gazachic 35 km. SW. of
Miniaca, Pennell 18955 (US holotype): A.

M. macrotis (Piper) Hitche. MEXICO: Janisco: Pringle 2360 (NY): KE.
Guerrero: Petlacala, S. of Reyes, Mexia 9098 (NY): A, L.

M. macroura (H. B. K.) Hitche. MEXICO: MicnoacAn: Quinceo, near Morelia,
Arsene, Nov. 11, 1909 (F): KE; 14 miles E. of Morelia, Soderstrom 551 (US): L.
Curapas: Chiquihuite, Vol. Tacana, Matuda 2829 (NY): A.

M. matudae Sohns. MEXICO: Moretos: Lago de Zempoala, Matuda 25601
(US holotype): A.

M. metcalfei Jones. USA: New Mexico: Grant County, Santa Rita Mountains,
Metcalfe 1485; (US holotype): A.

M. mexicana forma ambigua (Torr.) Fernald. USA: Connecricut: New Haven
(no voucher): L.

M. montana (Nutt.) Hitche. MEXICO: Sonora: Trail from El Tigre to El
Rancho de Robles, Santos 1924 (NY): A. Muicnoachn: Morelia, Arsene 5811
(NY): L.

M. aff. montana (Nutt.) Hiteche. MEXICO: Duranao: 7 miles E. of El Salto,
Reeder 2572 (YU): A(R).

M. nigra Hitchc. MEXICO: Disrriro Feperat: Serrania de Ajusco, Pringle
6676 (NY): A. Near Cima, Pringle 11739 (NY): L. GUATEMALA:
Voledn de Agua, Cook, May 12, 1905 (NY): E.

M. orophila Swallen. GUATEMALA: Hvgnverenanco: Cerro Chemal,
summit of Sierra de los Cuchumatanes, Steyermark 50309 (US, holotype): A.

M. palmeri Vasey. MEXICO: Cuinuanua: Sierra Madre, Pringle 1417 (NY):
A, L.

M. peruviana (Beauv.) Steud. PERU: Arxrquira: July 11, 1946, Pearson 1:
(YU): A(R).

M. plumbea Scribn. in Beal. MEXICO: Mexico: 17 miles E. of Toluca, Reeder
2218 (YU): A(R).

M. pubescens (H. B. K.) Hitche. MEXICO: Jatisco: 15 miles SE. of Enear-
nacién de Diaz, Soderstrom 692 (YU): A, 693 (YU): E. Tuaxcata: Sta.
Ana Chiautempan, Arsene, 10-11-1908 (NY): E. Pursia: 5 miles NE. of
Zacatepec, Soderstrom 483 (YU): L.

M. pulcherrima Scribn. in Beal. MEXICO: Duranco: 15 miles E. of Ciudad,
Reeder 2535 (YU): A(R).

M. purpusii Mez. MEXICO: Sawn Luts Porosf: Near km. 549, 36 mi. 8. of
Matehuala, Mc Vaugh 18204 (US): A.

M. quadridentata (H. B. K.) Kunth. MEXICO: Mexico: Temascaltepec, Las
Cruces, Hinton 4389 (NY): A, L.

M. racemosa (Michx.) B.S. P. USA: Enroute from Fort Duchesne to Dakotah,
Cornman, in 1888 (NY): L.

M. reverchonii Vasey & Seribn. USA: Texas: 8S. of Austin, Stlveus 2478 (YU):
A(R), L.

M. richardsonis (Trin.) Rydb. USA: Cotorapo: Leadville, Goodding M-3272
(YU): A(R).

SODERSTROM—PODOSEMUM AND EPICAMPES 181

M. rigens (Benth.) Hitche. USA: Arizona: Chiricahua Mts., Riggs’ Ranch,
Blumer 1491 (NY): BE. MEXICO: Chihuahua, 27 miles NW. of Madera,
Reeder 2662 (YU): A. Zacatecas: 25 miles NW. of Fresnillo, Soderstrom
731 (YU): A, L.

M. rigida (H. B. K.) Kunth. MEXICO: Jauisco: 2 miles W. of Guadalajara,
Soderstrom 640 (YU): L. Zacatecas: 2 miles W. of Sombrerete, Soderstrom
743 (YU): A, L.

M. robusta (Fourn.) Hitche. MEXICO: Jauisco: Near Guadalajara, Pringle
2321 (NY): E. Micuoacdn: Punguato, near Morelia, Arséne (NY): L.
Mexico: Temascaltepec, Carboneras, Hinton 1853 (NY): A. 3 miles NW. of
Tlamanaco, Soderstrom 382 (YU): L. Cutapas: 20 miles E. of Chiapa de
Corzo, Reeder 2128 (YU): A. GUATEMALA: CuIMALTENANGO: Near
Chimaltenango, Standley 59136 (F): E.

M. schreberi Gmel. USA: Connecticut: New Haven (no voucher): A(R), L.

M. scoparia Vasey. MEXICO: Sinatoa: Mesa Malqueson, Cerro Colorado,
Gentry 5178 (NY): L; 10 miles E. of Santa Lucfa, Reeder 2451 (YU): E.

M. setifolia Vasey. USA: Texas: Guadalupe Mountains, Havard 9 (US,
holotype): A.

M. aff. spectosa Vasey. MEXICO: Duranco: 21 miles E. of Santa Luefa,
Reeder 2476 (YU): A, L.

M. stricta (Presl) Kunth. MEXICO: Jauisco: La Granja, near Guadalajara,
Soderstrom 642 (YU): A, L.

M. subaristata Swallen. MEXICO: Durango: El Salto (Aserraderos), Sierra
Madre Occidental, Pennell 18572 (US holotype): A.

M. sylvatica (Torr.) Torr. ex Gray. USA: Connecricur: New Haven (no
voucher): L(R).

M. torreyana (Schult.) Hitche. USA: New Jersey: Egg Harbor, Bissell (NY):
A(R), L.

M. torreyi (Kunth) Hitche. USA: New Mexico: Capitan, Goodding M-189
(YU): L.

M. uniflora (Muhl.) Fernald. USA: New York: Wading River, Miller 130
(NY): L. New Hampsuire: Chatham, Johnson, August 7, 1927: A(R).

M. versicolor Swallen. MEXICO: Oaxaca: Sierra de San Felipe, Smith 927
(NY): L.

M. virletii (Fourn.) Soderstr. MEXICO: Mexico: Sultepec, Hinton 3219
(NY): A, L.

M. wrightti Vasey. USA: New Mexico: Soil Conservation Service Nursery,
Albuquerque (seed from Flagstaff, Arizona), Goodding and Hardies A-6498
(YU): L.

M. xerophila C. O. Goodding. USA: Arizona: Forest Cabin Canyon, Babo-
quivari Mountains, Goodding A—9493 (YU): L; Moristo Canyon, Baboqui-
vari Mts., Goodding 443-45 (NY): A.

GENERA OTHER THAN MUHLENBERGIA

Agrostis alba L. USA: Connecticut: Guilford, Dudley, July 17, 1904 (YU): L.

Agrostis scabra Willd. USA: Continental Divide, near Beardshead, Goodding,
July 25, 1939 (YU): E(R).

Blepharoneuron tricholepis (Torr.) Nash. USA: ARIZONA: Tiyo Plateau, Grand
Canyon, Reeder 1807 (YU): L(R).

Cinna arundinacea L. USA: Connecticut: New Haven (no voucher): L.
New JeRsEY: Plainfield, Tweedy, September 1881 (YU): E(R).

Lycurus phleoides H. B. KK. MEXICO: Jauisco: 11 miles N. of Ciudad Guzman,
Reeder 2352 (YU): L(R).

182 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Polypogon monspeliensis (L.) Desf. USA: Massacnuserts: Lowell, Swan, July
14, 1880 (YU): L. BELGIUM: Ripe fruits received from Hortus Botanicus
Bruxellensis in 1955: E(R).

Sporobolus asper (Michx.) Kunth. USA: Connecticut: East Haven, Nichols,
Aug. 31, 1912 (YU): L(R).

Sporobolus heterolepis (A. Gray) A. Gray. USA: Connecticut: West Rock,
Nichols, Sept. 5, 1911 (YU): L(R).

Sporobolus wrightii Munro ex Scribn. USA: Arizona: San Pedro, Towmey,
July 25, 1894 (YU): L; ripe fruits collected by L. N. Goodding, October 1955:
E(R).

Stipa avenacea L. USA: Massacnuserrs: Boston, Blue Hills Reservation,
Reeder 233 (YU): E(R), L(R).

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INDEX

(Synonyms in italics. Page numbers of principal entries in boldface.)

Agrosteae, 84
Agrostideae, 75, 78, 80, 81, 83, 93, 97,
100, 103, 108, 110
Agrostis, 75, 76, 78, 79, 81, 82, 83, 84,
93, 97, 100, 101, 107, 108, 110
alba, 105, 181
lanata, 77, 78, 148, 150, 151, 152
pubescens, 77, 78, 148, 150, 151, 152
seabra, 99, 181
Alopecurus, 84
Anthoxanthum, 101
Apera, 77
Arctagrostis, 81
Bauchea, 80
karwinskyi, 80
Bealia mexicana, 82
speciosa, 146
Blepharoneuron, 84, 107, 108
tricholepis, 105, 181
Calamagrostis, 84
Calamovilfa, 103
Cenchrus, 101
Chlorideae, 76
Cinna, 75, 76, 78, 81, 82, 83, 93, 97,
100, 101, 107, 108, 110
arundinacea, 99, 105, 181
lanata, 148
macroura, 80
phleoides, 79
pubescens, 148
stricta, 79, SO
Cinnastrum, sect., 112
Crypsinna, 80, 82, 83
Crypsis, 78, 83, 84
macroura, 78
phleoides, 78, 79
Dichelachne, 84
Echinopogon, 84
Epicampes, sect., 112
Epicampes anomalus, 119
argenteus, 177
arundinaceus, 177
berlandieri, 153, 154
bourgaei, 137
bourgaet var. grandis, 161
bourgaei var. muticus, 137

Epicampes—Continued
buchingeri, 177
clomena, 177
coeruleus, 79, 177
coeruleus var. submuticus, 177
crassiculmis, 177
distichophyllus, 81, 82, 83
distichophyllus var. muticus, 119
ehrenbergii, 137
emersleyt, 166
expansus, 136
giganteus, 137
gracilis, 78, 125
grandis, 161
kunthianus, 177
lanatus, 78, 148, 150
laxiusculus, 137
ligulatus, 82, 83, 119
macrotis, 128
macroura, 80, 81, 83, 177
mexicanus, 177
minutiflorus, 129, 130, 131
muticus, 136, 141
phleoides, 178
pubescens, 78, 148, 150, 151
rigens, 80, 81, 82, 83, 178
robustus, 153
speciosus, 146
strictus, 78, 84, 111, 112, 153, 154,

157

strictus var. distichophyllus, 132, 166
strictus var. muticus, 119
subpatens, 166, 173
virletit, 157

Eragrosteae, 84

Euagrosteae, 79, 80, 81, 82

Festuca, 103

Festuceae, 85, 105

Gastridium, 84

Glyceria, 101

Heleochloa, 84

Lagurus, 84

Lolium, 103

Lycurus, 93, 107, 108
phleoides, 105, 181

Melica, 101
187

188

Muhlenbergia

alta, 137, 139, 140

andina, 104, 106, 109, 178

angustata, 79, 90, 102, 112, 177, 178

angustifolia, 132, 133, 134

anomala, 119

arenicola, 90, 102, 106, 112, 178

argentea, 82, 177, 178

arizonica, 92, 93, 178

articulata, 90, 102, 112, 178

asperifolia, 92, 109, 178

attenuata, 175, 177

aurea, 91, 113, 115, 160, 161

berlandieri, 153

biloba, 82, 104, 177, 178

brachyphylla, 109

breviligula, 91, 118, 115, 148, 159,
160, 161, 172, 178

brevis, 104, 178

brevivaginata, 179

capillaris, 90, 102, 111, 112, 179

carinata, 143

clomena, 82

confusa, 92, 179

crispiseta, 92, 179

curvula, 179

depauperata, 104, 179

distans, 89, 90, 96, 103, 113, 115
128, 148, 153, 1638, 164, 165, 179

distichophylla, 89, 90, 96, 1038, 112,
113, 114, 125, 130, 132, 138, 134,
166, 179

distichophylla var. mutica, 128, 134

dubia, 89, 90, 102, 112, 179

dubioides, 90, 95, 100, 112, 179

elongata, 89, 90, 112, 179

emersleyi, 81, 82, 83, 89, 90, 97, 101,
102, 109, 110, 113, 115, 116, 117,
118, 119, 136, 142, 148, 156, 157,
160, 165, 166, 172, 173, 174, 175,
176, 177, 179

erectifolia, 179

eriophylla, 179

expansa, 90, 102, 112, 179

filiculmis, 179

filiformis, 109

firma, 90, 102, 112, 179

flexuosa, 179

fournieriana, 153, 154

frondosa, 104, 106, 179

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Muhlenbergia—Continued

gigantea, 86, 89, 90, 95, 96, 97, 98,
101, 103, 109, 110, 113, 114, 136.
137, 138, 189, 140, 141, 179

glabrata, 90, 102, 112, 179

glomerata, 104, 106, 179

gooddingii, 91, 113, 115, 117, 118,
119, 173

gracilis, 126

grandis, 89, 90, 96, 97, 98, 101, 108,
113, 115, 144, 145, 148, 161, 162,
179

hintonii, 179

holwayorum, 90, 112, 179

inaequalis, 91, 113, 114, 142, 148

involuta, 91, 102, 1138, 122, 123,
124, 125, 127, 128, 179

iridifolia, 91, 113, 114, 145

jaliscana, 90, 180

lanata, 148, 150

lehmanniana, 91, 118, 115, 172,
174, 175, 176, 177

ligulata, 119

lindheimeri, 78, 86, 87, 88, 89, 90,
95, 96, 103, 109, 110, 113, 122,
125, 127, 128, 180

longiglumis, 91, 97, 102, 113, 115,
165, 166, 180

longiligula, 83, 89, 90, 91, 98, 101,
103, 110, 113, 117, 118, 119, 122,
123, 180

lucida, 90, 112, 180

macrotis, 89, 90, 95, 96, 97, 98, 101,
103, 113, 114, 125, 128, 130, 131,
132, 133, 184, 156, 180

macroura, 80, 89, 94, 95, 96, 97,
99, 100. 101, 102, 106, 109, 111,
112, 177, 180

magna, 137, 139, 140

matudae, 180

metcalfei, 90, 180

mexicana, 109

mexicana f. ambigua, 104, 106, 180

meziana, 129, 130, 131, 1382

minutiflora, 129

montana, 104, 106, 180

aff. montana, 91, 92, 180

monticola, 109

multinodis, 175, 177

mutica, 91, 113, 114, 138, 139, 141

nigra, 90, 97, 99, 100, 101, 102, 106,
112, 180

INDEX

M uhlenbergia—Continued

orophila, 180

palmeri, 86, 87, 88, 102, 106, 112,
180

peruviana, 92, 177, 180

plumbea, 92, 93, 180

polycaulis, 109

porteri, 109

presliana, 153, 154, 157, 158

pubescens, 89, 90, 95, 96, 97, 98,
100, 101, 103, 109, 118, 114, 115,
128, 147, 148, 150, 153, 163, 164,
180

pubigluma, 91, 113, 122, 125, 127,
128

pulcherrima, 92, 180

pungens, 109

purpusii, 180

quadridentata, 104, 106, 180

racemosa, 106, 110, 180

reederorum, 91, 109, 110, 113, 122,
123, 125

repens, 110

reverchonii, 90, 102, 109, 112, 180

richardsonis, 92, 110, 180

rigens, 80, 82, 86, 89, 95, 96, 97, 98,
100, 101, 102, 106, 109, 110, 111,
112, 178, 181

rigida, 90, 102, 109, 112, 181

robusta, 89, 90, 96, 97, 98, 101, 103,
111, 112, 113, 114, 125, 130, 137,
139, 153, 156, 157, 158, 181

schreberi, 97, 99, 101, 104, 106, 107,
110, 111, 181

scoparia, 91, 113, 114, 143, 144,
145, 146, 161, 162, 181

setifolia, 90, 181

speciosa, 89, 90, 96,
114, 146, 147, 148, 150

aff. speciosa, 181

squarrosa, 110

stricta, 91, 102, 113, 153, 181

subaristata, 90, 181

sylvatica, 104, 106, 110, 181

103, 113,

189

Muhlenbergia—Continued
tenuifolia, 110
torreyana, 91, 92, 104, 106, 181
torreyi, 104, 181
uniflora, 104, 106, 110, 181
vaseyana, 83, 166
versicolor, 91, 103, 118, 114, 115,
128, 134, 136, 143, 164, 166, 181
virletii, 89, 90, 96, 113, 115, 156,
157, 158, 181
wrightii, 104, 106, 181
xanthodas, 91, 113, 115, 173, 174,
175, 177
xerophila, 90, 102, 112, 181
Muhlenbergia, subg., 111
Muhlenbergiinae, 84
Pennisetum, 101
Phleoideae, 79, 80
Phleum, 84
Podosemum, 78
captllare, 111
distichophyllum, 78, 132, 166
Podosemum, sect., 95, 102, 104, 111,
112, 113, 117, 122, 177, 178
Podosemum, subg., 111, 112, 113, 122
Polypogon, 81, 84, 97, 100, 101, 107,
108, 110
monspeliensis, 99, 105, 182
Sphaerocaryum, 84
Sporoboleae, 79, 80, 84
Sporobolinae, 84
Sporobolus, 78, 79, 81, 82, 83, 84, 93,
97, 100, 101, 107, 108, 109, 110,
111
asper, 105, 182
heterolepis, 105, 182
rigens, 177, 178
wrightii, 80, 99, 105, 177, 182
Stipa, 78, 97, 100, 101, 107, 108, 109
avenacea, 99, 105, 182
capillaris, 111
Stipeae, 79, 80, 81, 109
Triplachne, 84
Urochondra, 84
Vilfa rigens, 178

U.S. GOVERNMENT PRINTING OFFICE: 1967

CONTR. NAT. HERB. VOL. 34, PT. 4 SODERSTROM—PLATE 1

ee eR

sf ¥. oa 4
a 7 ‘
ia Big es

ld

Muhlenbergias, Podosemum seetion: Fic. 1, Muhlenbergia rigens among rocks in a dry
streambed, 12 mi. W. of Chihuahua, Chihuahua, Mexico, alt. 4,800 ft. (Soderstrom
927-929), Vic. 2.—AM. rigens, closeup of Soderstrom 927, Fic. 3.—M. macroura, pine-
zone area with volcanic soil, La Joya, Veracruz, Mexico, alt, 7,000 ft. (Soderstrom 478),
Fic. 4.-—MM. macroura on eroded bank at edge of pine-oak hillside, 14 mi. E. of Morelia,
Michoacan, Mexico, alt. 6,700 ft. (Soderstrom 551),

CONTR NAT. HERB. VOL. 34, PT. 4 SODERSTROM PLATE 2

Muhlenbergias, Hpicampes section: Pie. 1 Muhlenbergia emersteyt (awnless) near ledge
of arroyo, Laoami. Sli. of Mnearnacion de Diaz, Jalisco, Mexico, alt. 7,000 ft. (Soderstrom

O97). Pie. 2. Clump of MW. emerdeyi Gong awned) growing in protection of “maguey

silvestre” (dave sp.), plateau near San Praneiseo de Oro, Chihuahua, Nlexico, alt.
7,000 ft. (Soderstrom S70). Fig. 3. A. robusta, upper ledge of arroyo, 6 mi. I. of Santa
Cruz. Jalisco, Mexico, alt. 5,300 ft. (Soderstrom 366). Fic. 4.0 MM drstans, rocky ledyc

of arroyo, 15 mi. W. of Zacatecas, Zacatecas, Mexico, alt. 7,200 tt. (Soderstrom 724).

CONTR. NAT. HERB. VOL. 34, PT. 4 SODERSTROM-—PLATE 3

Habitats of Muhlenbergias, subgenus Podosemum: Fic. 1, Clumps of A. pubescens are
common on the rocky desert hillside near Durango, Durango, Mexico, alt. 6,200 ft. Fre.
2— Barranca de Oblatos near Guadalajara, Jalisco, Mexico. Many early C. G. Pringle

collections of Muhlenbergia came from this and nearby barraneas; an area of Mexico that

appears to be the center of distribution of the genus. Fic. 3. MW. rivens is found at the
base and MV. emersleyt higher on the ledges of a dry arroyo, 25 mi. NW. of Fresnillo, Zaca-

tecas, Mexico, alt, 7,300 ft. Pic. 4.—Large clumps of MM. robusta not vet in bloom, 3

mi. NW. of TPlamanaleo, Mexico, Mexico, alt. 7,000 ft. (Soderstrom 380 collected here).

CONTR. NAT. HERB. VOL. 34, PT. 4 SODERSTROM—PLATE 4

Dar

par

Anatomical structures of the leaves of Muhlenbergias, subgenus Podosemum: Vic. 1.—
Muhlenbergia lindheimeri, cross section through central part of leaf, showing heavily
sclerified central unit and mass of thick-walled parenchyma of keel (Burr 513), X50. Fic.
2.—M. lindheimert, cross section through lateral part of leaf, showing two 1° and two 2°
units (Burr 513), 134.) Fic. 3. AM. palmeri, cross section through central part of leaf,
including two 1° units and, between them, one 2° and two 3° units (Pringle 1417), > 134.
Abbreviations: par=parenchyma (thick-walled); s=sclerenchyma; ch—chlorenchyma
sheath (mesophyll); be =bulliform cells; os-=outer sheath; ce—colorless cells; is inner

sheath; x xylem; ph= phloem (line points to darkstained, active, nonsclerosed portion).

CONTR. NAT. HERB. VOL. 34, PT. 4 SODERSTROM—PLATE 5

Photomicrographs of leaf cross sections: Fic. 1.—Muhlenbergia rigens, young innovation
leaf, between ligule and point 2.5 cm. from ligule on blade (Soderstrom 731), «187. Fic.
2.—M. rigens, culm leaf, between ligule and point 1 em. from ligule on blade (Reeder &

§ , : p §

Reeder 2662), 150. Fic. 3.—M. dubia (Soderstrom 509), X 150.

CONTR. NAT. HERB. VOL. 34, PT. 4 SODERSTROM ~ PLATE 6

aoe
—s

Se
Sen er

we

een we :
uasenengretal ( ppomnnnsDeag,

Photomicrographs of leaf cross sections: Fic. 1 Muhlenbergia palmert (Pringle W417),
194.) iG. 2.--M. expansa (4Amer.Gr. Nat. Herb. 1374), < 156. Bic. 3. AM. holweayorum
(Rusby 201), <1 56,

CONTR. NAT. HERB. VOL. 34, PT. 4 SODERSTROM-— PLATE 7

: ik SOT aati
: @%
a

‘eo

Photomicrographs of leaf cross seetions: Fic. 1—AMuhlenbergia macrotis (Mexia 90°98),
“165. Fic. 2.—M. gigantea (Soderstrom 607), 58. Fic. 3.— AM. lindheimeri (Burr 513),
77 (cf. pl. 8, fig. 3).

CONTR. NAT. HERB. VOL. 34, PT. 4 SODERSTROM—PLATE 8

Photomicrographs of leaf cross sections: ie, l.— Muhlenbergia breviligula (Standley 76731),
M165. Fic. 2-—AL. tnooluta (Stlveus 780), “165.0 Bic. 3. AN. lindhetmert (Burr 513),
«206 (cf. pl. 7, fig. 3).

SODERSTROM—PLATE 9

CONTR. NAT. HERB. VOL. 34, PT. 4

24,
"He O5

Fic. 1.—Muhlenbergia gigantea, showing keel

Photomicrographs of leaf cross sections:

Fic. 3.

5.

128), «16

4

-M. robusta (Reeder 2 Reeder

Fic. 2.

(Soderstrom 607), 165,

M. torreyana (Bissell), «165,

10

PLATE

SODERSTROM

4

34, PT

HERB. VOL.

CONTR. NAT.

Se

Lan Me Og

3

Photomicrographs of leaf cross sections:

24),°

wmer \+

1
l

meiitgula (BI

]

eT a Le

L.—AMuhlenbe

396),

Pic.

gle

M. longiglumts (Pring

3. Fic. 3

lo:

capillaris (Stilveus 2

Pig. 2.0 0M.

65.
(

|
»)

CONTR. NAT. HERB. VOL. 34, PT. SODERSTROM-—PLATE 11

Photomicrographs of leaf cross sections: Fic. 1.—Muhlenbergia reverchoni (Silveus 2478), *
165, Fic. 2.—M. elongata (Pringle 938), 165, Fic. 3.—Muhlenbergia aff. montana

(Reeder & Reeder 2572), S165,

CONTR. NAT. HERB. VOL. 34, PT. 4 SODERSTROM—PLATE 12

Photomicrographs of leaf cross sections: Fic. 1—AMuhlenbergia peruviana (Pearson 1),
165. Fic. 2.—M. pulcherrima (Reeder 3 Reeder 2535), X 165, Fic. 3M. arizonica
(Reeder €F Reeder 2602), 165,

CONTR. NAT. HERB. VOL. 34, PT. 4 SODERSTROM—PLATE 13

Photomicrographs of leaf cross sections: Fic. 1.— Muhlenbergia confusa (Reeder 9 Reeder
2684), X 165. Fic, 2.—M. richardsonis (Goodding M-3272), 206, Fic. 3.—M. plumbea
(Reeder F Reeder 2218), < 165,

CONTR. NAT. HERB. VOL. 34, PT. 4 SODERSTROM—PLATE 14

Photomicrographs of leaf cross sections: Fic. 1. Muhilenbergia erty piseta (Reeder S Reeder
2545), 165.) Fic. 2--M. asperifolia Ulardies M103), > 163. Fic. 3M. uniflora
(Johnson), 206,

BULLETIN OF THE UNITED States NaTtTIonaL MUSEUM

CONTRIBUTIONS FROM THE UNITED STATES NATIONAL HERBARIUM

VoLumME 34, Part 5

THE GENUS MELOCHIA L.
(STERCULIACEAE)

By Aaron GOLDBERG

SMITHSONIAN INSTITUTION PRESS e¢ WASHINGTON, D.C. © 1967

THE GENUS MELOCHIA L. (STERCULIACEAE)
By Aaron GOLDBERG

History of the Genus

Linnaeus validated the name Melochia by including it in the first
edition of his Species Plantarum (37). In his Genera Plantarum (38)
of the following year under the description of Melochia he cited only
Dillenius (9), who had described and illustrated only Melochia cor-
chorifolia in the genus Melochia. According to the current Inter-
national Code of Botanical Nomenclature (33), it is agreed to associate
generic names which appear in Linnaeus’ Species Plantarum, edition 1,
1753, with the first subsequent description given under those names in
Linnaeus’ Genera Plantarum, edition 5, 1754. Therefore, Melochia
corchorifolia L. is considered to be the type species of the genus.

Melochia is a name altered from the Arabic name of Corchorus
olitorius L., a salad plant in the East, and according to Gray (20)
the name was borrowed by the Arabs, through the Syriac, from the
Greek word for the mallow.

Several genera have been published and subsequently reduced to
synonymy under Melochia. In 1777 Houttuyn (27) introduced the
name Visenia for his V. umbellata, a common tree of southeastern
Asia and the Pacific Islands. In 1854 Gray (19) expressed the opinion
that Visenia could not be generically distinguished from Melochia.
In 1895 Schumann (48) gave it its present status as a section of
Melochia.

In 1787 Medicus (42) introduced the name Moluchia for species
with pyramidal fruits—placed in the section Pyramis in this paper.
In 1803 Ventenat (64) introduced the name Riedlea, characterized by
a 5-valved globular capsule, filaments united in a column, and a calyx
subtended by 3 bracts. Riedlea is placed in the section Melochia in
this paper. In 1823 Humboldt, Bonpland, and Kunth (28) intro-
duced Mougeotia for species with a pentacoccic capsule. DeCandolle
(8) reserved the name Melochia for species placed in the section
Pyrams in this paper, and used the name Riedleia for Riedlea Vent.,
Mougeotia H. B. K., and Visenia Houtt. He also established the tribe
Hermannieae, which includes Hermannia L., MelochiaL., and Wailtheria L.
Two years later Sprengel (51) used Melochia as did DeCandolle, but
substituted Visenia for Riedleia, probably on the basis of priority.

191

192 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

In 1825 St.-Hilaire (46) reported on his rather extensive collections
of Melochia in southern Brazil, and gave reasons for not considering
Riedlea Vent. generically distinct from Melochia. He mentioned that
the flowers are very similar and that, as a rule, other characters being
the same, Linnaeus would not divide a genus if the flowers were the
same but the fruits different. He did not consider the presence of
bracts subtending the calyx a generic character and pointed out that
there are intermediate forms in the types of dehiscence of the fruit in
Melochia and Rvedlea.

In 1836 Presl (45) introduced Physodium, and mentioned that it
differed from Riedleia in having an inflated, closed calyx, linear
anthers, and an acute-angled, stipitate capsule. Bentham and Hooker
(3) maintained Melochia and Physodium, but, considering St.-Hilaire’s
observations and their own, they placed Riedlea in synonymy.
Hemsley (25) did not consider Physodium to be generically distinct
from Melochia but followed Bentham and Hooker in maintaining it.

In 1854 Triana and Karsten in Triana (58) introduced Anamorpha,
and in 1858 Turezaninow (60) introduced Physocodon. Both have
since been reduced to Melochia. They contain species now placed in
the section Mougeotia.

In 1864 Grisebach (22) divided the genus into the sections Humelo-
chia, Riedleia, and Mougeotia. As emended in description, and as
changed in nomenclature, in accord with the International Code of
Botanical Nomenclature, these sections are maintained in the present
paper.

Schumann (47), in the next major treatment of the genus, considered
all of the genera mentioned above, occurring in Brazil, to be synonyms
of Melochia, and he accepted Grisebach’s sectional division of the
genus.

In 1890 Garcke (17) discussed part of the extensive synonymy in
the genus.

In 1895 Schumann (48), in treating Melochia, added Visenia as a
section containing certain species of Asia and the Pacific Islands, and
reduced Physodium to synonymy.

In 1908 Fries (15) presented his concepts of the taxa in Melochia in
South America. He examined and cited specimens from several
European herbaria.

From time to time species and infraspecific taxa have been described
in the genus Melochia and its synonyms. These are considered in the
systematic section of this paper. Approximately 379 names and com-
binations have been used to describe the types of Melochia. Many are
synonyms and many of the descriptions are very inadequate. I have
made more than 30,000 observations of characters in the genus.
About half of them were measurements under the dissecting micro-

GOLDBERG—GENUS MELOCHIA 193

scope. Thus, I have supplemented the information in the old descrip-
tions. As a result of my studies, I consider 68 taxa to be adequate
to describe the entities in the genus. Two new species are described
and Physodium is placed in sectional status.

Division of the genus into several genera has been based on charac-
ters which show a gradual transition between species from section to
section as follows: (1) The carpels of some species of the section
Mougeotia have an obtusely rounded dorsal surface; in others the
carpels are somewhat carinate in the distal half of the fruit and transi-
tional to the section Pyramis. (2) In the size of the fruit from the
section Mougeotia to the section Pyramis. (3) One type of dehiscence
to another occurs between species. (4) The gradual change in degree
of accrescence of the calyx, degree of union of the filaments, and
length of the stipe. (5) In the size of bracts in all species and in the
length of the internode between each of them and between them and
the calyx.

In addition to the taxonomic treatments already mentioned, several
floras and lists have been published in which pertinent Melochia are
presented. Included among the regions treated are Mexico (52),
several countries of Central America (25, 53, 54, 55, 56), several of the
islands of the West Indies (4, 13, 22, 36, 62), Surinam (61), French
Guiana (35), Venezuela (44), Colombia (59), Peru (39), Bolivia (14),
tropical West Africa (29), Congo (18), Angola (11), the basin of the
Zambesi River (12), Madagascar (1), India (26), Indochina (16, 57),
Java (2), the Philippine Islands (43), and Fiji (49). Since the taxa
attributed to the various regions by the authors of these floras and
by me differ in some cases, I am including tables showing the distribu-
tion of the species of Melochia by countries (pp. 196 and 197).

Economic Importance

The genus Melochia has no outstanding economic importance.
Some species, such as M. pyramidata, have fine silky fibers, but at
present these are used only locally. Some species occur in pasture
lands, especially savannas. Gentry mentioned (his specimen no.
943M [MICH]) that burros are very fond of the plant (M. tomentelia).
A few species are weeds in cultivated fields. The plants have a
mucilaginous constituent that is used locally as a demulcent in enter-
itis due to dysentery. They are also used locally as a salad plant,
and for smoking. The flowers are a source of food for honeybees.

Melochia umbellata (Houtt.) Stapf, a rapidly growing species of
southern Asia and the Pacific Islands, is commonly planted as a shade
tree in that part of the world.

194 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Geographic Distribution

In America, Melochia is distributed over more than 5.6 million
square miles. It ranges from approximately 33° N. to 33° S. latitude
on the east coast, and from approximately 29° N. to 12°S. latitude on
the west coast. The distribution of the genus in South and North
America is shown in figures 1 and 2, respectively. The coordinates

SCALE

9 100 700 300 400 500 600 MILES

200 400 600 800 RILOMETERS
so" 1000 FOOT CONTOUR
SINUSOIDAL EQUAL-AREA PROJECTION

Ficure 1.—Distribution of Melochia in South America.

of many of the localities of the specimens studied were determined
from the “Index to the Map of Hispanic America” (31).

195

GOLDBERG—GENUS MELOCHIA

*BOTLIOUIYY YWUoN ul 14 I077fT JO uoTnqiisiq—Z FANT

ey

206

——
Yo}ausaI5

ogy oge_oge

oot

9

SalIW 40 3179S

196

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

TABLE 1.—Distribution of American Species of Melochia by Countries

MELOCHIA

adenodes
anomala
arenosa
argentina
bernoulliana
betonicifolia
caracasana
chamaedrys
colombiana
corchorifolia *
corymbosa
crenata
gardneri
graminifolia
hassleriana
hermannioides
illicioides
kerriifolia
lanata
lanceolata
leucantha
lupulina
manducata
melissifolia
mollis
morongii
nodiflora
nudiflora
parvifolia
pilosa
pyramidata
rhodocalyx
savannarum
simplex
splendens
thymifolia
tomentella
tomentosa
tragiifolia
ulmifolia
villosa
werdermannii

United States

Mexico

Guatemala

|
|
|

xX
xX x

xxxX XX

x XxX

British Honduras

Honduras
Salvador

Nicaragua

Costa Rica
Panama

x XX

Bahamas
Cuba

x xX xX xX

Hispaniola
Jamaica

|
|

xx

Puerto Rico

|

Lesser Antilles

|

Trinidad-Tobago
French Guiana
Surinam

|
|
|
|
|
|
|

British Guiana

Venezuela

m

xxx xX xX

xX
xX

Colombia
Ecuador

Peru

xxXX KX XK XX

x_X

xX

xX

Bolivia

North Brazil

|

|

x xX xX

xX xX xX X*X

xx

South Brazil
Uruguay

Paraguay
Argentina

xXx

x xX

x x x
x xX xX

*Adventive.

The genus occurs in more than 6.7 million square miles of the Old
World, from 37° N. in eastern Asia to 24°S. in the Pacific Islands,
and from 18° N. to 16°S. in western Africa and 2°S, to 20°S. in

eastern Africa.

The distribution of the genus in Asia, the East

Indies, and Africa is shown in figures 3, 4, and 5, respectively. The

GOLDBERG—GENUS MELOCHIA 197

TABLE 2,— Distribution of Extra-American Species of Melochia

MELOCHIA

New Caledonia

Fiji Islands
East Pacific

Philippines
Caroline &

Solomon Islands to
Marianas

Madagascar
East Indies,
New Guinea
Australia
Samoa, Tonga &

Africa
Asia

aristata x
bracteosa x
compacta
corchorifolia x1x}] x] xX |x I xX
degeneriana
grayana
lanceolata x
longepetiolata
melissifolia x
mollipila
odorata x x
pyramidata * x x | xX
roseiflora x
tomentosa * x
umbellata x |x | x
villosissima x
vitiensis x

x

x x

x eK KX

* Adventive.

coordinates of many of the localities were determined from the Co-
lumbia Lippincott Gazetteer of the World (5).

Distribution of the various taxa is given in greater detail at the
end of the respective descriptions. Distribution of species of Melochia
by countries is also given in tables 1 and 2.

Thirty-nine species are considered to be endemic to America, one
to southern Asia and the southwest Pacific islands, ten to the south-
west Pacific islands, one to Africa, two to Africa and South America,
and one to Asia, Africa, and the southwest Pacific islands.

Two American species occur as weeds in the Eastern Hemisphere
and one species has been transported from the Eastern Hemisphere
to America, perhaps as a weed seed in shipments of rice.

Some species occur in deserts with less than 10 inches of annual
rainfall, others grow in tropical rain forest where the annual pre-
cipitation is 126 inches.

FROM THE NATIONAL HERBARIUM

CONTRIBUTIONS

198

ainjelay] sy] Wolj=— WV }X9] Ul p9lb =@

‘[(9Z) J9yOOR{] uopsaa pur ‘eso¥"]eyy
‘WIYyYIG Oo} uoeWUINy wWoly ‘elpu] jo syied Jo}0Y 94} Ul pelnqisisip Ajjeiouas sI 0170/1404 9409 p1y20jajy

“eBISy Ul piyIJojJapy Jo woTNdIysIq—'¢ aanoly

06 oes

199

GOLDBERG—GENUS MELOCHIA

ainielaiy] ay} Woy = Xa} UI psp =@

"saIpuy ISeY Yi Ul yIyI07apy JO UONNqISIG—'F TANI

200 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Boo Lets
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409 aco. 1200 HILOMET ERS,

SINUSOIDAL PROJECTION

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- oe es ee Se
v 10 20 30 4

Ficure 5.—Distribution of Melochia in Africa.

@= cited in text A= from the literature

Taxonomic Characteristics

Most species of Melochia are perennial shrubs or subshrubs; a few
are herbs or small trees. A taproot is usually present, but secondary
roots are often numerous. The young stems sometimes are hollow or

GOLDBERG—GENUS MELOCHIA 201

have a large pith and are often provided with small, round lenticels.
They may be pubescent or glabrous. The bast fibers of the stem are
tough.

Several kinds of hairs occur in the genus. They may be stellate,
with 3 to 13 rays, forked, simple, or glandular. The last are several
cells long, with the distal cell enlarged.

The leaves are simple and alternate and vary greatly in size, shape,
and pubescence between species, sometimes within species. Their
margins are usually irregularly crenate-serrate.

The inflorescences are axillary, terminal, leaf-opposed (the peduncle
leaving the stem 180° from the insertion of the petiole at the same
node), or 90° from the leaf insertion. They are cymose. The pe-
duncle may be simple, branched, or absent. The bracts and stipules
are usually similar in shape, size, and pubescence.

The flowers are polypetalous, actinomorphic, perfect, and hypog-
ynous. They are sometimes matutinal and sometimes fragrant and
visited by honeybees. They are usually dimorphic, the gynoecium
and androecium differing in length in flowers of different plants of a
species. The condition may be slight or marked. In figure 6, 14, 1B,
3A, and 3B illustrate the longistylous condition, and 24, 2B, 4c, and 4p
the brevistylous condition.

The five fertile stamens are antepetalous, and the filaments are free
of the petels or partially attached to them. The filaments are par-
tially or completely attached to one another. Staminodes are usually
absent; sometimes there are five, usually dentiform, alternate with the
fertile stamens. The anthers are dithecous, extrorse, and attached to
the filaments at the middle. The pollen is smooth, usually globular
and 34-49u in diameter. Sometimes, according to Erdtman (10), it
is subprolate, and 51y long and 44u wide.

There are usually 5, rarely 6-8, carpels opposite the petals. The
placentation is axile and the ovules are attached one-fourth to one-third
the way up from the base of the carpel. The ovules are superposed
and close to one another, and anatropous, with the funicle ascending
on the adaxial surface of the ovule.

The calyx is campanulate, quinquefid, valvate in aestivation, and
persistent. Sometimes glandular tissue is between the calyx and the
corolla and between the bases of the petals.

The 5 petals are imbricate and contorted in aestivation and often
marcescent. They are flat and veined.

The fruit may be subglobular, and more or less pentagonal (figure
6, 1p, 2D) or pyramidal and pentapterous (figure 6, 3c). A rostrum
may be present (figure 6, 3c), or absent, or the apex of the capsule may
be retuse (figure 6,48). A fairly long stipe may be present (figure 6,
4E) or it may be absent. Dehiscence of the fruit may be (1) septicidal

202 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

into cocci with a narrow ventral slit, (2) loculicidal all along the ven-
tral suture, the cells gaping ventrally, and a fifth to half way along the
dorsal suture, and sometimes eventually septicidal (figure 6, 1c), or
(3) loculicidal all along the dorsal suture and then often septicidal
(figure 6, 2D).

The embryo is straight, 1.5-2.7 mm. long, and embedded in fleshy
endosperm. ‘The cotyledons are suborbicular, 0.7-1.7 mm. in diam-
eter, flattened, subcordate at the base and rounded at the apex. Some-
times 2 veins are on each side of the middle of the cotyledon; one pair
is basal. The radicle is terete, 0.7-1.1 mm. long, 0.25-0.5 mm. wide,
and inferior. A plumule is not evident in the seed stage. Venkata
Rao (63) studied the early stages in the development of the embryo
of M. corchorifolia L.

Cristébal (6) studied the chromosomes of two species of Melochia.
In both the diploid number was 18. According to Mangenot & Man-
genot (41) the diploid chromosome number of M. bracteosa is 14.

Plate 5 illustrates a typical species of Melochia. Some floral and
fruit characters of a species of each section are illustrated in figure 6.

Taxonomic Position

Melochia is a genus of the family Sterculiaceae sensu lato, tribe
Hermannieae.

Schumann (48) included eight tribes in the family. The flowers of
the Sterculieae are polygamous, those of the other tribes are hermaph-
roditic. The petals of the Hermannieae and Helictereae are flat and
usually longer than the calyx; those of the Buettnerieae are cucullate
at the base and embrace the anthers, often ligulate at the apex, and
usually not longer than the calyx. They are absent from the Ster-
culieae and Fremontieae, absent or minute, glandlike, or squamiform
in the Lasiopetaleae, and large in the Dombeyeae. In the Lasio-
petaleae the stamens are usually free; they are at least partly united
in the other tribes. The staminal tube is greatly elongated in the

Ficure 6.—Floral and Fruit Characters of Melochia. 1, Melochia chamaedrys St.-Hil. var.
chamaedrys, Pedersen 971 (US): a, Flower, one petal removed; B, flower with calyx re-
moved; c, fruit, lateral view; p, fruit, apical view. 2, Melochia arenosa Benth., flower,
Dahlgren &$ Sella 65 (US); fruit, Gardner 2489 (US), syntype: a, Flower, one petal removed;
B, flower with parts removed to show essential organs; c, fruit, lateral view; p, fruit, apical
view. 3, Melochia anomala Griseb., Venturi 2108 (US): a, Entire flower; B, flower opened
to show essential organs; c, fruit, lateral view; p, fruit, apical view. 4, Melochia adenodes
A. Goldberg var. adenodes, Hinton 16203 (US), type: a, Entire flower; B, flower with calyx
partly removed; c,p, flower with parts removed to show essential organs; £, fruit, lateral
view; F, fruit, apical view. 5, Melochia uwmbellata (Houtt.) Stapf, flower, Krukoff 4421
(US); fruit, Bartlett 8367 (US): a, Entire flower; B, flower with parts removed to show
essential organs; c, fruit, lateral view; D, fruit, apical view. (The scale at the right of
each illustration represents 1 mm.)

203

GOLDBERG—-GENUS MELOCHIA

204 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Eriolaeneae and Fremontieae. The flowers of the Hermannieae have
five fertile stamens with dithecous anthers; the stamens of the other
tribes are sometimes more numerous and sometimes the anthers are
more than dithecous. The gynophore is usually long in the Ster-
culieae and often so in the Helictereae; it is usually short or absent in
the other tribes. The carpels of the Sterculieae are usually separate
at anthesis; they are at least partly united in the other tribes.

The tribe Hermannieae contains two subtribes, the Hermanninae
and Melochinae. The former contains the genus Hermannia. Its
pistil consists of five carpels opposite the calyx lobes. Each carpel
contains an indefinite number of ovules. The styles are sometimes
united at the apex. The seeds are reniform, the embryo is curved,
and the cotyledons are oblong and twice as long as wide. The
filaments are free or united part way, and the free part is often flat-
tened and broader than the anthers. The anthers are relatively long
(about 3mm. long). The subtribe Melochinae contains Melochia and
Waltheria. The pistil of Melochia consists of five antepetalous carpels.
Each carpel contains two ovules. The styles are free to the ovary or
united part way at the base. The seeds are not reniform, the embryo
is straight, and the cotyledons are suborbicular. The filaments are
united part way or up to the anthers and the free part is not broader
than the anthers. The anthers are shorter (less than 3 mm. long).
Waltheria differs from Melochia primarily in that its pistil consists of
one carpel. In Waltheria the apex of the style is fimbriate.

Evolution and Relationships

Melochia belongs to a relatively primitive family, the Sterculiaceae,
that probably originated at a time when the temperature did not go
below freezing, at least not for any considerable period, and moisture
was abundantly available in all latitudes. The family migrated from
continent to continent. With changes in the climate, it did not
survive in colder regions, but at present is well represented in tropical
and subtropical areas throughout the world. Melochia probably
originated in America and the great majority of the species occur here.
Members of the genus with primitive characters range from southern
Mexico to southern Brazil and northern Argentina, in southeastern
Asia, and the Pacific Islands. The three species indigenous to
Africa have relatively few primitive characters, and the one species
indigenous to Australia is advanced.

Some species flower for extended periods under a variety of condi-
tions; others reproduce only under particular conditions. ‘The former
species have a wide distribution.

Species of Melochia are usually perennial subshrubs or shrubs, but
toward the latitudinal extremes of the range of the genus, and where

GOLDBERG—GENUS MELOCHIA 205

available moisture is reduced, the species become herbaceous, and
a few are annuals which avoid the rigors of a cold or dry season. Other
species have become adapted, by means of a dense indument, to
withstand moisture stresses resulting from relatively cool or hot and
dry conditions. In species with dense pubescence, stellate hairs are
usually more abundant than other types. Stellate hairs occur most
commonly in the sections Pyramis and Physodium; simple ones are
most common in the sections Melochia and Mougeotia. The leaves
of species growing in xerophytic habitats are usually thicker than in
those that grow under mesophytic conditions. In some species,
when the shoot is killed due to unfavorable seasonal conditions, the
base of the stem and root remain alive and perpetuate the plant.

As a result of the change to a drier climate, mutant genes tending
to enhance survival under xerophytic conditions accumulated. The
plants became more compact, and thus conserved moisture by ex-
posing less surface to the environment. For example, leaves became
smaller and the internodes of the vegetative stems and inflorescences
became shorter. The stipe is well developed in the section Physodium
but is much reduced or absent in the other sections. The trend was
from corymbiform, paniculate, and umbellate inflorescences to
spicate, subcapitate, and capitate ones. As is frequently the case in
capitate inflorescences, the entire flower or some of its organs become
reduced in size. This is especially true of the calyx; in a few species
the reduced calyx has become membranous. The sinus between the
teeth of the primitive calyx is acute but rounded or truncate in the
species with a reduced calyx, and the teeth are short and distant.
This trend toward reduction of the calyx is most pronounced in the
section Melochia. In some species of this section the internodes
between the bracts have become reduced so that the latter immediately
subtend the flowers; in other species the bracts are enlarged, closely
invest the flowers, and take over the protective function of the calyx.

Although some species, such as Melochia corchorifolia, have ac-
cumulated mutations adapting them to xerophytic conditions, they
have retained their ability to grow in mesophytic and hydrophytic
habitats.

An evolutionary trend in many lines of organisms is the reduction
and loss of parts. In Melochia this has occurred in the flower. Stam-
inodes, the vestiges of one of the cycles of stamens, are elongated and
most evident in some species of the section Physodium. In several
species, particularly those of the other sections, they are merely
dentiform or entirely absent. The fertile stamens show a trend to-
ward the loss of connective between the anther sacs, so that the species
advanced in this respect have deeply emarginate anthers and di-
vergent anther sacs. This is especially the case in the sections Mou-

227-051—67——_2

206 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

geotia and Melochia. The common number of carpels is five, but
species primitive with regard to this character frequently have six
to eight. There are two ovules in each carpel, but in many species
one aborts and only one seed develops.

The primitive colors of the petals appear to be yellow and orange,
the most common colors in species of the section Afougeotia. In
general, plants with rose or purple flowers are more attractive to
hymenopterous insects. In a few species of Melochia with yellow
or orange petals the essential organs are only slightly dimorphic,
whereas the heterostylous condition in flowers of different plants of
the same species, often associated with insect pollination, is quite
evident in rose- and purple-flowered species, predominant in the
other sections, except species in which the flowers have become
considerably reduced in size.

It is the consensus among botanists that the organs of the flower
are homologous with leaves and that, like leaves, in the primitive
condition these organs are separate. All species of the genus have
partially united sepals; however, the extent of union varies between
species. In some species of the section Melochia two or three of the
bracts may be partially united. In the section Pyramis the petals
are attached to the staminal tube at the base. In some species of
the sections Mougeotia and Melochia the petals are free of the stami-
nal tube; in others they are attached to it at the base. In these
two sections the claw is usually sharply demarcated from the blade
of the petal, just as the petiole is demarcated from the blade of
the leaf. In the section Pyramis the claw is usually not sharply
demarcated from the blade of the petal. Ordinarily the petals are
free from one another, but in one specimen of a species in the section
Pyramis, I noticed that the claws were united. In the sections
Melochia and Mougeotia the filaments in the longistylous form of the
flower are usually united up to the anthers, but united to a lesser
extent in the other sections. In the section Physodiwm the styles
are free to the ovary, in the section Mougeotia they are usually free
to the ovary, and often free in the section Melochia, but united at
the base in the section Pyramis. The cells of the fruit vary in the
extent united in the different species. In species of the section
Melochia the cells are usually united for a greater distance along
their sides than in species of the other sections.

The fruits of some species of the section Mougeotia dehisce in the
most primitive manner, along the ventral suture, the site of union of
the infolded carpel phyllomes, and in some species of this section
dehiscence is also septicidal. In the section Visenia the fruit dehisces
ventrally, but it is incompletely septicidal. In other sections dehis-
cence is first loculicidal along the dorsal suture, the most advanced

GOLDBERG—GENUS MELOCHIA 207

means of dehiscence in the genus, and in a few species of the section
Melochaa it is no longer also septicidal.

In Melochia the differences between many species are small. I
believe that there has been little, if any, extinction of species in the
genus. Perhaps the genus has diversified mainly as species became
adapted to different climatic conditions and habitats.

Taking into account the above-mentioned evolutionary changes,
geographical distribution, and the genetic principle that the simi-
larity of structure among species of the same genus is primarily
indicative of genetic relationship, I have arranged the species of
Melochia according to my understanding of their relationships (fig. 7).

Acknowledgments

I wish to express my appreciation to Dr. Lyman B. Smith, Senior
Scientist, United States National Herbarium, for his effort and time
expended in my behalf and the patience he has shown in directing
my research.

It has been a privilege to use the facilities of the United States
National Herbarium for the study of specimens of Melochia. I am
also grateful to other herbaria in America and Europe and their
curators for the loan of specimens. Herbaria are abbreviated accord-
ing to Lanjouw and Stafleu (34).

The portion of the paper dealing with the American taxa was
submitted to the faculty of the Graduate Council of the George
Washington University in partial satisfaction of the requirements for
the degree of Doctor of Philosophy.

Figure 2 is based on map 19AA, McKinley Publishing Co., Phila-
delphia, Pa.; all other maps are based on Goode Base Maps, copyright
by the University of Chicago Press.

Systematic Treatment

Melochia L.

Melochia L. Sp. Pl. I. 2:674. 1753.
Visenia Houtt. Handleid. 8:308. 1777.
Moluchia Medic. Malv. 10. 1787.
Meluchia Medic. in Staatsw. Vorles. Churpf. Phys. Oek. Ges. 1:217. 1791
Riedlea Vent. Choix. 37, pl. 37. 1803.
Altheria Thou. Gen. Nov. Madagascar. 19. 1806.
Mougeotia H. B. K. Nov. Gen. & Sp. 5:326, pl. 483, 484. 1823.
Riedleia DC, Prodr. 1:490. 1824.
Aleurodendron Reinw. in Sylloge Ratisbonn 2:12. 1828.
Glossospermum Wall. Cat. n. 1153. 1829.
Riedlia Dumort. Anal. Famil. 46. 1829.
Polychlaena G. Don, Gen. Syst. 1:488. 1831.

208 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

M. corchorifolia M. pyramidata

M. manducata — M. tomentosa

M. lanceolata M. crenata
M. simplex

M. graminifolia

M. lanata
M. arenosa

M. caracasana
M. betonicifolia
M. illicioides

M. melissifolia
M. bracteosa

M. splendens —_

M. villosa

. ea M. werdermannii
M. savannarum M. anomala

M. morongii

M. tomentella M. parvifolia
M. tragiifolia
M. nodiflora

M. nudiflora

M. bernoulliana

M. hermannioides

M. thymifolia
M. leucantha
M. lupulina
M. mollis
M. colombiana

M. gardneri
M. ulmifolia
M. argentina
M. kerriifolia
M. chamaedrys
M. pilosa
M. hassleriana

M, villosissima
M. compacta

M. grayana M

Wh snccihoy . corymbosa
. rosel ora M. rhodocalyx
M. mollipila M. adenodes

M. degeneriana

M. aristata
M. longepetiolata
M. vitiensis
M. odorata
M. umbellata

Figure 7.—Relationships of the species of Melochia.

GOLDBERG—GENUS MELOCHIA 209

Physodium Presl, Rel. Haenk. 2:150. 1836.

Lochemia Arn. in Ann. Sc. Nat. 2 sér. 11:172. 1839.

Ridleia Endl. Gen. 1000. 1840, attributed to Vent.

Riedleja Hassk. Catal. Pl. Hort. Bog. Alt. 205. 1844.

Anamorpha Triana & Karst. in Triana, Nuev. Jen. i Esp. Fl. Neo-Granad. 9.
1854.

Physocodon Turcz. Bull. Nat. Mosc. 31(1):212. 1858.

Mostly shrubs or subshrubs, sometimes herbs or small trees, 0.08-
15.0 (-20) meters high, mostly perennial, often with a woody taproot,
sometimes 2-5 times thicker than the base of the stem and 0.4-2.2
cm. wide, the young stems subglabrous to cinereous or canescent,
hirsute, pilose, sericeous, or tomentulose, hairs stellate, simple, forked
or glandular, sometimes tan, golden, or ferruginous, 0.05-4.0 mm.
long, the internodes 0.2-18.5 cm. long; leaves simple, alternate;
stipules 1.5-10.3 mm. long, 0.2-4.0 (-12) mm. wide, deltoid-acuminate
or deltoid-acute, generally lanceolate or linear, sometimes ovate or
suborbicular, usually ciliate and hirsute, tomentulose or sericeous,
hairs simple, stellate, forked or glandular, 0.05-3.5 mm. long; petiole
0.07-14.5 (-20.5) cm. long; lamina 0.6-24 (-30) cm. long, 0.2~—21.5
(-27) cm. wide, ovate, lanceolate, linear, oblong-ovate, suborbicular,
or elliptic, one side sometimes larger than the other, the base usually
rounded to cordate, sometimes truncate, cuneate or oblique, the apex
usually acute to acuminate, sometimes rounded, both surfaces gla-
brous, hirsute, sericeous or tomentulose and canescent or cinereous or
brownish, sometimes velutinous or lanate, hairs stellate, simple, forked,
or a combination of these, sometimes also glandular, 0.05-4.0 mm.
long, rarely covered with resin-colored granules or pulverulent, the
margin usually irregularly crenate-serrate, serrations 0.3-12.0 mm.
wide, 0.2-3.6 mm. high, 4-15 pairs of primary veins, usually 2 being
basal, usually impressed above and prominent beneath.

Inflorescences axillary, terminal, leaf-opposed, or 90° from the leaf,
umbellate, subcapitate, paniculate, spicate or corymbiform, cymose,
0.6-34 cm. long, the peduncle 0-11 (-16) cm. long, sometimes with a
primary branch 0.2-5.2 (-7.0) em. long, a secondary one up to 2.0
(-4.0) cm. long, a tertiary one up to 1.6 (3.0) em. long, and rarely
a quaternary one up to 2 cm. long, the pedicel 0-14 (-19) mm. long;
bracts 0.5-12.2 mm. long, 0.1-3.9 (-14) mm. wide, sometimes 2-3
immediately subtending each flower, at other times at the base of the
pedicel, sometimes 2-3 united part way; flowers: few to numerous,
fertile stamens 5, antepetalous, staminodes usually absent, sometimes
5, alternate with the stamens, 0.1-1.2 mm. long, usually dentiform,
sometimes elongated, the anthers 0.4—-2.6 mm. long, 0.3-1.4 mm. wide,
oblong, emarginate at both ends, sometimes nearly to the middle from
the apex, the carpels 5, rarely 6-8, antepetalous, styles filiform,
sometimes slightly clavate at the apex, ovules 2 per carpel.

210 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Calyx quinquefid, valvate in aestivation, persistent, usually green,
sometimes rose, rose purple, yellow, orange, red, or pinkish brown,
sometimes canescent, sometimes membranous, 1.4-15.0 mm. long,
2.1-10 mm. wide, the teeth 0.4-7.8 mm. long and 0.2-4.8 (-5.5) mm.
wide, sometimes accrescent, becoming up to 36 mm. long and 19 mm.
wide, with teeth up to 12 mm. long and 10 mm. wide, the teeth usually
deltoid-acute or deltoid-acuminate, sometimes ovate or nearly linear,
hirsute, tomentulose, or sericeous, sometimes ciliate, hairs simple,
stellate, glandular, or forked, 0.05-2.3 (-4.0) mm. long, the sinus
usually acute, sometimes rounded to truncate; petals purple, pink,
purple distally and yellow or orange proximally, yellow, orange, white,
blue, red, white with red stripes, or violet, 2.7-17.4 mm. long, 0.8-9.5
mm. wide, usually obovate or oblanceolate, sometimes obdeltoid,
obovate-oblong, elliptic, oval, ovate or oblong, cuneate or spatulate,
free of the staminal tube or attached to it for 0.3-5.0 mm., the claw
0-6.5 mm. long, rarely glandular tissue present between the calyx and
corolla, with lobes extending between the bases of the petals, some-
times a few glandular, simple, forked, or stellate hairs on the petals.

Longistylous form. Stamens 2.4-7.0 (-9.0) mm. long, the fila-
ments united up to the anthers or free for 0.2-3.0 mm., sometimes a
few hairs at the base of the staminal tube; pistil 2.6-10.9 (-12) mm.
long, the styles free for 1.2-6.7 mm., to the ovary or united for 0.3-4.0
mm., papillose at the apex for 0.5-2.5 mm.

Brevistylous form. Stamens 2.8-11.3 mm. long, sometimes with a
few hairs at the base of the staminal tube, the filaments free for
0.5-7.5 mm.; pistil 2.3-6.9 (-8.0) mm. long, the styles free for 0.8-5.3
mm., to the ovary or united for 0.2-3.0 mm., papillose for 0.3-1.0
mm. at the apex.

The flowers of several species in the section Melochia not dimorphic,
stamens and pistils 1.7-3.2 mm. long, the filaments free for 0-1.2 mm.,
the styles free for 0.6—1.2 mm., to the ovary or united for 0.1-0.3 mm.

Fruit schizocarp or capsule, 2.4-10.5 (-12) mm. long, 2.9-17.0
(-22) mm. wide, subglobular and pentagonal, pyramidal and pentap-
terous, ovoid or oblong, the sulci between the carpels shallow to deep,
extending a fifth to half way to the center of the fruit, usually hirsute,
tomentulose, or sericeous, subglabrous to canescent, hairs stellate or
simple, sometimes also forked or glandular, 0.1-1.6 mm. long, the
stipe 0-6.7 mm. long; dehiscence septicidal, loculicidal, or both;
seeds brown to black, 1 or 2 per cell, 1.4-3.9 mm. long, 0.9-2.5 mm.
wide, usually trigonal, the dorsal surface rounded and the sides flat,
sometimes obovate, sometimes somewhat flattened, in one section the
seed may have an ascending membranous wing 0.3-5.3 mm. high
and 0.7-3.0 mm. wide.

GOLDBERG—GENUS MELOCHIA 211

The flowering and fruiting periods are usually long, sometimes
throughout the year.

Type spxcies: In the current International Code of Botanical
Nomenclature (33), it is agreed to associate generic names which
appear in Linnaeus’ Species Plantarum, edition 1, 1753, with the
first subsequent description given under those names in Linnaeus’
Genera Plantarum. In edition 5, 1754, of the latter, Linnaeus cited
Dillenius (9) under Melochia, and the latter only described Melochia
corchorifolia in Melochia. I therefore consider Melochia corchorifolia
L. to be the type species of the genus. The proposal of M. pyrami-
data L. as lectotype by Hitchcock and Green (International Botanical
Congress, 1930) has no official standing.

Disrrisution: In tropical and subtropical regions, from southern
United States to northern Argentina and Uruguay, and the West
Indies, India and southeast Asia, Indonesia, the Philippines, and other
Pacific Islands to the Tuamotu Islands, in savanna, thickets, fields,
or forests, sometimes on hillsides, usually in sun, in dry or wet habi-
tats, on various types of soil, at 0-3,150 meters elevation.

Key to Sections of Melochia

1. Fruit slightly retuse at the apex, stipe 4.3-6.7 mm. long; calyx accrescent,
becoming 15-36 mm. long; staminodes often present, up to 1.2 mm. long.
Section V. Physodium

1, Fruit not retuse at the apex, stipe 0-1.5 mm. long; calyx not accrescent or, if
accrescent, not exceeding 13 mm. in length; staminodes rarely present, to

0.4 mm. long.

2. Fruit pyramidal (ovoid in M. crenata), pteratus, loculicidally dehiscent along
the entire dorsal sutures (M. hermannioides tardily dehiscent), sometimes
also dehiscent along the ventral sutures and septicidal; inflorescences
usually leaf-opposed. .........2.. . . Section IV. Pyramis

2. Fruit subglobular or ovoid, sometimes acute at the apex, apterous but dorsum
of carpels sometimes keeled, dehiscence various; inflorescences axillary,
terminal or both (sometimes also leaf-opposed in M. degeneriana).

3. Fruit first loculicidally dehiscent all along the dorsal suture, usually later
septicidal; sinus between calyx teeth usually rounded to truncate,
rarely acute, teeth often distant; dorsum of carpels of fruit rounded.

Section III. Melochia

3. Fruit (1) septicidally dehiscent into cocci with a narrow ventral slit or
(2) loculicidal all along the ventral suture, the cells gaping ventrally,
and usually also 0.2-0.5 the way along the dorsal suture, sometimes
eventually septicidal; sinus between calyx teeth usually acute, some-
times narrowly rounded; dorsum of carpels of fruit often somewhat
keeled, especially at the distal end.

4. Fruit often falling apart, the dehiscence often completely septicidal,
carpels usually 5, rarely 6, seeds wingless ; peduncle sometimes absent,
usually without branches from it; herbs or shrubs, rarely a tree to 5
meters high. .........2.2... .Section II. Mougeotia

212 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

4. Fruit persistent, the dehiscence incompletely septicidal, carpels often
6-8, seeds often winged, the wing 0.3-5.3 mm. high, 0.7-3.0 mm.
wide; peduncle always present, usually branches from it; shrubs or
trees to 15 (-20) meters high ..... . . . Section I. Visenia

Key to Species of Melochia

SECTION I. VISENIA

1. Fruit suboblong, intercarpellary sulci extending the entire length of the fruit,
dehiscence usually only along the ventral sutures; stipules generally broad
(5-12 mm. wide), usually suborbicular, the apex rounded; seed wing large
(4-5.3 x 2-8 mm)... 6 ee ee . 1. M. umbellata

1. Fruit ovoid, intercarpellary sulci usually only marked in distal half of the fruit,
dehiscence partly along the dorsal sutures as well as along the ventral
sutures; stipules narrower (usually less than 4 mm. wide), deltoid, lanceolate
or ovate, the apex acute or acuminate; seed wingless or wing smaller (0.3-3.5
x 0.7-2.2 mm.).

2 Leaves lanceolate or elliptic, narrow, 1.1-2.4 (—4.5) em. wide, subcoriaceous;
shrub 1.5-2 meters high, . 2... ee ee ee M. grayana

2. Leaves mostly or all ovate, at least some 5 or more em. wide; plant often
more than 2 meters high.

3. Leaves persistently canescent, woolly on both surfaces, hairs to 1.2 mm.
long; young stems densely hairy, hairs to 2.0 mm. long; flowers in rather
compact cluster (i.e., pedicels short)... . . . .11.M. villosissima

3. Leaves glabrescent, hairs usually 0.5 mm. long or shorter (1 mm. long in

M. compacta) ; hairs on young stems 0.6 mm. long or shorter.
4. Seeds wingless or if wing present less than 1 mm. long and wide.

5. Inflorescences usually compact, primarily due to short pedicels
(mostly 0-2 mm. long); bracts relatively broad (0.8-3.0 mm. wide) ;
internodes short (0.5-1.0 cm. long); indument relatively long (to
1.0 mm.); in longistylous form of the flower the filaments united up
to the anthers or free for0.6mm. ... . . . .10. M. compacta

5. Inflorescences fairly loose, the pedicels to 7 mm. long in fruiting stage;
bracts narrower (0.5-1.7 mm. wide); internodes usually longer;
indument shorter (to 0.5 mm.); in longistylous form of the flower
the filaments free for 1.2-1.5 mm.

6. Calyx 5.8-8.6 mm. long; fruit 4-6 (-8) mm. long, sometimes without
an arista; bracts 3.7-5.5 mm. long; leaf serrations to 2.4 mm.
high; internodes 0.3-3.0 cm. long; seed not aristate.
2. M. odorata
6. Calyx 4.0-5.5 mm. long; fruit 6.2-10.5 mm. long, aristate; bracts 2
mm. long; leaf serrations to 1.5 mm. high; internodes 0.7-8.3
cm. long; seed sometimes aristate. . . . . . .5. M. aristata
4. Seeds winged, at least some wings 1 mm. or more long and wide (0.6-
3.5 x 0.7-2.2 mm.).

7. Inflorescences long (17.5-21.5 cm.); petioles long (2-19 em.); petals
salmon pink... 1 ee ee ee ee ee M. longepetiolata

7. Inflorescences shorter (3-14 em.); petioles shorter (0.3-9.5 cm.);
petals usually bright pink, yellow or white.

8. Petioles short (0.3-1.5em.) ......... .8M. roseiflora
8. Petioles generally longer (0.3-9.5 cm.).
9. Serrations of leaves up to 2.5 mm. high.

GOLDBERG—GENUS MELOCHIA 213

10. Fruit 4-6 mm. long, 4-6.5 (-8.0) mm. wide; calyx 5.8-8.6 mm,
long, 6.1-8.0 mm. wide; petals usually pink.
2. M. odorata
10. Fruit larger, 7-9 mm. long, 7.0-7.5 mm. wide; calyx generally
smaller 3.5-6.0 mm. long, 3.8-6.7 mm. wide; petals yellow
or white. . ... .... . .& M., vitiensis
9. Serrations of leaves up to 1. 1 mm. high.
11. Fruit 9-10 mm. long, 6-7.2 mm. wide; lower surface of leaf
subglabrous. . . . . . .6.M. degeneriana
11. Fruit smaller, relatively broader, ‘5-7.5 mm. long, 5.3-9.4
mm. wide; lower surface of leaf tomentulose.
7. M. mollipila

SECTION II. MOUGEOTIA

1. Primary dehiscence septicidal into cocci with a narrow slit along the ventral
suture, sometimes also for 0.33-0.5 the way along the dorsal suture, or
readily split for 0.25-0.5 the way along the dorsal suture.

2, Calyx considerably accrescent, becoming 7-12.3 mm. long, 7.7-12 mm. wide.
3. Calyx becoming 8-12.38 mm. long, and up to 12 mm. wide; peduncle
slender and short, typically 0.3-0.5 mm. wide, and 0.1-0. 7 cm. long,
often 2 or 3 per leaf axil; hairs not Prownish, predominantly simple
on the upper surface of the lamina. . . . . . 22. M. lupulina
3. Calyx becoming 7-9.5 mm. long, 7.7-10 mm. wide: peduncle relatively
thick and long, typically 0.5-1.5 mm. wide and 0. 5-8. 0 cm. long, usually
1 per leaf axil; hairs often brownish, stellate hairs predominant, or
as common as the simple ones, on the upper surface of the lamina;
lamina and stems more densely hairy than those of M. lupulina.
19. M. mollis
2. Calyx not accrescent or only slightly so, 3.1-7.0 mm. long, 3-6.5 mm. wide.
4. Inflorescences slender, open, somewhat paniculate; peduncles very slender,
0-2.0 em. long; pedicels slender and long, 3-7 mm. long; serrations of
lamina large, up to 3.5 mm. high; lamina not canescent or cinereous,
hairs simple and stellate on the upper surface . . . 24. M. bernoulliana
4. Inflorescences usually glomerate or corymbiform; peduncles sometimes
absent; pedicels shorter, typically 1-3 mm. long; serrations of lamina
shorter.

5. Inflorescences usually corymbiform, sometimes paniculate; young
stems and leaves canescent-cinereous, hairs stellate, 0.1-0.3 mm. long.
21. M. leucantha
5. Inflorescences neither corymbiform nor paniculate; young stems and
leaves usually not canescent-cinereous, simple and longer hairs usually

also present.
6. Internodes short, 0.4-0.7 cm. long; petioles short, 0.2-0.5 cm. long;

peduncles 1.2-1.6 cm. long, branched . . . . . 20. M. colombiana
6. Internodes and petioles usually longer than 1 cm.; peduncles often
absent.

7. Peduncles relatively thick, 0.5-1.5 mm. wide, and usually rela-
tively long, 0.5-8.0 cm. long, sometimes branched.

19. M. mollis

7. Peduncles slender, usually shorter and not branched, often absent.

8. Young stems predominantly covered with glandular hairs;

simple hairs on the upper surface of the lamina more or less

erect; peduncle absent. ...... . .27. M. tragiifolia

214 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

8. Young stems without glandular hairs or with very few; simple
hairs on the upper surface of the leaf appressed.
9. Peduncle 0-4.0 cm. long; pedicels 0.5-3 (-4) mm. long;
bracts narrow, 0.2-1.1 mm. wide. . . . 23. M. thymifolia
9. Peduncle 0-1.0 cm. long; pedicels usually shorter, 0-2 (-3) mm.
long, bracts narrow or wide.
10. Leaves predominantly lanceolate and acuminate, up to

3.9 em. wide. .... ... . . 28 M. tomentella
10. Leaves predominantly ovate, usually acute, up to 7.7 cm.
wide.

11. Bracts mostly broad, 0.6-3.9 mm. wide; lower surface
of leaf tomentulose; inflorescences commonly terminal
as well as axillary... .. . . .26. M. nudiflora

11. Bracts narrow, 0.2-1.1 mm. wide; lower surface of leaf
usually subglabrous; inflorescences glomerate, usually
of numerous flowers, at up to 20 axillary nodes,
usually not terminal too. . . . .25. M. nodiflora

1. Primary dehiscence of the fruit loculicidal along the ventral suture, so that
the cells are gaping ventrally, and 0.2-0.5 the way along the dorsal suture,
sometimes also finally septicidal.

12. Inflorescences corymbiform, open, fairly large, 6 cm. long, 5 cm. wide,
axillary; lamina large, 5-14 em. long, 2-8.2 em. wide; a tuft of hairs
0.1-0.2 mm. long at base of petal blade; calyx and corolla short (4.3-6.1
mm. long) . . ee .. . . 17. M. ulmifolia

12. Inflorescences not corymbiform, sometimes contracted, often smaller,
terminal, axillary, or both; lamina usually smaller, 0.8-9.5 (-12) cm.
long, 0.2-7.5 em. wide; no hairs at base of petal blade.

13. Inflorescence a dense terminal spike, sometimes also axillary, 4-5 cm.
long; calyx (7-8.3 mm.), petals (11.6-12 mm.), pistil in longistylous
form (11-12 mm.) and fruit (7 mm.) relatively long.

18. M. gardneri

13. Inflorescences axillary, or axillary and terminal, usually paniculate,
racemose, or umbellate; calyx, petals, pistil in longistylous form and
fruit often shorter.

14. Fruit large, 6.7-7.5 mm. long, 4.5-6.5 mm. wide; serrations of leaves
large, 0.7-3.6 mm. high; leaves subglabrous; stamens and pistils
subequal (4.8-5.6 and 4.2-5.1 mm., respectively).

16. M. argentina

14, Fruit smaller, 3.7-6.0 mm. long, (3-)3. 5-5.6 mm. wide; serrations of
leaves smaller, 0.3-2.7 mm. high; leaves sometimes quite hairy.

15. Inflorescences primarily or entirely axillary; lamina small, 1-4.3
(-6.5) em. long, 0.8-3 (—4.5) em. wide, usually broadly ovate
or suborbicular; plant procumbent or ascending, small, 0.1-0.5
meter long. .... .... .. . 14. M. chamaedrys

15. Inflorescences primarily terminal: lamina usually larger, usually
ovate or lanceolate; plant often erect and larger.

16. Fruits usually in pairs; leaves subglabrous; cay short, 3-4.7

mm. long... . ce ee . M. Kerriifolia

16. Fruits not in pairs; leaves “usually < siecous-pilose calyx usually
longer, (4-) 4.6-7.8 mm. long.

17. Lamina long-ciliate, hairs on surfaces also long, 1.3-4.0 mm.

long; peduncles 0.5-3.0 cm. long; pedicels 2.5-4.0 (-6) mm.

GOLDBERG—GENUS MELOCHIA 215

long; flowers evenly distributed about the inflorescence
branchlets; inflorescences up to 6.5 em. long.

13. M. hassleriana

17. Lamina not ciliate, hairs on surfaces up to 1.3 mm. long;

peduncles and pedicels usually shorter; flowers sometimes

more frequent on the upper side of the inflorescence branch-

lets; inflorescences to 20 (-31) em. long and usually flowers

more numerous. ............ 12. M. pilosa

SECTION III. MELOCHIA

1. Leaves linear, at least 10 times longer than wide, entire or obscurely serrate,
glabrous or subglabrous... . - ... . . 36. M. graminifolia
1, Leaves not linear, less than 10 times longer ‘than wide, serrations evident.

2. Leaves densely tomentulose, at least the lower surface canescent, all or
almost all hairs very short (0.1 mm.), stellate, the rays numerous (8-12
per hair) ; inflorescences axillary, pedunculate, umbellate . 34. M. arenosa

2. Leaves not tomentulose-canescent, or if so, inflorescences primarily terminal
or if axillary, not pedunculate.

3. Flowers not dimorphic, androccium and gynoecium equal in length;
calyx short, 1.5-2.6 mm. long.
4. Inflorescences principally or entirely terminal.
5. Inflorescences subcapitate, leaves mostly ovate . 39. M. corchorifolia
5. Inflorescences corymbiform, leaves lanceolate . . 38. M. lanceolata
4. Inflorescences principally or entirely axillary.

6. Fruit small, 2.4-2.5 mm. long, 3.2-3.3 mm. wide, with simple hairs;
inflorescences not pedunculate; lamina small, 0.7-3.5 em. long,
0.6-1.6 cm. wide. . . . . . . .33. M. melissifolia

6. Fruit larger, 2.7-4.0 mm. long, 3. 8-4, 8 mm. wide, with glandular and
simple hairs; inflorescences often pedunculate; lamina larger, 3-7.5
cm. long, 0.9-6.2 cm. wide. ...... .. .87. M. manducata

3. Flowers dimorphic on different plants, androecium and gynoecium differ-
ing in length; calyx usually longer, 2.4-6.0 mm. long.

7. Leaves lanceolate to suboblong, nearly linear near the apex of the stem,
a few scattered hairs on the upper surface; staminal tube glabrous;
peduncles 0.2-4.1 em. long... . - . .. .35. M. simplex
7. Leaves usually ovate, if primarily lanceolate, then densely tomentulose;
staminal tube usually puberulous; peduncles 0-0.1 cm. (rarely to 3. 0

cm.) long.

8. Plants small, usually prostrate, 0.2-0.6 meter long; lamina small,
1-2.2 cm. long, apex usually rounded (Isle of Pines and Pinar del
Rio, Cuba) . . . .. .. .30. M. savannarum

8. Plants usually larger, more : erect; ‘lamina usually larger, 1.5-10.5 cm.
long, apex usually acute.

9. Inflorescences primarily terminal. . .... . .29. M. villosa
9. Inflorescences primarily axillary.

10. Petiole short, 0.1-0.4 cm. long, base of lamina cuneate to
rounded, serrations short, 0.2-0.8 mm. high; pedicels absent
or very short, to 1 mm. long; calyx 5-5.2 mm. long; petals
narrow, 1.7-2.0 mm. wide. . ... . .31. M. splendens

10. Petiole longer, to 2.3 cm. long, base of lamina rounded to sub-
cordate, serrations higher, to 2.0 mm. high; pedicels present,
longer, to 3.5 mm. long; calyx shorter, less than 5 mm. long;
petals wider, to 3.5 mm. wide . . .. . . 32. M. bracteosa

216 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

SECTION IVY. PYRAMIS

1. Upper surface of leaves densely sericeous-tomentose, yellow-tan-canescent,
and lanate, hairs simple, appressed, 0.5-2.0 mm. long, lower surface densely
tomentose, yellow-tan-canescent, hairs stellate, forked and simple, 0.7-1.6
mm. long; peduncles relatively thick, 1.2 mm. wide. . . 48. M. lanata

1. Upper surface of leaves not densely sericeous-tomentose, nor yellow-tan-
canescent and lanate, hairs shorter and other types often also present, lower
surface often not densely tomentose, hairs shorter; peduncles thick or
slender.

2. Fruit large, 10-12 mm. long, 19-22 mm. wide, chartaceous, truncate or very
shortly rostrate (1 mm.) at the apex, alae relatively broad, and blunt at
the lateral angle. . 2... 2 eee ee ee ee ee 45. M. illicioides

2. Fruit smaller, 5-10.5 mm. long, 3.8-17 mm. wide, somewhat indurate, apex
usually acute, rostrum short or long, alae usually narrower, and acute,
acuminate or rounded at the lateral angle.

3. Fruit ovoid, pentapterous, 10-10.5 mm. long, 6.7-8.7 mm. wide, the
lateral angle of the alae obscure; pedicels long, 4-7 mm. long; lamina
oval to broadly ovate, the apex rounded... . . .50. M. crenata

3. Fruit pyramidal, pentapterous, often smaller, the lateral angle of the alae
evident; pedicels often shorter; lamina usually ovate or lanceolate,
the apex often acute.

4, Fruit narrow, both absolutely and relative to length, 6-7 mm. long,
3.8-4.8 mm. wide; entire plant canescent, tomentulose, hairs stellate,
0.3-1.0 mm. long; peduncles short and relatively thick, 0.2-0.8 cm.
long, 1.2-1.5 mm. wide; pedicels short, 1.5-2.0 mm. long.

44, M. werdermannii

4. Fruit usually broader, 4-14 mm. wide; plant often neither canescent nor
tomentulose, hairs sometimes simple, forked or glandular as well as
stellate; peduncles sometimes longer and slender; pedicels sometimes
longer, to 6 mm. long.

5. Plants prostrate, small, 0.1-0.5 meter long; lamina small, 0.9-2.1 cm.
long, 0.6-1.4 em. wide, usually rounded at both ends, sometimes
cuneate at the base, crenations relatively large, 0.5-1.9 mm. high,
not extending to the base of the lamina; fruit tardily dehiscent,
lateral angle of the alae above the middle of the fruit.

40. M. hermannioides

5. Plants usually erect, rarely decumbent, often larger; lamina usually
larger, often acute at the apex, crenations or serrations relstively
smaller, usually extending to the base of the lamina; fruit -eadily
dehiscent, lateral angle of the alae usually at, or below, the middle
of the fruit.

6. Lamina usually obovate or elliptic, rarely ovate, the base cuneate or
rounded, the apex rounded, relatively small, 0.6-4.5 cm. long,
0.3-2.8 em. wide; petiole short, 0.1-1.2 em. long; flowers usually
white. . 2... 0... ee ee ee ee ee eA. M. parvifolia

6. Lamina usually ovate or lanceolate, the base not cuneate, the apex
often acute, usually larger; petiole usually longer; flowers often
colored.

7. Fruit relatively short and narrow, 6-7 mm. long, 4-5.8 mm. wide;
peduncle often relatively long, up to 4 cm. long; calyx rela-
tively long, 6.7-10.2 mm. long... .. . .42.M. morongii

7. Fruit often longer and broader, typically 7—-8.5 (-10.5) mm. long

GOLDBERG—GENUS MELOCHIA 217

and 6.6-11 (-17) mm. wide; peduncle often shorter, typically
to 2.5 em. long; calyx usually shorter, typically 5-8.5 mm. long.
8. Peduncle short, typically 0-0.5 cm. long; pedicels short, 1-2

mm. long; fruit relatively narrow, (5.5-) 6.2-7.0 mm. wide;

serrations of lamina fairly large, to 2.5 mm. high, hairs of

lamina to 1.2mm.long. . . - . . . . 43. M. anomala
8. Peduncle usually longer, typically 1-2 cm. long; pedicels

usually 1-4 mm. long; fruit wider, typically 8 (-17) mm.

wide; serrations of lamina usually smaller, hairs of lamina

often shorter.

9. Peduncles slender, 0.4-0.6 mm. wide; inflorescences um-
bellate, typically 2-10 flowers per inflorescence; flowers
usually purple, pink, violet, blue, or purple distally and
yellow proximally.

10. Inflorescences mostly leaf-opposed; leaves usually
sparsely pubescent, hairs simple, forked, stellate and
sometimes also glandular; fruit often purple-blotched,
rostrum 0-2.1 mm. long, point of alae acute or acumi-
nate; plant 0.08-2.0 meters high; calyx 3.7-6.2 mm.
long. ... . . . .51. M. pyramidata

10. Inflorescences mostly “axillary; leaves often densely
tomentose and canescent, hairs stellate; fruit not
purple-blotched, rostrum 1.3-5.0 mm. long, point of
alae acute or rounded; plant 0.3-4.0 meters high;
calyx 5-8.6 mm. long. . . . . . . 49. M. tomentosa

9. Peduncles relatively thick, 0.8-1.4 mm. wide; inflorescences
subcapitate, typically 3-20 flowers per inflorescence;
flowers usually white or yellow, rarely purple.

11. Peduncle usually axillary, 0.8-2.9 em. long; fruit 11.5-17
mm. wide; lower surface of lamina sometimes canescent
(Brazil). . . . . . . .46. M. betonicifolia

11. Peduncle often leaf-opposed, 0.4-2.0 cm. long; fruit
6-11 mm. wide; lamina generally more densely
pubescent and hairs often yellowish (mostly Vene-
zuela and Colombia). . . .. . .47. M. caracasana

SECTION V. PHYSODIUM

1. Petals 16-17.4 mm. long, 5.6-9.5 mm. wide, markedly exceeding the calyx
in length at anthesis; hairs of calyx stellate, 0.2-0.5 mm. long; staminodes
none orrarely 0.1 mm.long ............ 53M. rhodocalyx

1. Petals 8-12 mm. long, 2-2.7 mm. wide, not exceeding or slightly exceeding
the calyx in length at anthesis; hairs of calyx stellate or glandular; staminodes
none or to 1.2 mm. long.

2. Calyx and pedicels covered with glandular hairs 0.2-0.6 mm. long and a few
stellate hairs 0.1-0.2 mm. long; petals 8-12 mm. long, 2-2.7 mm. wide,

not exceeding the calyx in length at anthesis; staminodes 0.2-1.2 mm.
long .... .... . 52. M. adenodes

2. Calyx and pedicels covered with stellate hairs 0.1-0.8 mm. long; petals 9.7
mm. long, 2.1 mm. wide, slightly exceeding the calyx at anthesis; stami-
mnodesnone...............4. 4... 54M. corymbosa

218 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Section I. Visenia K. Schum. in Engler & Prantl, Pfizfam. HI
6: 81. 1890

Shrubs or small trees 0.7-15 (-20) meters high, the young stems
and petioles tomentulose or tomentose, hairs fuscous or cinerescent,
stellate, and often also forked and simple, 0.05-2.0 mm. long, the
internodes 0.2-18.5 cm. long; stipules often soon deciduous, 2-10
mm. long, 0.5-12 mm. wide, deltoid-acute or deltoid-acuminate,
lanceolate, ovate or suborbicular, tomentulose, hirsute, sericeous-
tomentose, ciliate or subglabrous, hairs stellate, forked or simple,
0.05-2.0 mm. long; petiole 0.3-14.5 (-20.5) em. long.

Lamina 3.1-24 (-30) em. long, 1.1-21.5 (-27) cm. wide, usually
broadly ovate, sometimes lanceolate, rarely elliptic, the base rounded
to cordate, rarely cuneate or truncate, the apex acute, rarely acuminate
or rounded, the upper surface tomentulose, sericeous-tomentulose,
canescent and woolly or velvety to the touch, or glabrous, hairs
stellate and usually also forked and simple, 0.05-1.2 mm. long, some-
times also minute resin-colored granules scattered over the surface,
the lower surface with similar indument but never completely glabrous,
the margin usually irregularly crenate to serrate, crenations or serra-
tions 0.9-8.3 mm. wide, 0.3-3.0 mm. high, 5-10 pairs of primary veins,
1-3 are basal.

Inflorescences loose or compact, 2.5-21.5 (-39) em. long, in axils
of 1-6 upper leaves, sometimes apparently also terminal, rarely leaf-
opposed or 90° from a leaf axil, or not associated with a leaf, corymbi-
form, branched, the ultimate divisions umbellate, the umbels 2-5-
flowered, the flowers numerous, moderate or few, sometimes fragrant,
the peduncle 2-11 (-16) cm. long, usually with a primary branch
0.4-5.2 (-7.0) em. long, often with a secondary one 0.3-2.0 (4.0) cm.
long, and sometimes with a tertiary one 0.5-1.6 (—3.0) em. long, and
rarely with a quaternary one to 2 cm. long, the pedicel 0-7 mm. long;
bracts sometimes soon deciduous, rarely 2 united, 1.0—5.5 (-10) mm.
long, 0.1-3.0 (-14) mm. wide, ovate, lanceolate, or deltoid, rarely
linear, apex usually acute, sometimes rounded, tomentulose, hirsute
or ciliate, hairs stellate, sometimes also forked and simple, 0.1-0.4
(-1.3) mm. long.

Calyx sometimes orange to bright red, pinkish brown or yellow
green, 3.5-9.0 mm. long, 3.8-10 mm. wide, tomentulose, sometimes
also sericeous, hairs stellate, sometimes also forked, simple, or glandu-
lar, 0.05-0.5 (-1.5) mm. long, the teeth 1.2-5.5 mm. long, 1.38-4.8
mm. wide, deltoid to ovate, acute or acuminate, the sinus acute to
narrowly rounded; petals usually pink to red, sometimes white to
yellow, rarely pale blue, pale purple, violet, or lilac with a proximal
orange spot, 5.0-9.0 (-15.5) mm. long, 1.6-5.0 (-7.5) mm. wide,
obovate, oblanceolate, elliptic, oval, ovate or oblong, the apex usually

GOLDBERG—GENUS MELOCHIA 219

rounded, rarely emarginate, the base acute, attached to stamens for
0.8-3.0 mm., the claw 0.5-3.0 mm. long; sometimes glandular tissue
present with lobes extending between the bases of the petals.

Longistylous form. Stamens 4-6 mm. long, the filaments often
yellow, sometimes rose, united up to the anthers or free for 0.2-2.0
mm., the anthers yellow, 1.2-2.2 mm, long, 0.6-1.4 mm. wide, oblong,
emarginate at both ends, usually deeply so at the apex; pistil 5.2-9.0
mm. long, the styles filiform, often slightly clavate, papillose for
0.6-2.0 mm. at the apex, free for 2.9-6.0 mm., to the ovary, or united
for 0.5-1.0 mm., the ovary ovoid or elliptic, the ovary and base of
styles usually densely sericeous, sometimes tomentose or hirsute,
hairs simple, forked, and sometimes also stellate, 0.5-1.8 mm. long,
the carpels sometimes 6-8, stipe none.

Brevistylous form. Stamens 4.7-8.1 mm. long, the filaments free
for 1.8-5.0 mm., the anthers 1.0-2.5 mm. long, 0.7-1.4 mm. wide,
rarely staminodes present, 0.1-0.2 mm. long; pistil 3.6-5.3 mm. long,
the styles filiform, papillose for 0.6-0.8 mm., free for 1.4-2.6 mm.,
usually to the ovary, sometimes united for 1 mm., the ovary globular
or ovoid, usually densely sericeous, sometimes sericeous-tomentose
or hirsute, hairs simple and forked, 0.7-1.7 mm. long, the carpels
sometimes 6-8, stipe none.

Fruit a green, brown or purplish-red capsule, ovoid or oblong,
4-10.5 mm. long, of which the arista is 0-2.0 (-3.0) mm. long, 4—9.4
mm. wide, the dorsum of carpels obtuse or subacute, the sulci between
the carpels in the distal half of the fruit or extending for its entire
length, shallow or deep, extending a fifth to half way to the center of
the fruit, tomentose, tomentulose, or sericeous, hairs simple, forked,
and stellate, the last short, 0.1-1.6 mm. long, the stipe (base of
capsule) 0-1 mm. long; dehiscence all along the ventral sutures,
sometimes also a fifth to half way along the dorsal sutures and partly
septicidal; seeds 1 or 2 per cell, the body brown, trigonal, 1.8-3.5
mm. long, 1.1-2.0 mm. wide, sometimes testa loosened for 0.2-0.5
mm., or with an arista 0.7 mm. long, or with an ascending wing
0.3-5.3 mm. high, 0.7-3.0 mm. wide, obliquely deltoid or adaxial
edge straight and abaxial edge convex, the apex rounded or acute.

A few species produce flowers and fruit throughout the year,
others have a restricted flowering and fruiting period.

Type: Melochia umbellata (Houtt.) Stapf.

Disrrizution: India and Southeast Asia, Indonesia, the Philip-
pines, and other Pacific islands to the Tuamotu Islands, in forests,
thickets, and disturbed areas, under wet or dry conditions, on soils
or rocks of calcareous or volcanic origin, at 0-1,400 (-1,700) meters
elevation.

920) CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

1. Melochia umbellata (Houtt.) Stapf, Kew Bull. 317, 1913.
Visenia umbellata Houtt. Handleidung 8:309. 1777.
Wisenia indica Gmel. Syst. 2:515. 1791.
Riedleia tiliaefolia DC. Prod. 1:491. 1824.
Riedleia velutina DC. Prod. 1:491. 1824.
Aleurodendron album Reinw. Sylloge Pl. Ratisbonn, 2:12, 1828.
Glossospermum velutinum (DC.) Wall. Cat. n. 1153. 1829.
Pentaglottis tomentosa Wall. Cat. n. 1156. 1829.
?Glossospermum ?cordatum Wall. Cat. n. 1195, 1829.
Melochia arborea Blanco, FI. Filip. 524. 1837.
Hypericum pentandrum Blanco, Fl. Filip. 614. 1837.
Visenia Javanica Jungh. in Hoev. & DeVriese, Tijdschr. 7:302. 1840.
Riedleia Aleurodendron (Reinw.) Steud. Nom. ed, 2, 2:459. 1841.
Riedleia cordata (Wall.) Steud. Nom. ed. 2, 2:459. 1841.
Riedleia Wallichiana (Wall.) Steud. Nom. ed. 2, 2:460. 1841.
Visenia alba (Reinw.) Endl. ex Walp. Rep. 1:351. 1842,
Visenia tomentosa Miq. Anal. Bot. Ind. 3:4. 1852.
Visenia velutina (DC.) Turez. Bull. Soc. Nat. Mose. 31(1):212. 1858.
Melochia velutina Wall. ex. Bedd. For. Man. Bot. 35. 1869 or 1871.
Melochia indica Kurz, Prelim. Rep. For. Veg. Pegu, App. A, 28. 1875.
Melochia indica var. deglabrata Kds. & Val. Bijdragen tot de Kennis der
Boomsoorten van Java 2:194. 1895.
Melochia odorata var. Schlechteri Mildb. Engl. Bot. Jahrb. 62:350. 1929.

Usually a small tree, sometimes a shrub, 2-15 (-20) meters high,
4-15 (-35) cm. in diameter, the ramification ascending to horizontal,
the bark about 4 mm. thick, rather smooth to rough, pale brown to
dark grey brown, with some longitudinal ridges, the wood straw, the
young stems and petioles woolly to the touch, tomentulose, hairs
stellate, 0.1-1.0 mm. long, often yellowish brown at base, the inter-
nodes 1-7 em. long; stipules deciduous, 4.3-10 mm. long, (1.1-) 5-12
mm. wide, ovate to suborbicular, the apex usually rounded, covered
with simple, forked and stellate hairs 0.1-0.5 mm. long; petiole
2.5-11 (-20.5) cm. long.

Lamina 6.3-14.8 (30) em. long, (2-) 3.8-14 (-27) cm. wide, usually
broadly ovate, the base rounded to cordate, the apex acute, often
lanceolate and much reduced just below the inflorescence, both
surfaces of young leaves covered with stellate hairs, rarely also
simple and forked, 0.1-0.5 mm. long, woolly or velvety to the touch,
larger hairs sometimes yellow brown at base, often minute resin-
colored granules scattered on surface, young leaves somewhat canes-
cent, older ones glabrescent, the margin irregularly crenate-serrate,
serrations 1.8-6.0 mm. wide, 0.5-3.0 mm. high, 7-10 pairs of primary
veins, 2 or 3 are basal.

Inflorescences fairly loose, in axils of upper 3 or 4 or more leaves,
3.7-16 (-39) em. long, corymbiform, the flower number moderate, the
ultimate divisions umbellate, the umbels 3-5-flowered, the peduncle
(2.8-) 4-6.5 (-11) em. long, with a primary branch 0.5-2 (-5.5) cm.

GOLDBERG—GENUS MELOCHIA 221

long, a secondary one 0.5-2(-4) cm. long, sometimes a tertiary one
0.5-1.3 (—3) cm. long, rarely a quaternary one to 2 em. long, the
pedicels 1-3 mm. long; bracts (1-) 2.5-5.3 (-10) mm. long, 0.7-2.3
(-14) mm. wide, ovate to lanceolate, the apex rounded or acute,
covered with stellate hairs 0.1-0.3 mm. long.

Calyx yellow green to pinkish brown, 4.8-6.2 (-7.0) mm. long,
4-5.7 mm. wide, covered with stellate hairs 0.1 mm. long, and less
numerous simple, forked, and stellate hairs to 0.6 mm. long, the teeth
2—3.4 mm. long, 1.4~3.3 mm. wide, deltoid to ovate, acute or slightly
acuminate, the sinus acute; petals usually pink (light pink to red),
sometimes white to yellow and sometimes orange proximally, rarely
pale blue, pale purple or violet, (5.5-) 6-7.2 mm. long, (1.9-) 2.2-3.7
mm. wide, obovate, oblanceolate, or elliptic, attached to the stamens
for 1-2 mm., the claw 0.5-1.5 mm. long, usually glandular tissue
present with lobes extending between the bases of the petals; fila-
ments and styles cream, anthers yellow.

Longistylous form. Stamens 4-4.9 mm. long, the filaments united
up to the anthers or free for 0.2-0.9 mm., the anthers 1.2-1.6 mm.
long, 0.8-1.0 mm. wide, oblong, slightly narrower at apex, emarginate
at both ends, almost to the middle from the apex; pistil (6.2—) 6.6—-7.6
mm. long, the styles free for (3.5-) 4.2-5.2 mm., united for 0.5 mm.,
papillose and clavate for (1—) 1.3-1.7 mm. at the apex, the base
hirsute, the ovary ovoid, pentagonal, densely hirsute or sericeous,
hairs forked, simple, and stellate, 0.5-1.3 mm. long, stipe none.

Brevistylous form. Stamens 5.5-6.4 mm. long, the filaments free
for 3.3-5.0 mm., the anthers 1-1.2 mm. long, 0.7-1.0 mm. wide;
pistil 4.6-5.2 mm. long, the styles free for 1.8-2.5 mm., nearly to the
ovary, papillose for 0.6 mm., the ovary densely hirsute, hairs mostly
forked and simple, to 1 mm. long, stipe none.

Fruit green, brown or purplish red, 8-9.5 mm. long, 6.3-7.5 mm.
wide, oblongish, only slightly narrower at the apex, the sulci between
the carpels deep, extending nearly half way to the center of the fruit
at both apex and base, the sides of the carpels cohering only for a
very short distance at the base, covered with stellate hairs 0.1 mm.
long and simple, forked, and stellate hairs 0.5-1.0 mm. long, the
stipe 0-0.5 mm. long; dehiscence along the ventral sutures; seeds
winged, the body brown, 2-3.3 mm. long, 1.5-2.0 mm. wide, the wing
ascending, 4—5.3 mm. high, 2-3 mm. wide.

Flowering and fruiting specimens collected throughout the year,
peak of flowering May—August, fruiting June-August.

Type Locauity: East Indies, probably Java.

DistriBuTiIon: Indigenous from India to New Guinea, often
common in secondary thickets, in recent clearings, along the edges

227-051—67—_3

992 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

of forests and rivers, often in red calcareous soil, often in localities
affected by monsoons, at 0-1,400 (-1,700) meters elevation.

INDIA: Terr. Canara (KANaARA): Mercara and Dharwar, ? Metz 623a (5).
Car Nicobar, Prain March 1891 (US).

INDOCHINA: Avustra-CHOcCHINCHINAE: Dinh ad Baria gallicae, Pierre 544
(L, ?US). Tonkin: Rive droite de la rivitre Novie entre C. Thap. et les roches
de Notre-Dame, Balansa 3719 (L). ANNAM: Dabac, Hout Donai, Poilane,
s.n. (8).

MALAYA: Perak: Dr. King’s Collector 8401 (L). Kampong Tembok, 3 mi.
N. of Ipoh, Sinclair 9854 (L). ?PeRraK: Father Scortechinis.n. (US). SELANGOR:
Kepong Plantation, Symington Kepong Field No. 32872 (L). Pawana: Sabai
Estate, Bentong, Kemanshul Div., Shah 258 (L).

Inv. Or. Hers. Wallich 1156 (L). Isotype of Pentaglottis tomentosa Wall.

SUNDA ISLANDS: Sumatra: Singarong agong, Robinson & Kloss May 24,
1914 (US) ?Indrapoera, Korthals s.n. (L). Malintang, Bunnemeijer 3793 (L).
SIBOLANGIT: Karohoogvl by Sarinembah, Galoengi 16 (L). BERKOELEN: Kebana-
goeng, de Voogd 1200 (L). Djambi. 8. Lesing by Pahoe, Posthumus 1015 (L),
1059 (L). TAPIANOELI: TOBA: Toetoepan, Boeea 5978 (L). Toba, Duwchond
252 (L). Near Taroentoeng, Bartlett 8367 (L, US). Palembang, Bamas, Grashoff
170 (L), 1091 (L). ?Pace Palembang, Forbes 2546 (L). N. Sumarra: Deli,
Padang and Bedagei to Bila, Karohochebene, bei Jongging, am NW. Toba See,
Lérzing ?1758a (L), 7987 (L), 8075 (L), 15058 (L), 15965 (L).. West coast:
Koerintjo Bt. te Bakar, Binnemeyer 8174 (L). CENTRAL Sumatra: Mt. Sago
near Pajakumbuh, Maradjo (ass. W. Meijer) 14 (L). East coast: Padang,
Stomps 11-10-1923 (L). asanan: Peddaloeng, Bartlett & LaRue 136 (L), 139
(L). Hoeta Padang, Asahan, Krukoff 4421 (L, US). Adian Rindang, Asahan,
Boeea 8726 (L). Vicinity of Loemban Ria, Asahan, Hoeta Bagasan, Boeea 6863
(S, US), 7287 (US), 7530 (S), 7902 (S, US). LABOEHAN BATOE: Kota Pinang,
Langga Pajoeng on the Soengei Kanan, Toroes 3636 (L, 8, US). Si Mandi Angin
on the Soengei Kanan, Toroes 3957 (US). Kraxatau (Krakatoa): Brenlage
s.n. (L). P. Rakata, SE. side, v. Borssum Woolkes 1024 (L). Verlaten Hiland,
Backer s.n. (L). Java: ?Waittzs.n. (L). Junghuhn 408 (L). Zollingero 34 (L).
H. Lugo Batar s.n. (8S). ?Reinwardt 2350 (8). East Java, Reinwardt s.n. (8).
Schierbrand s.n. (W). Tjibodas, aap 766 (L). G. Malabar Poentjak gede,
Monterie 40 (L). Montis Sebu, Sort, Junghuhn s.n. (L). Bagelen, Vorderman
s.n. (L). Bandoeng, Popta 542/98 (L). Meester Cornelis, Backer s.n. (L)
Solak, Méller s.n. (S). Buitenzorg and vic., Danser 5573 (L), Sutrisno 25 (L),
Schuurman 33 (L), Soegandiredja 233 (L), Durand 818 (LL). BANTAM: TJIMARA:
Oedjoeng, Koorders, 7744 g* (L). G. Pangissan, Koorders 7744 6 (L). G.
Karang, Koorders 7743 8 (L). Batavia, Koorders 30901 6 (L), Backer 88 (L).
Res. Batavia G. Salak, Koorders 24411 8 (L). Depok, Hallier Aug. 24, 1896.
BANEAN: Tandjoeng Tajar, Karta 148 (L). Soerabaja, Doyelt 383 (L), 786 (L),
950 (L). Paroeroean, Backer 7810 (L), 37444 (L). P. Purmerend, middle of the
island, den Hoed 3063 (L). Japara, Soemanding, Neth. Ind. For. Service Ja: 3724
(L). maprioEN: Pandan, Kendeng, Trinil, Elbert 447 (L), 506 (LL). Lawoe,
Djogolaranga, Oosbooch, Kali Djeplog, Hilbert 278 (L). Vak 112 houtrestery
Toeloeng, Wind 6289 (L). G. Wilis Ngebel, Koorders 7746 8 (L). PROV.
PREANGER: Ploem s.n. (L). Takoka, Koorders 15155 6 (L). sEMARANG: Ke-
doengdjati, Koorders 7740 8 (L), 7745 B (L), 7748 B (L), 7749 6 (L). Pangent-
jongan, Koorders 7742 B (L), 14024 6(L). Palaboean ratoe, Koorders 7738 6 (L),
7739 B (L), 7741 6 (L). Telomojo, Koorders 27970 8 (L). Karangasem, Koorders
7747 6 (L). Tjiratjap Djampang koelon, Backer 17394 (L). Reg. Tjadas

GOLDBERG—GENUS MELOCHIA 223

Malang bij Tjidadap. L. v. Tjibeber Pasir Anger, Winckel 113 6 (L), 161 (L),
1356 6 (L), Bat. huizen v/d Brink 99a (L). Tjatjaban, and in the ravine of the
river Tji Taroem, Hochreutner 1667 (L). Res. Semarang. Telomojo, Koorders
27970 6 (L). Res. Kedoe G. Sendoro, Koorders 11354 6 (L). Res. Besoeki
Poeger, Banjoewangi, Koorders 7753 6 (L), 7756 8 (L). Prov. Banjumas. Prin-
gambo, Koorders 7752 8 (L), Koorders 37048 6 (L). Res. Besoeki Rogodjampi,
Koorders 29088 8 (L). Res. Kediri. N. v. Tritik grens Kediri-Rembang, Thorenaar
102 (L). Pekalongan. Soebah, Koorders 11608 6 (L), 11609 8 (L). Southeast
Java, Forbes 1343 (L). Centrau Java: G. Muria, Tjollo, N. of Kudus, Koster-
mans 6264 (K, L). West Java: 18 km. 8. of Rangkasbitung, Buwalda 2855 (L).
?Java, Tjyiomas 28/11/88 (L). Borneo: Winkler 2197 (L). ?G. Kunepay,
Hallier 1939 (L). ?Dversin, Korthals s.n. (L). Rajang, Sibu Kalong, Haviland
2836 (L). Nortu Borneo: Kinabalu Mt., Puasa N. Borneo Forestry Dept. No.
1541 (L). Tambunan, Wyatt-Smith Kepongfield No. 80412 (L, US). Kampong
Kinasaraban, Sinclair et al. 8994 (L). Cx LesBus: de Vriese & Tetjsmanns.n. (L).
TUKANG-BESI ISLANDS: Biriongko, Elbert 2588 (6083) (L). Kp. Palima, Noerkas
265 (L). Bonto Djai, Rachmat Exp. v. Vuuren 29 (L). Malili, Thawata,
Boschproefstation #Cel/V—164 (L). Rante Lemo, Pare-Pare, Kjellberg 3827 (S),
3827a (S). Labibia, Kjellberg 1363 (S). Makale, Kjellberg 1656 (L, S). G.
Bontham, Bunnemeyer 12354 (L). Minahassa, Koorders 18091 6 (L), 19718 8 (L);
Menado, Koorders 18089 (L). Sourueast: Landschaft Lankape, Elbert 7420
(3029) (L). Landschaft Rumbia, Hilbert 7528 (3071) (L). Burton Isuanp:
Lipumangau, Elbert 6502 (2745) (L). Limbo, Elbert 6616 (2785) (L). Mona
(Muna) Istanp: Raha, Elbert 7086 (2904) (L), Boschproefstation 1 1 5848 (L).
Tanette, Bunnemeyer 11760 (L). SW. CeLeses: Kamp: Biroro by Lombasang,
Biinnemeyer 11646 (L), 11456 (L). Lompox: Rindjani-Vulkangebirge, N. side
Labuan tjarik, Elbert 235 (535) (L). SW. Lomsor: Plambi, Elbert 5763 (2411)
(L). SumsBa (SoznmBa): Maomarroe, Iboet 486 (L), 538 (L), Lologooh, Iboet 278
(L). WexstSumsa: Waikalubah, Monad de Froidville 1869 (L). FLorxs: Endeh,
Jaag 1806 (L). Aor: Central highland, Lantoka-Pido, Jaag 1245 (L). Timor:
Aspecimen ex Herb. Mus. Paris, labeled Riedleia tiliaefolia (US 13124). Zippelius
s.n. (L). En Onderh. P. Alor, Bowman-Houtman 159 (L). <A. de Jussteu missit
1834 (8S), ?M. umbellata or M. odorata. E. Timor: Plateau of Baucau, Van
Steenis 18048 (L).

PHILIPPINES: Hallier 347a (L). Pautawan: Fanabay, Manalo 3 (Forestry
Bur. #7426) (US). Tawrrawt: Sulu Prov., Ramos & Edafio Bur. Sci. #44111 (L).
MINDANAO: ZAMBOANGA: Tetuan, Ahern 48 (US). pavao: Todaya (Mt. Apo),
Elmer 10864 (L, US), Clemens Bur Sci. #15581 (L); Santa Cruz, Williams 2799
(US). AGusaN: Cabadbaran (Mt. Urdaneta), Elmer 13651 (L). Surigao, Ahern
340 (US). NxEGROS: NEGROS ORIENTAL: Dumaguete (Cuernos Mts.), Elmer 9493
(L, US). Srarcao: Ramos & Pascasio, Bur. Sci. #34853 (L). Leyte: Wenzel
1352 (L). Samar: Ramos Bur. Sci. #17594 (US). Cunion: Merrill 619 (US).
Minporo: Baco River, McGregor 299 (US). Calapan, Escritor Bur. Sci. #21269
(L, US). Mt. Yagaw, SE. slope, Conklin 269 (L, US). East Mindoro, 18 km.
S. of Puerto Galena, San Teodoro, Vera Santos 5291 (L, US). Luzon: Morong,
Tanay, Merrill 2286 (US). sorsogon: Irosin, Mt. Bangko, Edafio & Gutierrez
161 (L); Mt. Bulusan, Elmer 16505 (L, 8, US); Patag, Sinclair & Edafto 9613 (L).
Batangas, Ramos & Deroy Bur Sci. #22665 (US). Laguna: Los Bafios, Quisum-
bing 466 (US), Ramundo s.n. (S); Mt. Maquiling, Elmer 17920 (L, 8, US), Mendoza
Phil. Nat. Herb. 12201 (US). Rizau: Ramos 1086 (US); Ahern’s Collector For-
estry Bur. #3152 (US); Manila, Navaliches, Loher 155 (US); Montalban, Loher
5574 (US); Bosoboso, Merrill 2829 (US), 2840 (US). Bataan: Dinalupijan,
Merrill 1476 (US). zamBALeEs: Subig, Hallier 4347 (L). TayaBas: Casiguran,

224 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Mearns Bur. Sci. #2972 (US). BENGUET: Sablan, Elmer 6086 (US). CAGAYAN:
Littoc, Lagum, vic. of Pefiablanca, Adduru 161 (US), 228 (US).

MOLUCCAS: Tataup: Karakelong, Lam 2570 (L). Mororar: Kostermans
1541 (L). Haumauera: Idjan & Moehtar 365 (L). Kmp. Goal., Pleyte 157 (L).
S. peninsula, Anang 553 (L). TErnate: Kabora, Anang 135 (L). Foramadiahi,
Beguin 1103 (L), 1594 (L). Lagoena, Beguin 628 (L). Dessa Takome, Bish.
23 (L). East coast, Pleyte 82 (L). Sorna (Suna): Sanana, Tandjong Baliha,
Bloembergen 4367 (L). Mangoli Island and N. of Mangoli, Neth. Ind. For. Serv.
bb. 29860 (L). Buru (Borroe): de Vriese Feb. 1860? (L). AmBorna (AMBON):
Poka, exp. v. Hulstyn, Saanan 105 (L). Kar (Ker): Ohoitiel bij Toeal, Jensen
312 (L). Aros (Aru): P. Trangan, KP. Ngaibor, Buwalda 5334 (L). TANIMBAR:
Neth. Ind. For. Serv. bb 24350 (L). Basar: puLaAu wotTaNn: E. Herloh, van
Borssum Waalkos 3028 (L).

NEW GUINEA: Near Kp. Saban (Wentholt exp.), Anta 261 (L). Jappen-
Biak, Mentembae, Aet & Idjan 104 (L). Japen: Seroei, Neth. Ind. For. Serv.
bb. 30566 (L). Rapsan Ampat: Batanta Island, W. of Sorong, Marchesa Bay,
Amdoei village, van Royen, 3533 (L). Sorong, Misool, W. of Fakal, bank of
Kasim River, Pleyte 1128 (L). NHollandia (Hamadi) Onderafd. Hollandia,
Kalkman #BW3677 (L). Stephansort, (?M. uwmbellata, fruit immature), Lew-
andowsky 34 (lL, 8). Papua: Schlechter 16143 (L, S), isotypes of M. odorata var.
Schlecteri Mildb.; Koitaki, Carr 12672 (L); Lake Daviumbu, Middle Fly River,
Brass 7734 (L); Northern Dist., ca. 3 mi. from Aku on Kuruaku tract Tufi subdist.,
Saunders 30 (K, L); MILNE BAY DistT.: Cape Vogel peninsula near Tapio village,
Hoogland 4288 (A, L).

HAWAII: Oanu: Kahuku mauka, Lam 7285a (L), Degener et al. 19548 (L, 8S,
US).

This species can be distinguished from the other species in the section
Visenia by its oblong fruit with intercarpellary sulci extending its
entire length, large suborbicular stipules, and relatively large-winged
seeds.

2. Melochia odorata L. f. Supp. Plant. 302. 1781.

A shrub or spreading, umbrella-crowned tree (1.2—) 3.3-15 meters
high, 9-10 cm. in diameter, the young stems and petioles rather
densely puberulous, hairs stellate and forked, 0.1-0.5 mm. long, the
internodes 0.3-3.0 cm. long; stipules deciduous, 2.5-4.0 (-9.0) mm.
long, 0.8-1.1 (-4.0) mm. wide, deltoid-acute or deltoid-acuminate,
covered with forked, stellate, and simple hairs 0.1-0.3 mm. long;
petiole 1.0—9.0 cm. long.

Lamina 4—20.7 cm. long, 2-18.7 cm. wide, usually ovate, the base
rounded to cordate or truncate, the apex acute to acuminate, leaves
subtending the inflorescence sometimes lanceolate-ovate, the base
cuneate to rounded, and at the lower end of the range in size, both sur-
faces of young leaves silvery to greenish- or tan-canescent, densely
puberulous, sometimes rough, hairs stellate, forked, and simple, 0.1-0.5
mm. long, soon glabrate, a few scattered hairs remaining, especially
on the veins, the margin irregularly crenate-serrate, serrations 1.7—-6.6
mm. wide, 0.5—-2.4 mm. high, 6-10 pairs of primary veins, usually 2
(1-3) are basal.

GOLDBERG—GENUS MELOCHIA 225

Inflorescences usually loose, 2.5-7.0 (-14) em. long, in axils of upper
2—4 leaves, corymbiform, the flower number moderate, the ultimate
divisions umbellate, the umbels 3-5-flowered, the flowers scented, the
peduncle 2.5-4.5 (-13) cm. long, with a primary branch 0.8-1 cm. long,
and sometimes a secondary one 0.5-0.8 cm. long, the pedicels (1-)
2-7 mm. long; bracts 3.7-5.5 mm. long, 0.5-1.7 mm. wide, lanceolate,
covered with stellate and forked hairs 0.1-0.4 mm. long.

Calyx pink or pale purple, 5.8-8.6 mm. long, 6.1-8.0 mm. wide,
densely covered with forked, stellate, and simple hairs 0.1-0.5 mm.
long, the teeth 3.7-5.0 mm. long, 2.5-3.0 mm. wide, deltoid or ovate,
acuminate, the sinus acute; petals usually pink, rarely pale lilac, pale
lilac with a proximal orange spot, cream with yellow proximally, or
white, 7.8-12.5 mm. long, 3.0-4.2 mm. wide, ovobate, the claw 1.3-1.7
mm. long, attached to the stamens for 1.4-2.5 mm., undulating ring
of glandular tissue 0.2 mm. wide between the calyx and corolla
ascending between the petals and descending at their bases.

Longistylous form. Stamens 5-5.5 mm. long, the filaments free
for 1.3-1.5 mm., the anthers 1.5-1.7 mm. long, 1.0-1.2 mm. wide,
oblong, usually emarginate at both ends, usually deeply so at the
apex; pistil 7.5-8.2 mm. long, the styles 4.5-5.7 mm. long, free to
the ovary, papillose and clavate at the apex for 1-1.5 mm., the ovary
elliptical to ovoid, densely sericeous, hairs 0.5-1.5 mm. long, the
carpels sometimes 7 or 8, stipe none.

Brevistylous form. Stamens 6.5-6.8 mm. long, ?staminodes 0.1
mm. long sometimes present, the filaments free for 3.7 mm., the
anthers 1.5-1.6 mm. long, 0.8-0.9 mm. wide; pistil 4.5 mm. long, the
styles 2.5 mm. long, free to the ovary, the carpels sometimes 6 or 7.

Fruit ovoid, 4-6 mm. long, 4-6.5 (-8.0) mm. wide, the arista rem-
nant of styles 0-2.0 mm. long, the sulci between the carpels shallow
to fairly deep, sericeous to tomentulose, hairs 0.5-1.5 mm. long;
dehiscence all along the ventral sutures and a third to half way along
the dorsal sutures, very slightly septicidal; seeds 1 or 2 per carpel,
1.9 mm. long, 1.1 mm. wide, red brown, wing or other projection
usually absent, sometimes with a wing 0.6-2.0 mm. high, 1.0 mm.
wide.

Flowering specimens collected throughout the year except in
January; fruiting ones throughout the year except in January, May,
September, and December.

TYPE LocaALity: Tanna Island, New Hebrides.

Distrisution: New Guinea to New Caledonia, common in second
growth rain forest, at the edge of the rain forest, along river banks,
in savanna, littoral, in disturbed ground, on limestone or sandy soil,
at 0-1,200 meters elevation.

996 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

NEW GUINEA: Mapana: Near Mawan village, Gogol Valley (ca. 25 km.
inland), Hoogland 4917 (A, L, US). Yllabe by Finschhafen, Weinland 254 (L,
US). Cycloop-gebergte. W. van Ifor orderafd. MHollandia, Kalkman 3504
(L). Morose: Lae, Dept. Forests Native Collector 5494 (L); Busu River near
Lae, White N.G.F. 9550 (L); Bupu River, near Lae, Millar N.G.F. 9298 (L);
Manum Waters on the Nadzab Road, Womersley & Floyd 5655 (L); Sattelberg,
Clemens 521 (L). Papua: Daru Island, Western Division, Brass 6346 (L).
Kokoda, Carr 16277 (L). Isuarava, Carr 16041 (L). Rouna, Carr 12405 (L).
Rosse Isuanp: Abaleti, Brass 28336 (L, US); Jinju, Brass 28533 (L, US). MILNne
Bay: Biniguni Camp, Gwariu River, Brass 23931 (A); Between Dabora and
Wabubu, Cape Vogel Peninsula, Brass 21871 (A); Peria Creek, Kwagira River,
Brass, 23995 (A).

SOLOMON ISLANDS: BouGaINvILLE: Kieta, Kajewski 1539 (L); Kugi-
maru, Buin, Kajewski 1797 (L). New Grorata: Waterhouse 131 (K). GuapaL-
caANAL: Nalimbu River, F. S. Walker & White B.S.I.P. 17 (L); Vulolo, Tutuve
Mt., Kajewski 2539 (L); Neighborhood of Tenawe River and the sea, Riley 9
(US). San Cristé6BaLt: Waimamura, Brass 2681 (L).

NEW HEBRIDES: Santa Cruz Istanps: Vanikoro Island, Kajewski 555
(A, UC, US). Tanna and Nova Amsterdam, Forster in 1779 (8S, W). ANEITYUM:
Collector not mentioned, 61 (S); Anelgauhat Bay, Kajewski 713 (US).

LOYALTY ISLANDS: Franc 1093 (UC).

NEW CALEDONIA: Prony 1822 A (UC). Deplanche 155 (L). Slope of
Ouen Toro, McKee 2247 (L, US). Chepenehe, McKee 5257 (L). Auf den
Bergen bei Paita, 10 mi. from Nouméa, Schlechter 14978 (L). Near native
village of Tribu Katricoin along trail northward from waterfall at fork of Bogen
River and Table Unio, Buchholz 1289 (US).

This species is close to M. vitiensis and M. aristata. The seeds of
M. ordorata are usually wingless, those of M. vitiensis are winged, and
those of M. aristata are usually aristate. The young leaves of M.
odorata are sometimes quite hairy, those of M. vittensis and M. aristata
are soon glabrate. The calyx and bracts of MM. odorata are longer than
those of M. aristata and M. vitiensis.

3. Melochia vitiensis A. Gray, Bot. U.S. Expl. Exped. 1:193. 1854.

A shrub to spreading tree 1.5-12 meters high, the young stems
fuscous-tomentulose, hairs stellate, a few forked, 0.1-0.3 mm. long,
the internodes 0.5-14.0 em. long; stipules deciduous, 2.4-7.0 mm.
long, 0.9-2.0 mm. wide, deltoid-acute to deltoid-acuminate, tomentu-
lose, hairs stellate and forked, 0.1-0.4 mm. long; petiole 2-9.5 cm.
long, indument similar to that on young stems but tending to occur
on one side.

Lamina 6-17 cm. long, 3.3-12.0 cm. wide, broadly ovate, the base
cordate, sometimes rounded to subtruncate in those subtending the
inflorescence, the apex acute to acuminate, young leaves sericeous-
tomentulose on both surfaces, hairs forked, simple, and _ stellate,
older leaves subglabrate, with few, mostly stellate hairs 0.1-0.6 mm.
long on the veins, the margin irregularly crenate-serrate, serrations
1.2-5.0 mm. wide, 0.3-2.5 mm. high, 7-10 pairs of primary veins, 2 or
3 are basal.

GOLDBERG—GENUS MELOCHIA 227

Inflorescences fairly loose, in axils of 1-4 upper leaves, 6-12.7 cm.
long, corymbiform, the flower number moderate, the ultimate divi-
sions umbellate, the umbels 2- or 3-flowered, the peduncle 6-9.5 cm.
long, with a primary branch 0.8-3.8 cm. long, a secondary one 0.7—
1.4 cm. long, and sometimes a tertiary one 1 cm. long, the pedicels
2.5-7.0 mm. long; bracts soon deciduous, 1.0-2.1 mm. long, 0.4-0.8
mm. wide, nearly linear to lanceolate and ovate, acute, densely tomen-
tulose, hairs stellate, 0.1-0.2 mm. long.

Calyx 3.5-6.0 mm. long, 3.8-6.7 mm. wide, densely tomentulose,
hairs stellate, 0.1-0.4 mm. long, the teeth 1.7-3.0 mm. long, 1.7—2.7
mm. wide, deltoid or ovate, acute, the sinus acute to narrowly rounded;
petals pale yellow or white, 5.0-7.6 mm. long, 1.7-2.8 mm. wide, oval,
elliptic, ovate or oblong, attached to the stamens for 1.5-2.0 mm., the
claw 1-2 mm. long.

Longistylous form. Stamens 4.0-5.2 mm. long, the filaments
united up to the anthers, the anthers bright yellow, 1.5-2.2 mm. long,
0.7-1.4 mm. wide, oblong, emarginate usually nearly to the middle
from both ends; pistil 7.2 mm. long, the styles free for 3.5-4.5 mm.,
united for 1 mm., sparsely sericeous, hairs 0.3-0.7 mm. long, filiform,
sometimes clavate, papillose for 0.6-1.0 mm., the ovary oblong, densely
sericeous, hairs 0.7-1.4 mm. long, the carpels sometimes 6.

Brevistylous form. Stamens 4.7 mm. long, the filaments free for
1.8 mm., the anthers 2.0 mm. long, 0.9 mm. wide; pistil 4.3 mm. long,
the styles free for 2.0 mm., united for 1 mm., filiform, the ovary
globular, densely sericeous, hairs 1 mm. long.

Fruit 7.0-9.0 mm. long, 7.0-7.5 mm. wide, the arista 1-1.5 mm.
long, globular-ovoid, pentagonal, the dorsum of carpels obtuse-angled,
the sulci between the carpels extending a fourth to a third the way to
the center of the distal half of the fruit, fuscous-sericeous-tomentose,
hairs stellate (short), forked, and simple, 0.1-1.5 mm. long, the stipe
0-0.5 mm. long; dehiscence all along the ventral sutures and a third
to half way along the dorsal sutures; seeds 1 or 2 per cell, winged, the
body brown, 2.1-2.8 mm. long, 1.3-1.6 mm. wide, the wing extending
1.3-2.0 mm. beyond the body, 1.3-1.7 mm. wide, the adaxial edge
straight, the abaxial edge convex, the apex rounded or acute.

Flowering specimens collected in March, July, October, and
November; budding ones in May; fruiting ones in October and
November.

Type tocauity: Fiji Islands: Vanua-levu, Somu-somu (on Ta-
veuni), Ovalau, Oneata (Lau, eastern group).

Distripution: Fiji Islands, in thin forest, thickets, clearings,
crass-covered “‘forehills,”’ on rocky limestone slopes and on low ground
inside the sandy beach, at 0-500 meters elevation.

228 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

FIJI ISLANDS: U.S. South Pacific Exploring Exped. under Capt. Wilkes
(GH, US 13128 lectotype). Seemann 37 (GH). Vanua Levu: MATHUATA: 5.
slopes of Mt. Numbuiloa, E. of Lambasa, A. C. Smith 6549 (A, L, 8, US). Lower
Wainunu River valley, Mbua, A. C. Smith 1727 (GH, 8. US). THAKAUNDROVE:
Nakawanga, Gressitt 2487 (US). Taveunt: Vicinity of Waiyur, Gillespie 4701
(GH). KamBara (Lau group): A. C. Smith 1301 (GH, 8, US). Maruxu:
Bryan 259 (GH).

This species is closest to M. odorata and M. aristata. It is dis-
tinguished from them under the description of M. odorata. In
addition, the leaf of M. vitiensis is usually broader and more frequently
cordate at the base than that of M. aristata.

4. Melochia longepetiolata A. C. Sm. Journ. Arn. Arb. 31:310. 1950.

A tree 4-8 meters high, the young stems minutely tomentulose,
hairs tan, stellate, 0.05-0.1 mm. long, the internodes 1-18.5 cm. long;
stipules 3-5 mm. long, 0.8-2.0 mm. wide, deltoid-acute, minutely
tomentulose, hairs stellate, 0.05-0.1 mm. long; petiole usually long,
2-19 cm. long, indument similar to that on young stems, a few hairs
to 0.2 mm. long.

Lamina 6.2-24 em. long, 3-21.5 em. wide, usually broadly ovate,
the base cordate, truncate or rounded, the apex acute, the upper
surface glabrous, the lower surface subglabrous, hairs scattered,
stellate, 0.1-0.2 (-0.4) mm. long on veins and in their axils, stellate
hairs on the interstices 0.05 mm. long, the margin irregularly crenate
to crenate-serrate, crenations or serrations 2.5-7.0 mm. wide, 0.5-1.9
mm. high, 6-10 pairs of primary veins, 2 or 3 are basal.

Inflorescences 17.5-21.5 em. long, loose, in the axil of the uppermost
leaf, sometimes not associated with a leaf or 90° from a leaf axil, at
1-2 nodes, few-flowered, corymbiform, the ultimate divisions umbel-
late, the umbels 2- or 3-flowered, the peduncle 7.5—11 cm. long, with
a primary branch 3.5-5.2 (—7.0) em. long, a secondary one 1-1.7
(-2.5) em. long, and sometimes a tertiary one to 0.8 cm. long, the
pedicels 5-7 mm. long; bracts deciduous, all fallen off.

Calyx 6-8.2 mm. long, 7.2-10.0 mm. wide, densely tomentulose,
hairs stellate and forked, 0.05-0.2 mm. long, the teeth 4—5.5 mm.
long, 3-4.8 mm. wide, ovate, acute, the sinus acute; petals salmon
pink, 7.3-10.1 mm. long, 2-2.8 mm. wide, oblanceolate or obovate-
elliptic, attached to the stamens for 1-2 mm., the claw 1-2.5 mm. long.

Longistylous form not seen.

Brevistylous form. Stamens 6.6—-7.7 mm. long, the filaments free
for 3.5-4.4 mm., the anthers 1.4-1.7 mm. long, 0.7-1.0 mm. wide,
oblong, emarginate at both ends, staminodes sometimes present,
0.2 mm. long; pistil 7.2-7.5 mm. long (somewhat developed), the
styles filiform, free for 2.6 mm., united for 1 mm., the base tomentose,
the ovary globular, densely tomentose-sericeous, hairs 1.5 mm. long.

GOLDBERG—GENUS MELOCHIA 229

Fruit 8-9 mm. long, 7.5-8.0 mm, wide, the arista 1-1.5 mm. long,
ovate, globular, obtuse-angled, the sulci between the carpels shallow,
deepest distally, extending one-fifth of way to the center of the fruit,
tomentose, hairs simple, forked, and stellate, 1-1.6 mm. long, also
stellate 0.1 mm. long, the carpels sometimes 6-7, stipe none; de-
hiscence all along the ventral sutures and one-fifth along the dorsal
sutures; seeds brown, winged, the body 2.6-2.8 mm. long, 1.6—1.7 mm.
wide, the wing 2.5-3.0'mm. high, 1.1-2.0 mm. wide, the adaxial edge
straight, the abaxial edge convex.

Flowering and fruiting specimens ‘collected in October.

Type Locauity: Kandavu, Fiji Islands, type cited below.

Distrisution: Kandavu, Fiji Islands, in dense forest, or among
reeds, at 200-500 meters elevation.

FIJI ISLANDS: Kanpavu: Mt. Mbuke Levu, A. C. Smith 218 (S, US, type).
Hills above Namalata and Ngaloa Bays, A. C. Smith 110 (8, US).

This species can be distinguished from the other species with
subglabrous leaves in the section Visenia by its long inflorescences,
long peduncles, long petioles, and relatively large calyxes. Its salmon
pink petals are also an aid to identification.

5. Melochia aristata A. Gray, Bot. U.S. Expl. Exped. 1:193. 1854.

A small tree 1.5-6.0 meters high, about 12 cm. in diameter, the
young stems and petioles fuscous-tomentulose, hairs stellate, 0.1-
0.2 mm. long, the internodes 0.7-8.3 cm. long; stipules soon deciduous,
2.44.0 mm. long, 0.8-1.5 mm. wide, deltoid or ovate, acute, densely
hirsute-tomentulose, hairs fuscous, stellate and forked, 0.1-0.2 mm.
long; petiole slender, 2-6 cm. long.

Lamina 5.5-12.0 (-15) cm. long, 2.0-8.9 (-12.4) cm. wide, broadly
ovate, to lanceolate in leaf subtending the inflorescence, the base
rounded, broadly subcordate, cordate, or nearly truncate, the apex
acute or acuminate, the upper surface tomentulose, velvety to the
touch, mostly stellate, to glabrate with hairs scattered, stellate, forked,
and simple, 0.1-0.2 mm. long, the lower surface subglabrous with
scattered stellate, forked, and simple hairs 0.1-0.4 mm. long, mostly
on the veins, the margin irregularly crenate-serrate, serrations 1.3-
5.6 mm. wide, 0.3-1.5 mm, high, 7-9 pairs of primary veins, 1-3 are
basal.

Inflorescences 5-12 cm. long, fairly loose, the flower number moder-
ate, in axils of 14 upper leaves, corymbiform, the ultimate divisions
umbellate, the umbels 2-4-flowered, the peduncle 2.5-5.2 (-7.0)
cm. long, with a primary branch 0.6-2.2 cm. long, a secondary one
0.5-0.8 (-2.2) ecm. long, and sometimes a tertiary one 0.5 cm. long,
the pedicels 1.5-3.0 mm. long in flower to 7.7 mm. in fruit; bracts
soon deciduous, 2 mm. long, 0.5-1.5 mm. wide, ovate or lanceolate,
acute, densely tomentulose, hairs stellate, 0.1-0.2 mm. long.

230 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Calyx 4.0-5.5 mm. long, 4.1-6.5 mm. wide, tomentulose, hairs
stellate, simple, a few forked, 0.1-0.4 mm. long, the teeth 1.6-3.0 mm.
long 1.8-2.5 mm. wide, deltoid to ovate, acute, the sinus acute to
narrowly rounded; petals red, pink, or white proximally and shading to
rose distally, 7.8-9.0 mm. long, 2.0-3.6 mm. wide, obovate, oblance-
olate, or elliptic, attached to the stamens for 2-2.8 mm., the claw
1-2.8 mm. long.

Longistylous form. Stamens 5.0 mm. long, the filaments free for
1.2-1.4 mm., the anthers 1.5-1.7 mm. long. 0.9-1.1 mm. wide,
oblong, emarginate at both ends; pistil 8.2 mm. long, the styles slightly
clavate, papillose for 0.8 mm., 5.0 mm. long, free to the ovary, the
ovary ovoid, densely sericeous, hairs 0.6-1.0 mm. long.

Brevistylous form. Stamens 6.2-7.0 mm. long, the filaments free
for 4.0 mm., the anthers 1.4-1.7 mm. long, 0.8-1.1 mm. wide; pistil
5.0 mm. long (slightly developed), the styles filiform, 2.6 mm. long,
free to the ovary, the ovary globular-ovoid, sericeous-tomentose,
hairs 0.7-1.1 mm. long.

Fruit 6.2-10.5 mm. long, of which the arista is 1-2 mm. long,
5.2-6.7 mm. wide, globular-ovoid, pentagonal, the dorsum of carpels
obtuse-angled, the sulci between the carpels shallow, in the distal
half of the fruit, extending a fifth to a fourth the way to its center,
tomentose, hairs stellate (short), simple, and forked, 0.1-1.3 mm.
long, the stipe 0-1.0 mm. long (base of capsule); dehiscence all along
the ventral sutures and a third to half way along the dorsal sutures,
slightly septicidal; seeds 1 or 2 per cell, wingless to slightly winged,
the body 1.8-3.3 mm. long, 1.5-1.9 mm. wide, trigonal, brown,
sometimes with an arista 0.7 mm. long, testa loosened for 0.2-0.5 mm.,
or with a wing 0.3 mm. high, 0.7 mm. wide.

Flowering specimens collected in March, May, July to September,
and November; fruiting ones in March, May, and July to September.

Typp Locauity: Upolu, Samoa, type cited below.

Distripution: Samoa, Tonga, and Tuamotu Islands, in rain
forest, thickets, in coral sand, at 0-650 meters elevation.

TONGA: Vava’u Isuanp: Near Ngofe swamp, SW. side of the island, Yuncker
16029 (US). Vava’u and Lifuka, Harvey August and October 1855 (GH).

SAMOA: Savau: Safune, Bryan 140 (A). Vaipouli, Vaupel 360 (US).
Near Fagalele, Christophersen 3403 (L). Upouu: U.S. 5S. Pac. Exped. under
Capt. Wilkes (GH, US, type). K. & L. Rechinger 1443 (US). Falefa, Hochreutiner
3449 (L). Ridge above Malololelei, Christophersen 164 (US). Id. Singamana,
near Apia, Hochreutiner 3222 (L). Turviia: Setchell 512 (US).

TUAMOTU: Maxkarea: Wilder 1149 (A).

This species is close to AZ. odorata and M. vitiensis. It is dis-
tinguished from them under the description of M. odorata. In addition,
the serrations of the leaves of M. aristata are generally shorter than
those of the other two species.

GOLDBERG—GENUS MELOCHIA 931

6. Melochia degeneriana A. C. Sm. Journ. Arn. Arb. 31:307. 1950,

A slender tree 2-15 meters high, the young stems tomentulose or
with scattered stellate and forked hairs 0.05-0.2 mm. long, the
internodes 0.5-5.0 cm. long; stipules soon deciduous, 3.0-5.3 mm.
long, 0.5-1.0 mm. wide, lanceolate, tomentulose to subglabrous, hairs
stellate, 0.1-0.5 mm. long; petiole 0.3-3.0 cm. long, adaxial surface
tomentulose, hairs stellate, 0.1-0.3 mm. long.

Lamina thin, 4-11.5 em. long, 1.8-5.7 (-6.5) em. wide, usually
ovate, sometimes lanceolate-ovate, the base broadly rounded, rarely
also narrowly rounded, the apex acute, both surfaces subglabrous,
few scattered stellate, simple, and forked hairs 0.1-0.5 mm. long on
the veins and sometimes also on the interstices, the margin irregularly
and shallowly crenate-serrate, serrations distant, 1.3-8.3 mm. wide,
low, 0.3-1.1 mm. high, 6-10 pairs of primary veins, 1 or 2 are basal,
the second much stronger than the first.

Inflorescences fairly loose, the flower number moderate, sometimes
few, in the axils of the upper 4 leaves, sometimes apparently terminal,
or leaf-opposed, 6-14 cm. long, corymbiform, the ultimate divisions
umbellate, the umbels 2-4-flowered, the peduncle 2.5-4.5 (-6.0)
em. long, with a primary branch 0.5-1.5 cm. long, a secondary one
0.3-0.5 cm. long, and sometimes a tertiary one to 0.5 cm. long, the
pedicels 2.0-6.0 mm. long, densely tomentulose, hairs stellate, 0.1
mm. long and glandular, 0.2 mm. long; bracts soon deciduous, 1.1-
1.6 mm. long, 0.4 mm. wide, lanceolate, tomentulose, hairs simple,
forked, and stellate, 0.1 mm. long.

Calyx brownish pink 4.1-6.0 mm. long, 5.1-6.6 mm. wide, tomen-
tulose, hairs stellate, 0.05-0.2 mm. long, sometimes also glandular,
0.2 mm. long, the teeth 1.2-3.1 mm. long, 1.6-3.1 mm. wide, deltoid,
acute, the sinus acute to slightly rounded; petals pale yellow, 6.0-9.1
mm. long, 2.7-3.1 mm. wide, elliptic to obovate, attached to the
stamens for 1.0-2.5 mm., the claw 1.0-2.5 mm. long.

Longistylous form. Stamens 5.2-5.5 mm. long, the filaments
united up to the anthers, the anthers 1.8-2.0 mm. long, 0.9-1.4 mm.
wide, oblong, emarginate at both ends, nearly to the middle from the
apex; pistil 7.6 mm. long, the styles filiform, papillose for 1.5 mm.,
free for 5.0 mm., united for 1.0 mm., the ovary ovoid densely sericeous,
hairs 0.7-1.0 mm. long, stipe none.

Brevistylous form. Stamens 5.5-7.0 mm. long, the filaments pale
yellow, free for 2.5-3.5 mm., the anthers 2.1-2.5 mm. long, 1.3-1.4
mm. wide; pistil 4.0 mm. long, the styles 1.4-1.6 mm. long, free
to the ovary, papillose for 0.8 mm., the ovary globular, densely
tomentose-sericeous, hairs 1.0-1.2 mm. long, the carpels sometimes
6 or 7.

232 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Fruit tough, 9-10 mm. long, of which the arista is 1.5 mm. long,
6-7.2 mm. wide, globular-ovoid, the dorsum of carpels obtuse-angled,
the sulci between the carpels extending a fourth to half way to the
center of the distal half, tomentose when young, hairs simple and
forked, 0.3-1.2 mm. long, and stellate, 0.1 mm. long, stipe none;
dehiscence all along the ventral sutures and a third along the dorsal
sutures; seeds (immature) brown, winged, the body 2.3 mm. long,
1.3 mm. wide, the wing 1.3-3.0 high, 0.7-1.9 mm. wide.

Flowering specimens collected in January, May to September, and
December; fruiting ones January to March, May to September, and
December.

Typr LocaLity: Viti Levu, Fiji Islands, type cited below.

Disrripution: Viti Levu, Fiji Islands, in forest, often on dry
crests, at 0-1,150 meters elevation.

FIJI ISLANDS: Vitrt Levu: Greenwood 742 (K). mMBa (formerly Tholo
North): Western and southern slopes of Mt. Tomanivi (Mt. Victoria), A. C.
Smith 5095 (L, US, type). THOLO NORTH: VIC. OF NANDARIVATU: Nauwanga,
Degener 15399 (US). Lauroxa: N. of Lomolomo, Degener & Ordonez 13643
(S, US). NANDRONGA AND NAVOSA (formerly iTholo# West) : Southern slopes of
Nausori Highlands, A. C. Smith 4585 (L, 8, US). Naduruloulou Agr. Exp.
Garden, Lam 6933 (L). Namosi, Gillespie 2825 (US). Leviti Creek, Suva,
Mrs. Parham 21 (A).

This species is closest to M. mollipila. The differences are given
under the description of the latter species.

7. Melochia mollipila A. C. Sm. Journ. Arn. Arb. 31:308. 1950.

A branched shrub or small tree 3-7 meters high, the young stems
densely fuscous-canescent, tomentulose, hairs stellate, 0.05-0.2 mm.
long, the internodes 0.5-6.0 cm. long; stipules soon deciduous, 3.2-
5.0 mm. long, 0.9-1.2 mm. wide, deltoid, acute, tomentulose, hairs
stellate, 0.1-0.2 mm. long; petiole 0.5-6.0 cm. long, indument similar
to that on young stems but hairs to 0.5 mm. long, average 0.2 mm.

Lamina 3.5-13.0 em. long, 1.8-11.5 cm. wide, the small leaves being
close to the inflorescence, usually broadly ovate, the base rounded
to subcordate, rarely cuneate or subtruncate, the apex acute, the
upper surface subglabrous, hairs few, scattered, stellate and forked,
0.1-0.2 mm. long, the lower surface slightly canescent, velvety to the
touch, tomentulose, hairs stellate, 0.1-0.4 (most 0.2) mm. long, the
margin irregularly and shallowly crenate or crenate-serrate, crena-
tions or serrations 2-7 mm. wide, 0.3-0.8 mm. high, 8 or 9 pairs of
primary veins, 2 are basal.

Inflorescences compact to fairly loose, the flower number few to
moderate, in the axils of the upper 2 or 3 leaves and 3-9.5 cm. long,
sometimes also terminal and 3.5 cm. long, corymbiform, the ultimate
divisions umbellate, the umbels 2- or 3-flowered, the peduncle 2-4

GOLDBERG—-GENUS MELOCHIA 233

(-5.5) em. long, with a primary branch 0.4—1.4 cm. long and a sec-
ondary one 0.4—0.6 cm. long, the pedicels 1-3 mm. long in flower, to
5 mm. long in fruit; bracts soon deciduous, 2-3 mm. long, 0.7-1.0 mm.
wide, deltoid, acute, tomentulose, hairs stellate, 0.1-0.2 mm. long.

Calyx pink, thick, 5.1-5.8 mm. long, 5.8-7.3 mm. wide, densely
tomentulose, hairs stellate, fewer forked, 0.1-0.3 mm. long, the
teeth 2.6-3.0 mm. long, 2.5-3.3 mm. wide, deltoid to ovate, acute,
the sinus acute to somewhat rounded; petals pale yellow, 6.7-8.3
mm. long, 2.0-3.2 mm. wide, elliptic to oval, attached to the stamens
for 0.8-1.5 mm., the claw 1-1.5 mm. long.

Longistylous form. Stamens pale yellow, 4.3-5.0 mm. long, the
filaments united up to the anthers or free for 0.5 mm., the anthers
1.4-1.9 mm. long, 1.2-1.3 mm. wide, oblong, emarginate at both
ends, sometimes fairly deeply so; pistil 6.3-6.7 mm. long, the styles
filiform, papillose for 1 mm., free to the ovary, 3.5-4.0 mm. long, the
ovary globular-ovoid, the ovary and base of styles sericeous-tomentose,
hairs 0.5-1.2 mm. long, carpels sometimes 7-8, stipe none.

Brevistylous form. Stamens 6.2 mm. long, the filaments free for
4.0 mm., the anthers 1.6-1.7 mm. long, 1.0 mm. wide; pistil 4.5 mm.
long, the styles 2.0 mm. long, filiform, free to the ovary, the ovary
ovoid-globular, densely sericeous, hairs 0.7 mm. long, the carpels
sometimes 6-8.

Fruit globular-ovoid, 5-7.5 mm. long, 5.3-9.4 mm. wide, the
arista soft, 0.7 mm. long., the dorsum of carpels obtuse-angled, the
sulci between the carpels deepest near the apex, where they extend
a third to half way to the center of the fruit, sericeous-tomentose,
canescent, hairs simple and forked, 0.5-1.2 mm. long, stellate 0.2
mm. long, the stipe 0-1.0 mm. long; dehiscence all along the ventral
sutures and one-fourth the way along the dorsal sutures, also partly
septicidal; seeds winged, the body 2-2.5 mm. long, 1.4-1.5 mm. wide,
the wing 2.6-3.5 mm. high, extending 1.8 mm. beyond the body, 1.7-
2.2 mm. wide just above the body, adaxial edge straight, abaxial
edge convex.

Flowering specimens collected August to October; fruiting ones in
September and October.

TypE LocaLity: Viti Levu, Fiji Islands, type cited below.

Distrisution: Viti Levu, Fiji Islands, in hillside and grassland
thickets, usually in the sun, at 550-800 meters elevation.

FIJI ISLANDS: Vir1 Levu: mBa (formerly Tholo North): Slopes of the
escarpment N. of Nandarivatu, A. C. Smith 6040 (L, US, type). NANDRONGA
AND NaAvosa: N. portion of Rairaimatuka Plateau, A. C. Smith 5588 (L, S, US).
Mt. Evans Range, Greenwood 1229 (US), 1239 (US).

This species is closest to M. degeneriana. The lower surface of the
leaf of the latter is subglabrous, that of M. mollipila is persistently

234 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

tomentulose. The capsule of AZ. mollipila is shorter than that of
M. degeneriana, and broader than long; capsule of MZ. degeneriana is
longer than broad.

8. Melochia roseiflora A. C. Sm. Journ. Arn. Arb. 31:3381. 1950.

A shrub or tree 1-8 meters high, the young stems and petioles
tomentulose, hairs brownish, stellate, 0.05-0.5 mm. long, average
0.2 mm., the internodes 0.3-7.0 cm. long; stipules 2.5-5.5 mm. long,
1.0-1.5 mm. wide, deltoid, acute, the abaxial surface tomentulose-
hirsute, hairs stellate, 0.1-0.3 mm. long; petiole short, 0.3-1.5 cm.
long.

Lamina 3.7-9.0 cm. long, 2.2-6.2 em. wide, broadly ovate to
lanceolate-ovate, the base usually rounded, sometimes cuneate, the
apex acute to somewhat rounded, the upper surface somewhat shiny
and rather dark green, subglabrous, hairs on the veins few, scattered,
stellate, 0.1-0.2 mm. long, the lower surface lighter green, hairs on
the veins scattered, stellate, 0.1-0.4 mm. long, the margin irregularly
and rather shallowly crenate or crenate-serrate, crenations or serra-
tions 1.2-5.5 mm. wide, 0.38-1.0 mm. high, 6-8 pairs of primary
veins, 1 or 2 are basal, the second much stronger than the first.

Inflorescences fairly compact, 4-6.5 em. long, in the axils of the
upper 2 or 38 leaves, corymbiform, the flowers few to moderate in
number, the ultimate divisions umbellate, the umbels 2- or 3-flowered,
the peduncle 2.3-3.2 cm. long, with a primary branch 0.6-1.4 cm.
long, and sometimes a secondary one to 0.5 em. long, the pedicels
deep pink, 1.5-3.0 mm. long in flower, to 5 mm. long in fruit; bracts
1.0-3.1 mm. long, 0.5-0.7 mm. wide, deltoid to lanceolate, acute,
tomentulose-hirsute, hairs stellate, 0.1-0.2 mm. long.

Calyx deep pink, 4.5-5.0 mm. long, 6.0-6.5 mm. wide, tomentulose,
hairs stellate, 0.05-0.2 mm. long, also a few glandular, 0.1-0.4 mm.
long on the young calyx, the teeth 2.2-2.5 mm. long, 2.5-2.8 mm. wide,
deltoid, acute to acuminate, the sides rounded, the sinus acute; petals
bright pink, slightly paler than the calyx, 6.5-7.9 mm. long, 2.2-2.9
mm, wide, elliptic to slightly obovate, attached to the stamens for
1.5-1.7 mm., the claw 1.5-2.0 mm. long.

Longistylous form. Stamens 4.7-5.0 mm. long, the filaments free
for 0.56 mm., the anthers yellow, 1.5-2.0 mm. long, 1.1-1.3 mm. wide,
oblong, emarginate nearly to the middle from both ends; pistil 7.8-8.3
mm. long, the styles rich pink, filiform, papilose for 1-1.2 mm., free
for 5.0-5.2 mm., united for 0.5-0.7 mm., proximal half tomentose, the
ovary globular-ovoid to elliptic, densely tomentose, hairs 0.7-1.0 mm.
long, the carpels sometimes 6, stipe none.

Brevistylous form. Stamens 8.1 mm. long, the filaments nearly
white, free for 4.5 mm., the anthers 1.7—1.8 mm. long, 0.7-0.9 mm. wide;

GOLDBERG—GENUS MELOCHIA 935

pistil 4.8 mm. long, the styles recurved, 2 mm. long, filiform, free to
the ovary, the ovary ovoid, densely sericeous, hairs 1 mm. long, the
carpels sometimes 7.

Fruit 7.0-7.7 mm. long, of which the arista is 0.5 mm. long, 6.0-6.3
mm. wide, globular-ovoid, the dorsum of carpels obtuse-angled, the
sulci between the carpels shallow, except in the distal half, where they
extend a third of way to the center of the fruit, tomentose, hairs
simple, forked and stellate, 0.3-1.2 mm. long, the stellate ones also 0.1
mm. long, stipe none; dehiscence all along the ventral sutures and a
fourth to half way along the dorsal sutures, also partly septicidal;
seeds brown, winged, the body 2.0-2.5 mm. long, 1.3-1.7 mm. wide,
the wing 1.3-1.7 mm. high, 1.0-1.5 mm. wide, obliquely deltoid, apex
acute or rounded.

Flowering specimens collected in April and May; fruiting ones in
May.

Typs Locauity: Viti Levu, Fiji Islands, isotypes cited below.

Distrisution: Viti Levu, Fiji Islands, in grassland-forest transition
and in dense ridge forest and thickets, at 700—-1,195 meters elevation.

FIJI ISLANDS: Vitrt Levu: mspa: Northern portion of Mt. Evans Range,
between Mt. Vatuyanitu and Mt. Natondra, A. C. Smith 4361 (L, US, isotypes).
Summit of Mt. Koroyanitu, high point of Mt. Evans Range, A. C. Smith 4236
(L, 8, US). Mt. Evans, Greenwood 316 (K).

This species appears to be closest to M. degeneriana and M. mollipila.
The latter two have yellow flowers, those of M. roseiflora are bright
pink. The petioles of M. roseiflora are usually shorter.

9. Melochia grayana A. C. Sm. Journ. Arn. Arb. 31:309. 1950.
Melochia vitiensis var. 8B A. Gray, Bot. U.S. Expl. Exped. 1:198. 1854.

A branched shrub 1.5-2 meters high, the young stems and petioles
minutely tomentulose and cinereous or hairs scattered, stellate, forked,
and simple, 0.05-0.3 mm. long, the internodes 0.2—-3.2 cm. long;
stipules 3.6-6.0 mm. long, 0.9-1.6 mm. wide, deltoid to lanceolate,
acute to acuminate, subglabrous or ciliate, hairs few, simple, 0.1-0.3
mm. long; petiole 0.5—2.4 cm. long.

Lamina subcoriaceous, 3.1-7.3 (-9.0) em. long, 1.1-2.4 (-4.5) cm.
wide, lanceolate to elliptic, the base cuneate to narrowly rounded, the
apex acute, the upper surface glabrous, the lower surface with few,
scattered, simple, forked and stellate hairs 0.1-0.3 mm. long on the
veins, the margin irregularly and shallowly serrate to crenate-serrate,
serrations 1.3-5.7 mm. wide, 0.4-1.2 mm. high, 5-8 pairs of primary
veins, 1 is basal.

Inflorescences loose, 5-10.5 cm. long, in axils of upper 3-5 leaves,
corymbiform to subpaniculate, the flower number usually few, some-
times moderate, the ultimate divisions umbellate, the umbels 2—4-
flowered, the peduncle 2-3.5 cm. long, with a primary branch 0.7-2.4

236 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

cm. long, sometimes a secondary one to 1.0 cm. long and a tertiary one
to 1.6 cm. long, the pedicels 1-3.5 mm. long; bracts soon deciduous,
1.2—-2.0 mm. long, 0.1-0.8 mm. wide, lanceolate to linear, tomentulose
and ciliate, hairs stellate and forked, 0.1-0.2 mm. long.

Calyx orange-tinged to bright red, 5.5-6.7 mm. long, 4.0-5.5 mm.
wide, tomentulose, hairs stellate and forked, usually 0.05—-0.2, rarely
to 0.5 mm. long, the teeth 2.2-3.2 mm. long, 1.5-2.5 mm. wide, deltoid,
acute to acuminate, the sinus acute; petals yellow, 5-9 mm. long,
1.6-3.0 mm. wide, elliptic, suboval or oblong, the apex rounded, the
base acute, attached to the stamens for 2.6-3.0 mm., the claw 1-2 mm.
long.

Longistylous form. Stamens yellow, 4.5-5.2 mm. long, the fila-
ments free for 1-2 mm., the anthers 1.7 mm. long, 1.1-1.2 mm. wide,
oblong, deeply emarginate at both ends, sacs separate at the apex,
pistil 5.7-7.1 mm. long, the styles slightly clavate, papillose for
0.8-1.0 mm., 2.9-4.5 mm. long, free to the ovary, the ovary narrowly
ovoid, densely tomentose-sericeous, the carpels sometimes 7-8, stipe
none.

Brevistylous form not seen.

Fruit. Only parasitized fruit seen.

Flowering specimens collected in October and November; fruiting
ones in October.

TypE Locanity: Vanua Levu, Fiji Islands, isotype cited below.

DistriBuTION: Vanua Levu, Fiji Islands, in forest, at 60-590 meters
elevation.

FIJI ISLANDS: Horne 635 (GH) ?M. grayana, early bud stage. VANUA
Levu: MarHuaTaA: Summit ridge of Mt. Numbuiloa, I. of Lambasa, A. C. Smith
6525 (US, isotype). Muthuata, U.S. Expl. Exped. under Capt. Wilkes (US
13126, type of M. vitiensis var. 8). Labasa, Macuata Coast, Greenwood 620A
(K). Mt. Labasa, Greenwood 620 (Kx).

This species can be distinguished from the other species in the
section Visenia by its lanceolate or elliptic ‘leaves;*the other species
have at least some ovate leaves,’'and they are usually considerably
broader.

10. Melochia compacta Hochr. Ann. Conserv. Jard. Bot. Genéve 21:429. 1920.

A shrub or small tree 1-6 meters high, the young stems and petioles
tomentulose, hairs tan, stellate, and usually also forked and simple,
0.1-0.6 mm. long, the internodes 0.5-1.0 (—4.5) cm. long; stipules
3.3-8.0 (—9.0) mm. long, 1.7—4.1 (—-5.0) mm. wide, lanceolate to ovate,
acuminate, hirsute or tomentulose, hairs forked and simple, 0.1-
0.5 mm. long; petiole 1-6.5 (-13) em. long.

Lamina 6.2-16 (—28) em. long, 4.8-12.5 (—24) em. wide, ovate, the
base rounded to cordate, the apex acute, often densely tan-canescent
when young, both surfaces of older leaves glabrescent, hairs scattered,

GOLDBERG—GENUS MELOCHIA 237

stellate and usually also forked and simple, 0.1-1.0 mm. long, the
margin irregularly crenate-serrate, serrations 0.9-8.0 mm. wide,
0.3-1.7 mm. high, 8-10 pairs of primary veins, 2 or 3 are basal.

Inflorescences 3.5-20 cm. long, usually compact, flower number
moderate to fairly abundant, in the axils of the upper 3-5 leaves,
corymbiform, the ultimate divisions umbellate, the umbels about
5-flowered, the penduncle 2-16 cm. long, with a usually short primary
branch 0.5-4.0 cm. long, sometimes a secondary one 0.3-1.5 cm. long,
and a tertiary one 0.5-1.0 cm. long, the pedicels short, 0-3 mm. long;
bracts rather broad, sometimes 2 united, 2-4.5 mm. long, 0.8-3.0 mm.
wide, ovate to lanceolate, tomentulose or hirsute and ciliate, hairs
simple, forked, and stellate, 0.1-0.9 mm. long.

Calyx 6-9 mm. long, 5.3-8.0 mm. wide, tomentulose and sericeous,
hairs stellate and forked, 0.1-1.0 mm. long, the teeth 2.3-5.4 mm.
long, 2-4 mm. wide, deltoid to ovate, acuminate or acute, the sinus
acute; petals pink, red proximally and pink or white distally, rarely
rose and yellow, or white, 6.3-15.5 mm. long, 2.3-7.5 mm. wide,
obovate-cuneate, attached to the stamens for 1-2.5 mm., the claw
1.5-3.0 mm. long.

Longistylous form. Stamens 5-6 mm. long, the filaments united
up to the anthers or free for 0.5 mm., the anthers 1.2-1.9 mm. long,
0.6-1.2 mm. wide, oblong, emarginate at both ends; pistil 5.2-9.0 mm.
long, the styles filiform, slightly clavate, papillose for 0.8-2.0 mm.,
free for 2.9-6.0 mm., to the ovary, or united for 0.5 mm., the ovary
ovoid or elliptical, densely sericeous, hairs simple and forked, 0.5~1.5
mm. long, stipe none.

Brevistylous form. Stamens 6.2 mm. long, the filaments free for 2-3
mm., the anthers 1.2-1.3 mm. long, 0.7 mm. wide; pistil 3.6 mm. long,
the styles filiform, 2.0 mm. long, free to the ovary, the ovary densely
sericeous, hairs simple and forked, 1-1.4 mm. long.

Fruit (4-) 5.2-9.0 (-10) mm. long, with an arista 0.5-2.0 mm. long,
4.2-6.5 mm. wide, globular-ovoid, the dorsum of carpels rounded, the
sulci between the carpels shallow, extending one-fifth the way to the
center of the fruit, tomentulose and sericeous, hairs simple, forked and
stellate, 0.1-1.5 mm. long, the stipe 0-0.9 mm. long; dehiscence all
along the ventral sutures and a third to half way along the dorsal
sutures, partly septicidal; seeds wingless, 2.4-3.5 mm. long, 1.2-1.9
mm. wide.

Flowering specimens collected in March, April, and June to Decem-
ber; fruiting ones in March and June to December.

Tyre Locauity: Marianas Islands, probably Guam. The type was
collected by Gaudichaud and is cited below.

DistriBuTion: Palau, Yap, and Marianas Islands, in forest or on
cleared land, on coastal plain, on limestone, at 1-170 meters elevation.

227-051—67_4

238 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

CAROLINE ISLANDS: Patau Istanps: Peleliu Island, SW. side, Fosberg
25983 (BISH, US). Urukthapel Island, E. end, around lighthouse on top of peak,
Fosberg 32039 (BISH, US). Yap Isuanps: yap IsLanp: /Tallier 547 (L). Kanehira
1121 (US). Low E. ridge of Mt. Matade, slopes just above Yap Town, Fosberg
25574 (BISH, US). RumuNa: Ross Oct. 5, 1949 (US). Tomi IsLanp: Hosaka
3293 (BISH, US). vunrrmi aTott: Mogmog Island, Knox 829 (US); Fosberg &
Wong 25471 (BISH, US). Potangeras Island, Hosaka 3237 (BISH, US).

MARIANAS ISLANDS: Gaudichaud 66 (G, type, US neg. 5415). Guam:
Oca Point, Moore 43 (US). Machanao Dist., Bryan 1197 (US). N. of NW. Field,
Steere 1 (US). N. end of Harmon Field, Fosberg 39252 (BISH, US). Ritidian
Point, Fosberg 25317 (BISH, US); Hosaka 3091 (BISH, US); Anderson 218
(BISH, US). Vicinity of Agafia, Anderson 174 (BISH, US). Rora: Rota and
vicinity, Fosberg 25068 (BISH, US). NW. coast of W. end of main part of island,
between Rota and Tataacho Point, Fosberg 25017 (US). Sarpan: W. of Lake
Susupe, Fosberg 25264 (BISH, US).

This species is closest to M. villosissima. Differences are given
under the description of the latter species.

11. Melochia villosissima (Pres!) Merr. Philip. Journ. Sci. 15:543. 1919.
Riedlea villosissima Presl, Rel. Haenk. 2:146. 1835.
?Melochia Lessoniana K. Schum. ex Volkens, Eng. Bot. Jahrb. 31:468.
1902, nomen nudum.
Melochia hirsutissima Merr. Philip. Journ. Sci. 9:113. 1914.

Usually a shrub, sometimes a small tree, 0.7-7.6 meters high, the
young stems and petioles densely fuscous-rufescent tomentose, hairs
forked, simple, and stellate, 0.3-2.0 mm. long, the internodes 0.3-14
em. long; stipules 2-5 (-9) mm. long, 1.0-2.5 (-7.0) mm. wide, lance-
olate to ovate, acute, sericeous-tomentose and ciliate, hairs forked,
stellate, and simple, 0.7-2.0 mm. long; petiole 1.1-14.5 em. long.

Lamina 4.4-30 em. long, 3.0-21.5 cm. wide, ovate, largest blades
sometimes with a pair of lateral lobes 2 em. high and 2 em. wide, the
base rounded to cordate, the apex acute, typically canescent and
densely woolly on both surfaces, hairs forked and stellate (shorter ones),
0.2-1.2 mm. long, the margin irregularly crenate-serrate, serrations
1.1-3.2 mm. wide, 0.5-1.1 mm. high, 6-8 pairs of primary veins, 2 are
basal.

Inflorescences usually compact, 4-14 cm. long, in axils of upper
4-6 leaves, corymbiform, flowers fragrant, number usually moderate,
sometimes few or numerous, the peduncle thick, 2—3.5 mm. wide,
2.8-9.0 (-15.5) cm. long, usually with a short primary branch
to 1.0 (-5.5) em. long, rarely a secondary one 0.5 (-2.5) em. long,
rarely a tertiary one 0.5 (-3.0) cm. long, the pedicels usually very
short, 1-2 (4) mm. long; bracts 2-3.5 mm. long, 1.1-2.5 mm. wide,
ovate, acute to rounded at the apex, hirsute and ciliate, hairs forked,
stellate, and simple, 0.3-1.3 mm. long.

Calyx thick, 7.9-8.7 mm. long, 5.2-7.0 mm. wide, densely sericeous-
tomentulose, hairs ferruginous, stellate and forked, 0.2-1.5 mm.
long, the teeth 3.3-4.0 mm. long, 2.7-3.3 mm. wide, deltoid to ovate,

GOLDBERG—GENUS MELOCHIA 939

acute to slightly acuminate, the sinus acute; petals pink to red, some-
times fading distally, rarely magenta, relatively thick, 7.4-8.9 mm.
long, 3-5 mm. wide, obovate, sometimes emarginate, attached to the
staminal tube for 1-1.7 mm., the claw 1.5-2.0 mm. long.

Longistylous form. Stamens 5.2 mm. long, the filaments united
up to the anthers, the staminal tube rose colored, the anthers 1.7-1.9
mm. long, 1.2-1.4 mm. wide, oblong, emarginate nearly to the middle
from both ends; pistil 8.5 mm. long, the styles filiform, papillose for
1.5 mm., free to the ovary, 5.0 mm. long, the ovary ovoid, densely
sericeous, hairs 0.9-1.8 mm. long, stipe none.

Brevistylous form. Stamens 5.8-6.1 mm. long, the filaments free
for 3 mm., the anthers 1.8-1.9 mm. long, 0.7-0.9 mm. wide; pistil 5.3
mm. long (some development), the styles 2 mm. long, the ovary
globular-ovoid, densely sericeous-tomentose, hairs to 1.7 mm. long.

Fruit (4-) 6.4-7.0 (-8.0) mm. long, of which 1-1.5 (-3) mm. is
arista, 5.2-6.0 mm. wide, globular-ovoid, the dorsum of carpels
obtuse-angled, the sulci between the carpels shallow, in the distal
half of the fruit, extending a fifth to half way to its center, densely
tomentose, hairs simple, forked, and stellate, 0.3-1.3 mm. long, the
stipe 0-0.5 mm. long; dehiscence all along the ventral sutures and a
fifth to half way along the dorsal sutures, also partly septicidal;
seeds generally wingless, 3.0-3.5 mm. long, 1.5-2.0 mm. wide, trigonal,
of 10 seeds examined only one had a faint indication of a wing, 0.3 mm.
high and 1 mm. wide, the apex rounded.

Flowering and fruiting specimens collected throughout the year
except in November, most commonly in June to September.

TYPE LOCALITY: Guam.

DistrrisutTion: Caroline and Marianas Islands, usually in disturbed,
open, shrubby areas previously cultivated, on soils or rocks of volcanic
or limestone origin, at 1-360 meters elevation.

CAROLINE ISLANDS: Tjaperna plateau (?Kusaie), Hallier Aug. 27, 1903
(L). Pauau Isuanps: Arukoron-ségen, Hosokawa 7044 (A, US). BABELDAOB
ISLAND: KE, coast, hills between Melekiok and Lake Ngardok, Fosberg 32596
(BISH, US). NGARAKABESANG (ARAKABESAN) ISLAND: S. side of W. peninsula,
Fosberg 32131 (BISH, US). cGaspan: Hosokawa 9641 (A). TruK IsLANDs:
MOEN ISLAND: E. end, Hosaka 2795 (BISH, US); Slopes of Mt. Tonaachau
(Toladjau), Fosberg 24431 (BISH, US). puBLoNn Istanp: Fosberg 24536 (BISH,
US). SeNyavin IsLANDS: PONAPE: Matalanim, Hosaka 3548 (BISH, US).
KUSAIE: Kanehira 1343 (US). Lelehafen’s Kénigsland, Hallier Sept. 3, 1903
(L). Lowest slopes and foot of S. side of Mt. Matanta (Buache) N. of head of
Lela Harbor, Fosberg 26570 (BISH, US). LEeLuistanp: Clarke Ku-78 (US).

MARIANAS ISLANDS: Guam: McGregor 456 (L, US, isotypes of M.
hirsutissima). Nelson 1 (US), 353 (US). Faepi Point, approach to Mt. Lamlam,
Necker 374 (US). Near Telefofo (?Talofofo) River, Rodin 683 (US). Agat,
Achugao, Hosaka 3192 (BISH, US). Mt. Tenjo and Mt. Reconnaisance area,
Moore 217 (US). S. of Asan Point and Piti, Anderson 73 (BISH, US). Lower

240 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Fonte River, W. of Agafia, Fosberg 31238 (BISH, US). Chalandao Mt. 1 km.
SE. of Jumujong Manglo Mt., Fosberg 35379 (BISH, US). Anao (?Anae),
above cliffs, Fosberg 38639 (BISH, US). Com.-Marianas Fuel Storage Area,
Fosberg 35213 (BISH, US). Mataguac, Moran 4449 (US). Rora: Necker
R52 (US). Kanehira 1762 (US). Hosokawa 7556 (A, US). Airstrip, N. of
Shimapaaru, Fosberg 25158 (BISH, US). Sarpan: Oleai, Fosberg 31283 (BISH,
US). Raishingan, Hosokawa 8028 (US). Pagan: Anderson 501 (BISH,
US). Aarinan (AcrRIGAN): Trail around S. side of island on coastal strip, a
narrow shelf, Fosberg 31567 (BISH, US).

This species is closest to M. compacta. The leaves of the latter
are glabrescent, those of M. villosissima are persistently fairly densely
hairy. The hairs of the young stems of M. villosissima are twice as
long as those of M. compacta.

Section II. Mougeotia Griseb. Fl. Brit. W. Ind. 94. 1859
Section Anamorpha Triana & Planch. Prodr. Flo. Novogran. 215. 1862.

Perennial, rarely annual, herbs to shrubs, rarely a small tree,
0.1-5.0 meters high, the taproot not thicker than the base of the stem
or to 2.5 times thicker and 0.4—2.2 cm. wide, the young stems tomen-
tulose, hirsute or pilose, sometimes canescent, hairs sometimes tan,
golden or ferruginous, simple, stellate, forked, or glandular, 0.1-4.0
mm. long, internodes 0.4-11.0 cm. long; stipules 1.5-10.3 mm. long,
0.2-4.0 (-5.0) mm. wide, usually deltoid-acuminate or deltoid-acute
or lanceolate, usually ciliate, and hirsute, sericeous or tomentulose,
hairs simple, stellate or forked, rarely also glandular, 0.1-3.5 mm.
long; petiole 0.1-6.7 (-8.0) cm. long.

Lamina 0.8-14.0 (—22) em. long, 0.2-8. 8 (—10.5) em. wide, usually
ovate or lanceolate, sometimes oblong-ovate or nearly orbicular, the
base rounded to cordate, sometimes truncate, the apex usually acute
or acuminate, sometimes rounded, both surfaces glabrate, tomen-
tulose, sericeous or velutinous, hairs simple, forked, or stellate, rarely
also glandular, 0.1-4.0 mm. long, the margin irregularly crenate-
serrate, serrations 0.5-8.5 mm. wide, 0.3-3.6 mm. high, 5-14 (-15)
pairs of primary veins, usually 2 are basal.

Inflorescences axillary, terminal, or both, 0.6-21.0 (-31) cm. long,
spicate, subcapitate, paniculate, umbellate, or corymbiform, the
flowers sometimes very fragrant, the peduncle 0-8.5 cm. long, rarely
@ primary branch 0.4-3.5 cm. long and a secondary one 0-1.5 cm.
long, the pedicel 0-9.3 mm. long; bracts 0.5-9.5 mm. long, 0.1-3.9
(-4.8) mm. wide, usually narrower than the stipules, sometimes
several immediately subtending each flower and sometimes 2 or 3
united part way, pubescence similar to that of the stipules.

Calyx at anthesis 3.0-7.8 (-8.5) mm. long, 2.4-6.5 mm. wide,
sometimes accrescent, becoming up to 12.3 mm. long, 12.0 mm. wide,

GOLDBERG—GENUS MELOCHIA 241

tomentulose, hirsute or sericeous, sometimes ciliate, hairs simple,
stellate, glandular or forked, 0.1-1.3 (-4.0) mm. long, the teeth
1.3-5.3 (-6.5) mm. long, 0.5-2.8 (-5.5) mm. wide, the sinus usually
acute, sometimes narrowly rounded; petals usually yellow to orange,
or white, sometimes pink, reddish, white with red veins or purple,
3.7-12.0 mm. long, 1.4-5.0 mm. wide, usually obovate or oblanceo-
late, cuneate, free of the staminal tube or attached to it for 0.3-1.5
(-2.7) mm., the claw 0-1.5 (-4.0) mm. long, rarely a few hairs just
above the claw; anther 0.7-2.1 mm. long, 0.4-1.1 mm. wide, oblong,
emarginate at both ends, usually to the middle from the apex, stam-
inodes rarely present, 0.1-0.4 mm. long.

Longistylous form. Stamens 2.4-5.9 mm. long, the filaments
united up to the anthers, but separate for 0.5-1.0 mm. in M. ulmifolia,
rarely tufts of hair 0.4-1.3 mm. long present at the base of the staminal
tube, alternate with the filaments, hairs 0.1-0.2 mm. long; pistil
2.6-8.8 (-12) mm. long, the styles free for 1.2-5.3 mm., usually to the
ovary, sometimes united for 0.3-4.0 mm., papillose at the apex for
0.5-1.2 mm.

Brevistylous form. Stamens 2.8-8.2 mm. long, the filaments free
for 0.5-5.0 mm.; pistil 2.3-6.0 mm. long, the styles free for 1.1-2.5
mm., usually to the ovary, sometimes united for 0.2-0.7 mm., papillose
for 0.3-0.7 mm. at the apex.

Fruit 2.5-7.5 mm. long, 2.9-6.5 mm. wide, globular, pentagonal,
the carpels usually obtuse at the base and carinate above the middle,
the sulci usually shallow, extending one-fifth to one-third, rarely
half, the way to the center of the fruit, usually tomentulose or
hirsute, hairs usually stellate, forked, and simple, sometimes also
glandular, 0.1-1.0 mm. long, the stipe 0-1.0 mm. long; dehiscence
usually septicidal into cocci, or sometimes along the entire ventral
suture and a third to half way along the dorsal suture and slightly
septicidal or tardily septicidal and the fruit falling apart; seeds 1.5-3.0
mm. long, 0.9-2.0 mm. wide, rarely the chalazal end loosened, mem-
branous, and almost winglike for 0.1-0.3 mm., usually 1 per cell
but often 2.

LecroryreE: Melochia pilosa (Mill.) Fawe. & Rendle.

DistriputTion: Mexico, West Indies, Central America, and South
America to Argentina, in thickets, llanos, fields, savanna and woods,
at 0-3,150 meters elevation.

12. Melochia pilosa (Mill.) Fawc. & Rendle, Fl. Jamaica 5:164. 1926.
Sida pilosa Mill. Gard. Dict. ed. 8. 1768.
Melochia venosa Sw. Prodr. Veg. Ind. Occ. 97. 1788.
Mougeotia polystachya H. B. K. Nov. Gen. & Sp. 5:328, pl. 483. 1823.
Riedleia polystachya (H. B. K.) DC. Prodr. 1:490. 1824.
Riedleia Jamaicensis DC. Prodr. 1:492. 1824.

242 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Melochia Jamaicensis Balb. ex DC. Prodr. 1:492. 1824.

Riedleia venosa (Sw.) DC. Prodr. 1:492. 1824.

Melochia sericea St.-Hil. Fl. Bras. Mer. 1:160. 1825.

Melochia ulmarioides St.-Hil. Fl. Bras. Mer. 1:159. 1825.

Visenia polystachya (H. B. K.) Spreng. Syst. 3:30. 1826.

Visenia venosa (Sw.) Spreng. Syst. 3:30. 1826.

Visenia sericea (St.-Hil.) Spreng. Syst. 4:Cur. Post. 248. 1827.

Visenia Ulmariae Spreng. Syst. 4:Cur. Post. 248. 1827.

Riedleia virgultosa Macfad. Fl. Jamaica 1:104. 1837.

Riedleia interrupta Schlecht. Linnaea 11:375. 1837,

Riedleia sericea (St.-Hil.) Steud. Nom. II, 2:460. 1841.

Riedleia Ulmariae (Spreng.) Steud. Nom. II, 2:460. 1841.

Melochia gracilis St.-Hil. & Naud. Ann. Sci. Nat. II, 18:35, 1842.

Melochia nitidula St.-Hil. & Naud. Ann. Sci. Nat. I], 18:35, 1842.

Melochia polystachya (H. B. K.) Triana & Planch. Ann. Sci. Nat. IV,
17:341. 1862.

Melochia interrupta (Schlecht.) Hemsl. Biol. Centr. Amer. Bot. 1:131. 1879.

Riedleia argentea Poepp. ex Schum. in Mart. Fl. Bras. 12(3) :37. 1886, in
synonymy.

Melochia betonicaefolia Ruiz & Pavon ex Schum. in Mart. Fl. Bras. 12(8) :37.
1886.

Melochia paniculata Willd. ex Schum. in Mart. Fl. Bras. 12(3):37. 1886.

Melochia venosa Sw. var. typica Schum. in Mart. Fl. Bras. 12(3) :37. 1886.

Melochia venosa Sw. var. polystachya (H. B. K.) Schum. in Mart. Fl. Bras.
12(3):37. 1886.

Melochia venosa Sw. var. betonicifolia Schum. in Mart. Fl. Bras. 12(3) :37.
1886.

Melochia venosa Sw. var. sericea (St.-Hil.) Schum. in Mart. Fl. Bras, 12(3) :38.
1886.

Melochia venosa Sw. var. Widgrenii Schum. in Mart. Fl. Bras. 12(3):38. 1886.

Melochia ‘nervosa’ (=venosa Sw.) 6 polystachya (H. B. K.) Schum. var.
flava O. Ktze. Rev. Gen. 3(2):25. 1898.

Melochia venosa Sw. var. squarrosa Schum. & Hassl. Bull. Herb. Boiss. H,
4:71, in 1903 for 1904.

An erect herb or suffrutex, sometimes frutex, 0.4-3.0 (4.0) meters
high, simple or branched, the branches slender, to 15 cm. long, some-
times 2 or more shoots from near the base, the root thickened, to 2.5
times the diameter of the base of the stem, 0.8-1.2 em. wide, woody;
young stems sometimes hollow, hirsute, pilose or tomentose, some-
times woolly, hairs tan or golden, sometimes ferruginous, mostly
simple, sometimes also glandular and a few forked or stellate, 0.1-2.2
mm. long, internodes 0.8-8.0 cm. long; stipules deciduous, (3-) 4-10
mm. long, 0.7-4.0 mm. wide, deltoid-acuminate, ovate-acute, or lan-
ceolate, sometimes slightly oblique, sericeous, ciliate, hairs simple, 0.2-
1.5 mm. long; petiole 0.2-2.7 (-5.0) em. long, pubescence similar to
that of the young stems.

Lamina 2.2-9.5 (-11) em. long, 0.9-7.5 cm. wide, ovate, oblong-
ovate, ovate-lanceolate, lanceolate, or subelliptic, the base rounded
to subcordate, the apex acute to somewhat rounded, leaves im-

GOLDBERG—GENUS MELOCHIA 243

mediately below the inflorescence reduced, the upper surface usually
green, sericeous, hirsute or puberulous, sometimes glabrescent, some-
times scabrous or woolly to the touch, hairs simple, sometimes a
few forked or stellate, 0.2-1.0 mm. long, the lower surface green, or
covered with brownish, golden or ferruginous hairs, sericeous or
pilose on the veins, hairs simple, to 1.3 mm. long, sometimes densely
tomentulose between the veins with hairs erect, stellate, forked, and
simple, rarely also glandular, 0.2-0.7 mm. long, velvety or woolly to
the touch, sometimes glabrescent, the margin irregularly crenate-serrate,
serrations 0.6-4.8 mm. wide, 0.3-2.7 mm. high, primary veins numer-
ous, (7-) 8-11 (-18) pairs, 2 or 3 are basal, more or less parallel,
straightish and ascending at a rather acute angle (about 40°), im-
pressed above, prominent beneath.

Inflorescences principally terminal, extending (2—) 10-20 (-31) em.
beyond the first leaf, at 3-11 nodes, paniculate, contracted or open,
sometimes spicate, the flowers tending to be more numerous on the
upper side of the brachlets, in subcapitate umbels of (1—-) 3-12 flowers,
the branchlets 1.5-3.5 em. long, usually also axillary, 1-11.8 em. long,
in the axils of 1-5 or more reduced leaves, at 1-3 nodes, the peduncle
0—-0.3 cm. long, the pedicel usually short, 1-3 (-5) mm. long; bracts
usually separate, rarely 2 or 3 united at the base, 1.5-5.0 mm. long,
0.2-2.1 mm. wide, deltoid-acute to deltoid-acuminate, ovate, lanceolate
or linear, ciliate, sericeous, hairs simple, 0.1-1.0 mm. long.

Calyx (4.0-) 4.6-7.5 mm. long, (2.5—-) 2.9-5.3 mm. wide, not accres-
cent, external surface hirsute, sericeous, or tomentulose, hairs simple,
sometimes also glandular, stellate or forked, 0.1-1.0 mm. long, the
internal surface densely puberulous at the apex, the teeth 2.5-4.7 mm.
long, 0.9-2.2 mm. wide, deltoid-acuminate or deltoid-acute, the sinus
acute or narrowly rounded; petals usually yellow, sometimes orange,
5.9-10.5 mm. long, 2.0-4.5 mm. wide, obovate to oblanceolate, some-
times cuneate, the claw 1-2 mm. long, attached to the staminal tube
for 0.5-1.0 mm.

Longistylous form. Stamens 3.4-5.3 mm. long, the filaments united
up to the anthers, the anther 0.9-1.5 mm. long, 0.5-0.7 mm. wide,
oblong, emarginate at both ends, nearly to the middle from the apex;
pistil 4.6-8.3 mm. long, the styles 2.6—-5.0 mm. long, free to the ovary
or united for 0.5 mm., stigmatic for 0.5-1.0 mm., the ovary oblong,
ellipsoid, ovoid or globular, densely hirsute-sericeous at the apex, less
so below, hairs 0.2-1.5 mm. long, stipe none.

Brevistylous form. Stamens (5.1—-) 6.3-7.5 mm. long, the fila-
ments free for 2.64.2 mm., glabrous, the anther 1.1-1.3 mm. long,
0.6—-0.8 mm. wide, shape same as that above, sometimes very small
staminodal teeth present; pistil (3.3-) 3.7-4.7 mm. long, the styles
(1.1—-) 1.7-2.2 mm. long, free to the ovary or united for 0.2-0.5 mm.,

244 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

stigmatic for 0.3-0.5 mm., the ovary globular-ovoid, densely hirsute-
sericeous, hairs simple, 0.2-1.0 mm. long.

Fruit globular or suboblong, obtusely pentagonal, (3.7-) 4.2-6.0
mm. long, (3-) 4.2-5.0 mm. wide, usually slightly longer than wide,
the rostrum 0.3 mm. long, hirsute-tomentulose, hairs simple, 0.3-1.1
mm. long, sometimes also forked and stellate, rarely glandular, 0.1-
0.2 mm. long, the sulci between the carpels very shallow, extending
a fifth to a fourth the way to the center of the fruit, the base sometimes
narrowed into a stipe 0.5 mm. long; dehiscence all along the ventral
suture and a third to half way along the dorsal suture, eventually
also septicidal, the fruit falling apart; seeds 1.6-2.7 mm. long, 1.1-1.6
mm. wide, dark brown, 1 or 2 per cell.

Flowering and fruiting specimens collected throughout the year.

Type Locaity: Jamaica. Miller’s type is in the herbarium of
the British Museum.

Disrripution: Vera Cruz, Mexico, Jamaica, Venezuela, Colombia,
southern and western Brazil, Ecuador, Peru, Bolivia, Paraguay and
northern Argentina, in campo and thickets, usually in dry, open
habitats, in savanna, on hillsides, in rocky, calcareous soil, at 224—
2,000 meters elevation, usually above 500 meters.

MEXICO: Vera Cruz: Hacienda de la Laguna, Schiede s.n. (P, isotype of
Riedleia interrupta Schlecht.). F neg. 35381 ex P of type of Riedleia interrupta
Schlecht. (US). ?VeRa Cruz: Pacho (?Pacho Viejo), Liebmann 11874 (US).

JAMAICA: Swartz s.n. (8, type of Melochia venosa Sw., also US neg. 5606).
Balbis s.n. (8). Constitution Hill, Harris 6815 (US), 9026 (US). Along the
road to Newcastle, Rishbeth s.n. (1).

VENEZUELA: Vogl 334 (8). Mrranpa: San Blas?, Gines 4414 (US).
Feperau Disrricr: Caracas and vicinity, Pittier 6151 (US); Curran & Haman
1063 (US); Martin? (8S). Road from Caracas to La Guayra, Pultier 5898 (US),
9557 (US). Lower Catuche wood above Caracas, Pittier 7027 (US). Lower
Cotiza near Caracas, Pittier 7888 (MICH, US). Rio Catuche, Eggers 13062 or
13162, July 9, 1891 (L). Araaua: Colonia Tovar, La Victoria, Fendler 111
(GH), 1929 (GH). Cosxrpxs: San Carlos, Velez 2753 (US). Mérrpa: Tabay,
Gehriger 444 (US).

COLOMBIA: Triana s.n. (W). Macpa.ena: Sierra Nevada de Santa
Marta, Barkley & Araque M. 38 (US). Between Pueblo Viejo and San Miguel,
Seifriz 354 (US). Santa Marta, H. H. Smith 471 (L, 8, US). Norrs pp San-
TANDER: Ocafia, Schlim 73 (P). Sanranper: Upper Rio Lebrija valley, NW.
of Bucaramanga, Killip & Smith 16307 (US). CuNDINAMARCA: Sasaima,
Bro. A. Marie 20E (US). Torma: Honda, Humboldt & Bonpland 1803 (P,
isotype of Mougeotia polystachya H. B. K., also US neg. 5669). Mariquita,
Pennell 3627 (US). Huma: E. of Neiva, Rusby & Pennell 496 (US). San
Antonio Fortalecillas, Fosberg 19837 (US). Antioquia: Bello, Archer 138 (US).
Caupas: Anserma, Pennell 10637 (US). Supia, Pennell 10696 (US). VALLE
pEL Cauca: Palmira, Pennell & Killip 6033 (US). Puerto Caldas, Kullep &
Hazen 11033 (US). Cauca: Rio Patia, von Sneidern 2401 (8, US).

ECUADOR: “Yapio cerca Quilichao,”’ Jameson s.n. (US).

PERU: Mathews 3307 (8). Vitoe and Chacahuassi, F neg. 9609 of Ruiz

GOLDBERG—GENUS MELOCHIA 245

s.n. ex B, presumably of type of Melochia venosa Sw. var. betonicifolia Schum.
(US). Casamarca: Jaen, Weberbauer 6200 (US). Valley of Tabaconas River,
Weberbauer 6244 (US). San Marrin: Tarapoto, Ferreyra 5104 (US). Alto
Rio Huallaga, Ll. Williams 6685 (US), 5802 (S). Loreto: Near Tarapoto,
Ule 6436 (L). Hudnuco: Cuchero, Poeppig addenda 71 (W), 1269, in part
(W). Cusco: Lares Valley between Calea and Pasto Grande, Weberbauer
7932 (8, US).

BOLIVIA: Chiquiacé, Fiebrig 2722 (L). Florida and Palissa, Herzog 1294
(L, 8). Ixiamas, Cardenas 1916 (MICH). Guanai, Rusby 1846 (MICH).
Yungas and vicinity, Rusby 1847 (MICH); Bang 517 (MICH, US), 2810 (MICH,
8S, US). Norra Yuneas: Milluguaya, Buchtien 789 (S), 6021 (S). Bent:
Trinidad-Missiones Guarayos, Werdermann 2504 (S). Santa Cruz: Sara:
Buena Vista, Steinbach 7413 (S).

BRAZIL: Pohl 560 or 160 (W). Minas Gerais: Near Sao Antonio de Rio
to Cima not far from Villa Rica (=Ouro Preto), St.-Hilaire 2295 (P, isotype of
Melochia sericea St.-Hil., also US neg. 5673). Caldas, Regnell I 20 (R, 8, US),
I 20a (US); Widgren 245 (GH, 8); Lindberg 285a (S), 284a (S). Corinto, Fazenda
do Diamante, Mexia 5549 (GH, MICH, R, 8, US). Ituiutaba, Macedo 4159
(US). Itabira do Campo, Schwacke s.n. (R). Belo Horizonte, Colonia Bias
Fortes (Jardim Bot. Bel. Horiz. 5356). Belo Horizonte, Barreto 7862 (R), 10380
(R). Caxambu near das Aguas, Glaziow 15848 (R). Mantiqueira pelo vale do
rio Verde até Trés Coragdes, de Saldanha 8724 (R). Lagoa Santa, Warming
s.n. (8). Between Sitio and Dr. Sa Fortes, Serra Mantiquerra, Chase 8675 (GH).
SANTA LUZIA: Venda Nova, L. O. Williams & Assis 6511 (US). CAMPINA
VERDE: Manga, Macedo 362 (US). Gords: Near the city of Goids, Burchell
6993 (K). Sao Domingos, Gardner 4105 (GH). Corallinho, Pohl 1310 (W).
Caretao, Pohl (W). Javat: Queixada, Macedo 1549 (US). Mato Grosso:
Coxipo da Ponte, Hoehne 3026 (726)(R). Maribondo St. Laurengo, Hoehne
2995 (721)(R). Tapirapoan, Hoehne 1511 (R). Porto Esperidido, Jauri, Hoehne
704 (327)(R). Casa da Pedra, Armond (R 78143). Coxipo Mirim near Cuyaba,
Malme s.n. (S). S&o Pauto: Burchell 5618 (K). Itu, Riedel 1993 (W). Near
Mogyguassi Mosén 1127 (5). ParanA: Jaguariahyva, Dusén 16172 (8, US);
Jonsson 353a (S). Restinga Secca, Dusén, 4238 B(R). Seugés-Fabio Rego,
Dusén 10979 (8). Capao Grande, Dusén 9167 (S). Morungava, Dusén 17384
(S). Itaperussi, Dusén 13855 (S). tTrBaci: Fda. Monte Alegre, Harmonia,
Hatschbach 2872 (US), 2972 (US). campo MouRAo: Campo Mourao, Hatschbach
8770 (US). Santa Catarina: Rio Capinzal, Dusén 17851 (GH, 8). curt-
TIBANOS: Butia campo, 19 km. E. of Lebon Regis, L. B. Smith & Klein 11052
(US). Curitibanos, Miller 83 (R). cagapor: 31 km. W. of Cagador, L. B.
Smith & Klein 10921 (US). campos Novos: 33 km. SE. of Campos Novos,
L. B. Smith & Klein 11205 (US). Tle Sainte Catherine Gaudichaud s.n. (P).
Rio GRANDE vo Suu: Santa Maria, Allenoa Vidal (R 78201, US); ?Vida (R);
Malme 1236 (S). Séo Leopoldo, Leite 2518 (A); Rambo (PACA 1161), (PACA
1289); Theissen 398 (PACA), 582 (PACA), (PACA 25115); Ritter (PACA 35405).
Sapucaia near Sio Leopoldo, Rambo (PACA 38174), (PACA 40768), (PACA
44277). Cristo Rei near Sio Leopoldo, Rambo (PACA 46206). Vicinity of
TUPANCIRETA: Jari, Rambo (PACA 9142), (PACA 9485). IZjusinho, Rambo
(PACA 9989). At Ivaf River, Rambo (PACA 9474). Vicinity of sio Lurz:
Caaré, Rambo (PACA 53454). Cerro Largo, Sehnem (PACA 48091). Lagoa
Vermelha and vicinity, Friderichs (PACA 11134), (PACA 11859). Passo do
Socorro, Rambo (PACA 51506). Nonoai near Uruguay River, Rambo (PACA
28272). Cachoeira, Malme 988 (S). Santo Angelo, Rambo (PACA 53001).
Kappesberg near Montenegro, Sehnem (PACA 48376). Sao Borja, Rambo (PACA

246 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

3029). Palmeira, Rambo (PACA 50011). Neu-Wiirttemberg (Panambi), Born-
mueller 433 (GH). Near an abrupt descent, called Salto, in the swifty flowing
Uruguay River, St.-Hilaire 2526 (P, isotype of Melochia ulmartoides St.-Hil.,
also F neg. 35387 ex P, type of Melochia ulmariotdes St.-Hil. (US).

URUGUAY: Cufiapird (Brazilian frontier), Rambo (PACA 4024).

PARAGUAY: San Luis between Rio Apa and Rio Aquidaban, Fiebrig 4412
(GH, K, L). Upper part of Rio Apa, Hassler 8058 (GH, 8, isotypes of Melochza
venosa Sw. var. squarrosa Schum. & Hassl.). Near Bellavista (Rio Apa), Hassler
8184a (GH, 8). Villarrica, Jorgensen 3810 (8S, US). Guarapi, Balansa 3254
(L, P). In valley of Y-acd River, Hassler 6587 (GH, 8). Near Igatimi, Hassler
5529a(S). Est. Caballero, Anisits 374 (8). Est. Santa Maria, Antsits 2412 (S).
Central Paraguay, Morong 403 (US).

ARGENTINA: SALTA: ROSARIO DE LERMA: Campo Quijana, Venturt 8213
(GH). TucumA&n: Mufiecas, Schreiter 1935 (8S). ramarnia: San Pablo, Venturt
1175 (8S, US). caprrau: Rio Sali, Venturt 1175a (GH, US). Tart: Cumbre de
Taficillo, Venturi 6047 (US). BuRRoyAco: Cerro del Campo, Venturt 10326
(S). Formosa: Laishi, Jorgensen 2324 (GH, US). Mustonns: Posadas, Bon-
pland, ‘La Granja,’ Ekman 181 (S), 182 (8, US). SAN IGNACIO: San Ignacio,
Vattuone & Bianchi 143 (US); Montes 479 (UC). Gobernador Roca, Montes
562 (UC). cANDELARIA: Puerto Santa Ana, Schwarz 623 (UC). Loreto, Montes
439 (UC). Piaido Finlandesa, Schwarz 690 (UC). Arroyo ‘‘Martin chico,”
Schwarz s.n, (UC).

Because the differences between the varieties described in this
species are so minor, and the transition between them so gradual,
and because the varieties are more or less sympatric, I do not con-
sider the varieties worthy of retention. However, in general, the
species is hairier in the southern portion of its range than in the
northern.

This species is most closely related to M. hassleriana Chod., but
the pubescence, peduncles, and pedicels are generally longer in the
latter species. AZ. pilosa usually has a longer terminal inflorescence,
a ereater total number of flowers, and more per cluster, and the
flowers tend to be more frequent on the upper side of the branchlets;
the flowers are rather evenly distributed in M. hassleriana.

M. pilosa is also related to M. chamaedrys St.-Hil. The major
differences between them are mentioned under the description of the
latter species.

13. Melochia hassleriana Chod. Bull. Herb. Boiss. II, 1:403. 1901.
Melochia ulmarioides St.-Hil. f. sericea, f. Hassleriana (Chod.), and f. brevi-
pedunculata, all Schum. & Hassl. Bull. Herb. Boiss. HW, 4:71. 1903.
Melochia ulmarioides St.-Hil. f. albo-villosa Rk. EK. Fries, Svensk. Vet. Acad.
Handl. 42(12):10. 1908.

An ascendent or procumbent suffrutex 0.5-1.5 meters high, with
several stems from the base, the root woody, thickened, 0.7-1.0 cm.
wide, the young stems and petioles pilose, hairs simple, straight,
1.5-4.0 mm, long, also shorter, recurved, 0.8-0.5 mm. long, internodes
1.5-7.0 em. long; stipules 6-10.38 mm. long, 0.8-1.0 mm, wide, lance-

GOLDBERG—GENUS MELOCHIA 947

olate, ciliate and pilose or sericeous, hairs simple, 2-3.5 mm, long;
petiole short, 0.1-1.0 cm. long.

Lamina 2.5-5.5 cm. long, 0.9-3.0 em. wide, lanceolate to ovate,
the base rounded, the apex acute, the upper surface sericeous-pilose,
hairs tan, simple, 1.3-4.0 mm. long, the lower surface sericeous-pilose
on the veins, hairs tan, simple, 1.74.0 mm. long, usually also stellate,
0.2 mm. long, between the veins, the margin long-ciliate, irregularly
crenate-serrate, serrations 1.2-5.2 mm. wide, 0.6-1.5 mm. high, 8-13
pairs of straight, parallel primary veins, 2 are basal.

Inflorescences axillary, 3.5-5.5 cm. long, in the axils of the upper
2-4 leaves, also terminal, extending 3-6.5 em. beyond the first leaf,
the peduncle 0.5-3.0 cm. long, the ultimate divisions umbellate, 2-5
flowers per umbel, more than one umbel per peduncle, the pedicels
2.5-4.0 mm. long at anthesis, up to 6.0 mm. long in the fruiting state;
bracts 3.5-9.5 mm. long, 0.3-1.5 mm. wide, lanceolate to nearly
linear, ciliate and sericeous, hairs simple, 1-3 mm. long.

Calyx 6.3-7.8 mm. long, 3.4-4.4 mm. wide, not accrescent, pilose-
sericeous and ciliate, hairs simple, 1-4 mm. long, also stellate, 0.1
mm. long, the teeth 3.7-5.5 mm. long, 1.4—-1.8 mm. wide, deltoid-
acuminate, the sinus acute; petals yellow to orange, 10-11.5 mm.
long, 3.2-3.4 mm. wide, obovate-cuneate, attached to stamens for
1-1.5 mm., the claw 1 mm. long.

Longistylous form. Stamens 5.0 mm. long, the filaments united up
to the anthers, the anther 1.2-1.3 mm. long, 0.6 mm. wide, oblong,
emarginate at both ends, nearly to the middle from the apex; pistil
8.8 mm. long, the styles filiform, papillose for 1.2 mm. at the apex,
free for 4.6 mm., united for 1.5 mm., the ovary ovoid, pentagonal,
sericeous, hairs simple, 1-2 mm. long.

Brevistylous form. Stamens 7.4-7.8 mm. long, the filaments free
for 4-5 mm., the anther 1.1-1.6 mm. long, 0.7 mm. wide, shape same
as that above; pistil 5.5-6.0 mm. long, the styles papillose for 0.7 mm.
at the apex, 2.2-2.5 mm. long, free to the ovary, the ovary ellipsoid to
globular, pentagonal, sericeous, hairs simple, 1-2.3 mm. long.

Fruit 5 mm. long, 4.8 mm. wide, remnant of styles 1 mm. long,
globular, obtusely pentagonal, the sulci between the carpels rather
shallow, extending a fifth to a fourth the way to the center of the fruit,
sericeous, hairs simple, 1-2 mm. long, also stellate, 0.1 mm. long, the
stipe 0.5 mm. long; dehiscence all along the ventral suture and half
way along the dorsal suture, also slightly septicidal; seeds 2.1-2.5
mm. long, 1.3-1.6 mm. wide, dark brown, 1 or 2 per cell.

Flowering specimens collected in October and December to March;
fruiting ones, from December to March.

Typ Locatity: The type was collected by Hassler, no. 1739, in
thickets near Itacurubi, Paraguay. An isotype is cited below.

248 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

DistripuTion: Paraguay and Parand, Brazil, in campo, in sandy
fields, in arid rocky places, at about 790 meters elevation.

BRAZIL: Parani: Capdo Bonito, Dusén 16903 (8).

PARAGUAY: Near Itacurubi, Hassler 1739 (NY, isotype). Along the road
between Caacupé and Tobaty, Archer & Rojas 4856 (US). Road to Paraguay
River, Hincinde ad Monte Calvario prope Villa occidental, Lorentz 33 (US).
Capibary, Hassler 5917 (8). River Apa, Hassler 8074 (5, type of Melochia
ulmarioides St.-Hil. f. sericea Schum. & Hassl.). Sierra de Maracaju, Hassler 5596
(S). In altaplanitie “Sierra de Amambay,” Hassler 12038 (S). Near Igatimi,
Hassler 4793 (S, type of Melochia ulmarioides St.-Hil. f. brevipedunculata Schum, &
Hassl.), (W). Est. Armonia, Anisits 1913 (8, type of Melochia ulmarioides St.-Hil.
f. albo-villosa KR. E. Fries). San Luis between Rio Apa and Rio Aquidaban,
Fiebrig 4451 (L).

This species has been misinterpreted as Melochia ulmarioides St.-Hil.
I have seen the isotype of the latter species and consider it to be the
same as Melochia pilosa (Mill.) Fawce. & Rendle.

This species is close to Melochia pilosa (Mill.) Fawe. & Rendle but
differs in that the margins of the leaves are long-ciliate, the stems
are usually less hairy, and the flowers are fewer and more loosely
clustered. M. pilosa usually has a larger terminal inflorescence than
M. hassleriana, and the flowers tend to be more frequent on the upper
side of the branchlets but more evenly distributed in M. hassleriana.

14. Melochia chamaedrys St.-Hil. Fl. Bras. Mer. 1:161. 1825.

A small, procumbent or ascending, spreading herb to suffrutex
0.1-0.5 meter high, sometimes simple but usually with several slender
shoots (2-10) from the root, the taproot thickened, woody, sometimes
branched, 0.5-2.2 cm. wide, the young stems hirsute or rather densely
covered with straight simple hairs 0.1-2.0 mm. long, curved simple
ones 0.2-0.3 mm. long in a longitudinal line, and glandular ones 0.2—0.6
mm. long, internodes 1-3.3 (—7.5) cm. long; stipules 2.5-5.0 (-8.8)
mm. long, 0.5-1.0 (-5.0) mm. wide, lanceolate, deltoid-acuminate or
ovate, mostly ciliate, hairs simple, 0.2-2.0 mm. long, occasionally a few
glandular, 0.2-0.3 mm. long; petiole 0.2-1.5 (-2.7) cm. long, indument
sunilar to that of the young stems.

Lamina 1-4.3 (-6.5) cm. long, 0.8-3.0 (-4.5) cm. wide, ovate to
nearly orbicular, rarely also oblong and lanceolate, the base rounded
to cordate, the apex usually obtuse, sometimes also acute, the upper
surface green, pilose, hairs scattered to rather dense, somewhat
appressed, simple, usually also a few glandular, 0.1-1.6 mm. long,
the lower surface green, similarly pubescent but more densely so,
rarely stellate hairy, hairs 0.1-1.8 mm. long, the margin slightly
irregularly crenate to crenate-serrate, crenations or serrations 0.5-3.2
mm. wide, 0.4-2.0 mm. high, 5-11 pairs of primary veins, 2 are basal.

GOLDBERG—GENUS MELOCHIA 249

Inflorescences axillary, (1.2-) 1.6-4.5 em. long, in the axils of the
upper 1-6 (-8) leaves and occasionally also terminal, extending 3.5
cm. beyond the first leaf, the ultimate divisions umbellate, few-
flowered, 2-4 (-5) per umbel, the peduncle slender, 0.1-2.5 cm. long,
sometimes absent, sometimes with a branch 0.4-0.5 em. long, the
pedicels slender, 1.6-9.3 mm. long, mostly glandular, hairs 0.1-0.4
mm. long; bracts, 0.6-4.0 (-8.3) mm. long, 0.1-1.3 (-4.8) mm. wide,
lanceolate to ovate or linear, mostly ciliate, hairs simple, 0.1-1.7 mm.
long.

Calyx 4.6-7.8 (-8.5) mm. long, 3.1-4.8 (-5.1) mm. wide, not ac-
crescent, sericeous and ciliate, hairs simple and usually also glandular,
0.1-0.8 (-1.6) mm. long, the teeth 2.5-5.0 mm. long, 1.2-1.8 (2.5)
mm. wide, deltoid- to ovate-acute or ovate-acuminate, the sinus
acute; petals yellow to orange, Weberbauer no. 6220 from Peru was
rose with brown veins according to Macbride (39), 7.2-11 mm. long,
2.7-4.1 mm. wide, obovate-cuneate, attached to the staminal tube
for 0.5-1.0 mm., the claw 0.9-1.8 mm. long, 5 minute glands some-
times present between the bases of the petals.

Longistylous form. Stamens 3.8-4.2 mm. long, the filaments united
up to the anthers or separate for 0.5 mm., the anther 0.9-1.2 mm.
long, 0.5-0.8 mm. wide, oblong, emarginate at both ends, nearly to
the middle from the apex; pistil 4.8-7.5 mm. long, the styles filiform,
free for 2.0-5.3 mm., united for 0.3-1.0 mm., papillose at the apex
for 1 mm., the ovary ovoid to globular, pentagonal, hirsute to sericeous
at the apex, hairs simple, 0.1-1.2 mm. long.

Brevistylous form. Stamens 6.1-8.2 mm. long, the filaments free
for 3.2-5.1 mm., the anther 1.1-1.6 mm. long, 0.6-1.1 mm. wide,
shape the same as that above; pistil 4-5 mm. long, the styles 2 mm.
long, free to the ovary or united for 0.7 mm., papillose for 0.3-0.5
mm. at the apex, the ovary ovoid, pentagonal, hirsute-sericeous,
mostly near the apex, hairs 0.2-1.3 mm. long.

Fruit 4.4-5.6 mm. long, 4.4-4.8 (-5.3) mm. wide, globular, some-
times suboblong, obtusely pentagonal, the sulci between the carpels
shallow to moderately deep, extending a fifth to a third the way to
the center of the fruit, covered with simple hairs 0.1-1.5 mm. long,
sometimes a few forked and glandular hairs 0.1-0.4 mm. long, the
base narrowed to a stipe 0.5-1.0 mm. long; dehiscence all along the
ventral suture, a third to half way along the dorsal suture, and
slightly septicidal; seeds 1.5-2.6 mm. long, 1.1-1.5 mm. wide, trigonal,
dark brown, rugulose, 2 per cell.

Flowering specimens collected in August, September, November to
February, April, and May; fruiting ones, in September, November
to February, and April.

250 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Key to Varieties of M. chamaedrys

1. Lamina suborbicular, 1-2.3 em. long, 0.9-2.7 cm. wide, apex obtuse, crenations
up to 2 mm. high; pedicels 2.5-9.3 mm. long.
14a. M. chamaedrys var. chamaedrys
1. Lamina ovate, 1.2-4.5 (-6.5) em. long, 0.8-3.0 (-4.5) em. wide, apex usually
acute, crenations up to 1.2 mm. high; pedicels 1.6-3.0 mm. long.
14b. M. chamaedrys var. decumbens

14a. Melochia chamaedrys St.-Hil. var. chamaedrys
Riedleia Chamaedrys (St.-Hil.) G. Don, Gen. Syst. 1:529. 1831.
Melochia chamaedrys St.-Hil. f. typica Ekman, Arkiv. Bot. 9(4):46. 1910.
Melochia chamaedrys St.-Hil. var. loretensis and var. viscidula, both Hassl.
Addenda Pl. Hassl. 12. 1917 (without description).

Commonly, this variety is 0.1-0.3 meter long; stipules, 2.5-3.6 mm‘°
long, 0.5-0.8 mm. wide; petiole 0.2-0.9 em. long; lamina 1-2.3 cm.
long, 0.9-2.7 cm. wide, nearly orbicular, the apex obtuse, the crenations,
up to 2.0 mm. high, 5-7 pairs of primary veins; inflorescences 1.6—3.5
cm. long, the peduncle 0.6-1.7 cm. long, the pedicels 2.5-9.3 mm.
long, the bracts 0.6-2.5 mm. long, 0.1-0.5 mm. wide.

TypE LOCALITY: The type was collected by St.-Hilaire in herbaceous
fields near the estate commonly called Estancia do Salto S. Itaroro,
not far from the village of S. Francisci Borjensis, Misiones, Argentina.
The isotype is cited below.

Distrisution: Peru, Brazil, Uruguay, Paraguay, and Argentina,
in shrubby and herbaceous fields, in dry compo, on hills among dry
calcareous rocks, and in thickets among rocks, at 60-800 meters
elevation.

PERU: Casamarca: JAEN: Near the confluence of Chinchipe and Marafion
Weberbauer 6220 (US).

BRAZIL: Banta: Catinga de Moura, Schreiner in 1890 (R 78127). Maro
Grosso: Capio Bonito, Archer & Gehrt 86 (US). PARANA: PARANA: Campo
Mour&o: Rio da Vargem, Hatschbach 8757 (US). Sanra Catarina: Campos
Novos: 34 km. E. of Joagaba, L. B. Smith & Klein 11181 (US). Rio GRranpDE
po Sunt: Campinas near Sta. Rosa, Spies (PACA 36029). Caaré near S. Luiz,
Rambo 3056 (PACA), (PACA 53298), (PACA 53433). Cerro Largo near S. Luiz,
Buck (PACA 11136). Livramento, Rambo 3950 (PACA). Parecf near Monte-
negro, Rambo (PACA 42973).

URUGUAY: Satto: Arapey, Herter 2293 (5).

PARAGUAY: Villa Encarnacion, Rojas 7924 (8). Itapé, in canyon at the
foot of Cordillera de Villarrica, Jorgensen 4477 in part (S), 4477 (A, US).

ARGENTINA: CorRIENTES: SANTO TOMS: Est. ‘“Garruchos,’”’ Pedersen 971
(S, US). Mursrones: Near Est. do Salto 8. Itaroro near S. Francisci Borjensis,
St.-Hilaire (P, isotype). ?Mistones: F neg. 9600 ex B, presumably an isotype
of the species. Posadas, Ekman 183 (5S), 184 (S).. CANDELARIA: Loreto, Montes
560 (S).

This species is close to Melochia pilosa (Mill.) Fawe. & Rendle, but
the latter is more robust. MV. pilosa is erect and 0.43.0 meters high;

GOLDBERG—GENUS MELOCHIA 251

M. chamaedrys is procumbent or ascending and 0.1-0.5 meter high.
The leaves are often suborbicular, those of M, puosa are more elongate,
usually ovate to lanceolate, and larger. The inflorescences are pri-
marily axillary but primarily terminal in M. pilosa.
14b. Melochia chamaedrys St.-Hil. var. decumbens (Fries) A. Goldberg, comb.
et. stat. nov.
Melochia decumbens R. E. Fries, Svensk. Vet. Akad. Handl. 42(12):11. 1908.
Melochia chamaedrys St.-Hil. f. decumbens Ekman, Arkiv. Bot. 9(4):46.
1910.

This variety is more robust than the typical variety. It is up to 0.5
meter long; stipules 4-5 (-8.8) mm. long, 0.7-1.0 (-5.0) mm. wide;
petiole 0.2-1.5 (-2.7) em. long; lamina more densely hairy than that
of the typical variety, 1.2-4.5 (-6.5) em. long, 0.8-3.0 (4.5) cm. wide,
ovate, the apex usually acute, crenations to 1.2 mm. high, 5-11 pairs
of primary veins; inflorescences 1.2-4.5 cm. long, the peduncle (0-)
1-2.5 cm. long, the pedicel 1.6-3.0 mm. long, the bracts 2.6-4.0
(-8.3) mm. long, 0.6-1.3 (-4.8) mm. wide.

Tyre Locauity: The syntypes were collected by <Anisits, nos.
2509 and 2656, at Estancia Santa Maria, Paraguay. The latter is
selected as lectotype.

Distripution: Paraguay and Argentina on dry, calcareous,
rocky hills.

PARGUAY: Estancia Santa Maria, Anisits 2509 (S, paratype), 2656 (S,
lectotype, also US neg. 5617, syntypes of Melochia decumbens Fries),

ARGENTINA: Misiones: Posadas, Bonpland, Lillieskéld s.n. (S, also US
neg. 5618).

Ekman (see synonymy) considered Melochia decumbens Fries to be
a form of Melochia chamaedrys St.-Hil. I consider it sufficiently
distinct to have varietal status.

15. Melochia kerriifolia Triana & Planch. Ann. Sci. Nat. IV, 17:341. 1862.
Melochia Humboldtiana Steyerm. Ficldiana, Bot. 28(2):365. 1952.

An erect, slender-branched annual herb or suffrutex 0.2-0.8 meter
high, the root not thickened, the young stems hirsute, hairs scattered
and sometimes in a longitudinal line, simple, straight, and shorter
recurved hairs and very few glandular hairs, 0.2-1.3 mm. long,
internodes 1-11 em. long; stipules 1.5-4.0 mm. long, 0.3-1.2 mm wide,
deltoid-acute to deltoid-acuminate or lanceolate, mostly ciliate, hairs
simple, 0.2-1.3 mm. long; petiole 0.1-2.3 em. long, tomentose, especi-
ally on the adaxial surface, hairs mostly simple, some forked, few
glandular, 0.5-1.8 mm. long.

Lamina 0.8-9.3 em. long, 0.2-3.1 em. wide, narrowly lanceolate
to ovate, the base rounded to subcordate, the apex acute to slightly
acuminate, the upper surface subglabrous, with a few simple, ap-
pressed or recurved hairs 0.1-1.0 mm. long on the midrib, and some-

952 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

times a few glandular hairs, especially on the margin, the lower
surface with scattered, appressed, simple hairs 0.1-1.1 mm. long on
the veins, the margins irregularly crenate-serrate to serrate, serra-
tions 0.8-4.2 mm. wide, 0.5-1.9 mm. high, 5-14 pairs of primary
veins, 1 or 2 are basal.

Inflorescences 1-21 cm. long, terminal and axillary, rarely also
leaf-opposed, in the upper 1-4 axils, or on axillary branches up to 7.3
cm. long, with reduced leaves, slender, usually paniculate, the ulti-
mate divisions umbellate, 2 or 3 (1-5) flowers per umbel, the peduncle
slender, 0.6-8.5 cm. long, the pedicels 1.5-5.0 mm. long, with scattered
glandular hairs; bracts 1-3.5 mm. long, 0.2-0.8 mm. wide, deciduous,
deltoid-acute to deltoid-acuminate or lanceolate, mostly ciliate, hairs
simple, 0.2-0.8 mm. long.

Calyx 3.0-4.7 mm. long, 2.44.0 mm. wide, not accrescent, hirsute
and sometimes also ciliate, hairs simple, some glandular, rarely a few
forked, 0.1-1.2 mm. long, the teeth 2.2-3.3 mm, long, 0.8-1.4 mm.
wide, deltoid-acuminate, the sinus acute to narrowly rounded; petals
yellow, 5.0-8.3 mm. long, 2.2-3.1 mm. wide, obovate, obovate-
cuneate, or nearly elliptic, the claw 0.5-1.0 mm. long, attached to the
stamens for 0.3-1.0 mm.

Longistylous form. Stamens 2.5-3.2 mm. long, the filaments
united up to the anthers, the anther 0.8-1.1 mm. long, 0.4-0.6 mm.
wide, oblong, emarginate at both ends, nearly to the middle from the
apex, sacs slightly divergent at the apex, rarely 6 stamens; pistil
2.8-3.5 mm. long, the styles 1.2-1.7 mm. long, free to the ovary,
filiform, papillose for 0.5-0.7 mm. at the apex, the ovary ovoid to
elobular-pentagonal, hirsute-sericeous, hairs simple and sometimes
also forked and a few stellate, 0.2-0.7 mm. long, rarely 6 carpels,
stipe none but base of carpels narrowed for 0—0.5 mm,

Brevistylous form not seen.

Fruit usually in pairs, 3.7-5.5 mm. long, 3.5-5.6 mm. wide, globular
to somewhat oblong, obtusely pentagonal, the sulci between the
carpels shallow, extending a fifth to a fourth the way to the center
of the fruit, hiruste to tomentulose, hairs simple and sometimes
forked and stellate, 0.1-1.0 mm. long, base narrowed to a stipe 0.5—
1.0 mm. long; dehiscence all along the ventral suture and a third to
half way along the dorsal suture, slightly septicidal to completely
septicidal and the fruit falling apart; seeds 1.9-2.8 mm. long, 1.2-2.0
mm. wide, brown to black, trigonal, 2 per cell.

Flowering specimens collected in May, June, October, and Novem-
ber; fruiting ones, from April to June, and in October and November.

Typm Locatiry: Savannas d’Ibagué, basin of the Magdalena,
Colombia, elevation 1,300 meters. Type collected by Triana, cited
below.

GOLDBERG—GENUS MELOCHIA 253

Disrripution: Mexico, Guatemala according to Standley «&
Steyermark (56), Venezuela, Colombia, and Brazil, in Ilanos and
fields, on rocky open limestone slopes at 20-1,300 meters elevation.

MEXICO: Veracruz: Zacuapan, Purpus 8031 (UC). Zacuapan, Fortin,
Purpus 2048 (GH, US). Camarén, Purpus 11066 (US). Oaxaca: TUXTEPEC:
Chiltepee and vicinity, Martinez-Calderén 506 (A, US).

VENEZUELA: Sucre: Between Cocollar and Guamal, Steyermark 62436
(F, isotype of Melochia Humboldtiana Steyerm., also US neg. 5671).

COLOMBIA: Maapavena: Santa Marta, H. H. Smith 484 (A,8, US). Tota
Tbagué, Mariquita, Triana (P, type, also US neg. 5659; F fragment and neg.

23860 of type; W).
BRAZIL: Near border between Pard and Goids, Burchell 9055 (GH). Maro
Grosso: Corumbd4, Malme 3042 (S).

This species is usually characterized by a loose terminal inflorescence.
The fruits usually develop in pairs and the leaves are subglabrous.
It is close to M. pilosa (Mill.) Fawe. & Rendle, but the inflorescence
is much more lax, the leaves are not as hairy, and the calyx is shorter.
The fruits of MZ. pilosa are not in pairs.

Steyermark distinguishes his Mf. Humboldtiana from M. kerritfolia
by slight differences in vestiture.

16. Melochia argentina R. E. Fries, Meded. Rijk. Herb. Leid. No. 19: 42. 1913.

Melochia argentina Hassl. Repert. Sp. Nov. Fedde 13:237. 1914.

Melochia argentina Hassl. var. genuina Hassl. Repert. Sp. Nov. Fedde
13:238. 1914.

Melochia argentina Hassl. var. lanceolata Hassl. Repert. Sp. Nov. Fedde
13:238. 1914.

A branched subshrub to shrub 0.6-2.5 meters high, the young stems
hirsute in longitudinal lines, hairs simple, curved, 0.1-0.3 mm. long,
internodes 1—3.5 cm. long; stipules 3.6-7.5 mm. long, 0.2-1.0 mm. wide,
deltoid-acuminate to nearly linear, with few simple hairs, mostly
ciliate, 0.1-0.3 mm. long; petiole 0.8-2.5 cm. long, vestiture similar
to that of the young stems, but sometimes encircling the petiole.

Lamina 2.5-7.8 cm. long, 1.5-3.6 em. wide, up to 9 by 4 cm. ac-
cording to Hassler (24), usually ovate, also ovate-lanceolate to lanceo-
late, the base rounded, the apex acute to subacuminate, the upper
surface subglabrous, hairs simple appressed, 0.1-0.6 mm. long on the
midrib and sometimes present between the veins, the lower surface
glabrous or with few simple appressed hairs 0.1-0.5 mm. long on the
veins, the margin irregularly crenate-serrate, serrations relatively
high, 2-8.5 mm. wide, 0.7-3.6 mm. high, 6-9 pairs of primary veins,
2 are basal.

Inflorescences axillary, 14.8 cm. long, at 1-14 upper nodes, rarely
also terminating axillary branches 4-13.5 cm. long, the terminal
inflorescence of a branch extending 3-4 cm. beyond the first leaf, the

227T—-051—_67——-5

254 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

ultimate divisions umbellate with 2 or 3 (3-6, Hassler) flowers per
umbel, 1-4 umbels per inflorescence, the peduncle 0-3.3 cm. long,
with a branch 0-0.5 cm. long, the pedicels 1-4 mm. long; bracts
1.3-3.7 mm. long, 0.2-0.5 mm. wide, deltoid-acuminate to acicular,
mostly ciliate, hairs simple, 0.1-0.3 mm. long.

Calyx 5.5-7.5 mm. long, 4.7-5.8 mm. wide, not accrescent, mostly
ciliate, hairs simple, 0.1-0.2 mm. long, the teeth 2.5-3.6 mm. long,
1.8-2.8 mm. wide, deltoid-acuminate or deltoid-acute, the sinus acute;
petals yellow, 8.6-9.6 mm. long, 3.5-5.0 mm. wide, 10-12 by 5-7 mm.
according to Fries, obovate-cuneate, the claw 0.5 mm. long, attached
to the stamens for 0.5 mm.

Longistylous form not seen.

Brevistylous form. Difference in length of gynoecium and androe-
cium not marked, stamens 4.8-5.6 mm. long, the filaments free for
0.5-1.5 mm., the anther 1.7-2.1 mm. long, 1.0 mm. wide, oblong,
emarginate at both ends, almost to middle from the apex, pollen
larger than average for the genus; pistil 4.2-5.1 mm. long, the styles
free for 1.5-2.2 mm., united at the base, papillose at the apex for
0.7 mm., the ovary ovoid, hirsute, hairs simple and forked, 0.1-0.7 mm.
long, stipe none.

Fruit 6.5-7.5 mm. long, 4.5-6.5 mm. wide, obtusely pentagonal,
elobose, acute to subacuminate at the apex, moderately deeply sulcate
between the carpels, but the sulci extending less than half way to the
center of the fruit, sparsely hirsute, hairs simple and forked, 0.2-
0.5 mm. long, stipe none; dehiscence all along the ventral suture and
one-third along the dorsal suture, also slightly septicidal, but capsule
not falling apart, several loose fibers about 1 mm. long along the
ventral sutures; seeds 2.6-2.8 mm. long, 1.5-1.6 mm. wide, brown,
sometimes with short white longitudinal markings.

Flowering specimens collected in October, November, and January
to March; fruiting ones, in October, November, January, and Febru-
ary.

TypE LOCALITY: Salta, Argentina, in moist places in the forest
between Embarcacion and Miraflores. Type collected by Herzog,
no. 1029, cited below.

Disrrisution: Tarija, Bolivia to Cordoba, Argentina, in moist
places in forests, thickets on hillsides, and in dry gravel of riverwash,
at 650-1,700 meters elevation.

BOLIVIA: Tarisa: La Merced at Bermejo, Fiebrig 2190 (S).

PARAGUAY: Ibytymi, Rojas 13282 (8).

ARGENTINA: Jusuy: Cerro de Zapla, Venturi 5122(US). LEepEsMA: Sierra
de Calilegua, Venturi 5348 (US). san PEDRO: About 17 km. SW. of San Pedro

de Jujuy, Eyerdam & Beetle 22573 (GH). Saura: Between Embarcacion and
Miraflores, Herzog 1029 (L; S, type; F neg. 9598 ex Bin US). Capital, Schulz

GOLDBERG—GENUS MELOCHIA 255

900 (8). ordn: Carapari, Bridarolli 3267 (GH). Obra Grande, Venturi 5576
(S, US). Tucum4n: tari: Yerba Buena, Venturi 160 (US); Wall 12 (8).

This species is characterized by its rather large fruits, among the
largest in the section Mougeotia, and very reminiscent of those in the
section Visenia, by subglabrous leaves with high serrations, and by
the nearly equal length of the gynoecium and the androecium.

From their descriptions and the fact that each cited Stuckert 12757
as belonging to his species, it is evident that Fries’ and Hassler’s
species are the same. Hassler (24) separated Melochia argentina var.
lanceolata Hassler from the type on the basis of the shape of the leaf.
Since ovate and lanceolate leaves may occur on the same plant, I do
not consider that, in this species, a variety based on the shape of the
leaf is worthy of recognition.

17. Melochia ulmifolia Benth. in Hook. Journ. Bot. 4:129. 1841.
Riedleia ramuliflora Mig. Linnaea 19:132. 1847.
Riedleja dichotoma Turez. Bull. Soc. Nat. Mose. 31(1):208. 1858.
Melochia Benthami Schum. in Mart. Fl. Bras. 12(3):40, pl 7. 1886.
Melochia carthaginensis Willd. ex Schum. in Mart. Fl. Bras. 12(3):40. 1886.
Melochia ramuliflora (Miq.) R. E. Fries, Kgl. Svensk. Vet. Akad. Handl.
42(12):11. 1908.

A scrambling shrub to small tree 1.8-4.0 meters high, the young
stems and petioles tomentulose, hairs stellate, 0.1-0.2 mm. long, the
internodes 1.8-5.5 cm. long; stipules caducous, 1.7-2.5 mm. long,
0.2-0.6 mm. wide, deltoid-acuminate to nearly linear, hirsute on the
abaxial surface, hairs 0.1-0.4 mm. long; petiole 1-3.1 em. long, slightly
broadened at the distal end.

Lamina relatively large, 5.0-14.0 em. long, 2.0-8.2 cm. wide, ovate
to oblong-ovate, the base rounded, truncate or subcordate, the apex
acute to slightly acuminate, the upper surface green, with scattered,
nonoverlapping, stellate hairs 0.1-0.2 mm. long on the veins and
usually between them, the lower surface green, puberulous, velvety
to the touch, hairs stellate, 0.2-0.3 mm. long, the margin irregularly
crenate-serrate, serrations 2-4.5 mm. wide, 1-2.5 mm. high, 6-9 pairs
of primary veins, 2 are basal, one more prominent than the other,
prominent beneath.

Inflorescences axillary, at up to 8 upper nodes, up to 6 cm. long and
5 cm. broad, open, 3-8 flowers per umbel, 3-8 umbels at each axil,
the peduncle 0.2-1.0 cm. long, sometimes with a primary branch 0.6
em. long, and a secondary one 0.3-0.7 cm. long, the pedicel 1-5 mm.
long; bracts 1-3 mm. long, 0.2-0.5 mm. wide, deltoid-acuminate to
nearly linear, hirsute-tomentulose, hairs 0.1-0.3 mm. long; flowers
very fragrant.

Calyx 4.3-6.1 mm. long, 3.7-5.7 mm. wide, campanulate, not
accrescent, tomentulose, hairs forked and stellate, 0.1-0.6 mm. long,

256 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

the teeth 1.5-2.9 mm. long, 1.3-2.5 mm. wide, deltoid-acute, deltoid-
acuminate to ovate, the sinus acute to somewhat rounded; petals
white, yellowish white or yellow, 4.4-5.5 mm. long, 1.7-2.3 mm. wide,
obovate to obovate-oblong, or oblanceolate, separate from one
another and from the androecium, the blade rather sharply demar-
cated from the claw, claw 0.5-1.0 mm. long, hairs at the base of the
adaxial surface of the blade 0.1-0.2 mm. long.

Longistylous form. Stamens 3.2-4.0 mm. long, the membrane
between the filaments very thin, hairs near the base 0.6—-0.8 mm.
long or absent, the filaments free for 0.5-1.0 mm., the anther 0.8-1.2
mm. long, 0.7-0.8 mm. wide, oblong, emarginate at both ends; pistil
4.5-5.2 mm. long, the styles 2.7-3.7 mm. long, free to the ovary,
hirsute at the base, papillose at the apex for 0.7 mm., the ovary oval
to ovoid, hirsute mostly on its angles, hairs 0.5-1.0 mm. long, stipe
none.

Brevistylous form. Stamens 5.5 mm. long, the filaments free for
3.5 mm., hairs at the base of the filaments 0.1—-0.2 mm. long, the anther
1.1 mm. long, 0.6 mm. wide; pistil 3.0-3.5 mm. long, the styles free
to the ovary, 1.7 mm. long, the ovary ovoid, hirsute, hairs up to 1.2
mm. long.

Fruit a capsule 5-6 mm. long, 4.8-5.5 mm. wide, nearly globular,
mucronate at the apex, obtusely pentagonal, very shallowly sulcate
between the carpels, hirsute-tomentulose, hairs 0.1-1.0 mm. long,
the stipe 0.2-0.3 mm. long; dehiscence all along the ventral suture and
0.2-0.25 the way along the dorsal suture, only slightly septicidal;
seeds 1.9-2.4 mm. long, 1.1-1.2 mm. wide, trigonal, brown.

Flowering specimens collected in February, May, June, and from
August to December; fruiting ones, in February, May, June, August,
and from October to December.

Type Locauity: British Guiana, in savannas. Isotypes collected
by Schomburgk, no. 203, cited below.

Disrripution: Santander, Colombia to Maranhdo, Brazil, in
second-growth thickets, along trails, on river banks, at the edge of
clearings, in fields, on loamy soil, at 80-700 meters elevation.

FRENCH GUIANA: ?Le Prieur 302 (P). Melinon 166 (P), 277 (P), 487 (P).
Banks of River Maroni, Melinon 210 (P), 229 (P, US); Sagot 1072 (P).

SURINAM: Hostmann 226 (US). Hostmann & Kappler 1047 (S, W, isotypes
of Riedleja dichotoma Turez.). Near Victoria station in region on way to Surinam
River, Kappler 1695 (P, S). Near Blaauwe Berg, Splitgerber 890 (L).

BRITISH GUIANA: Schomburgk 203 (L, isotype). Orealla, Courantyne
River, Forest Dept. 6753 (MICH, 8).

VENEZUELA: Botfvar: La Unién, Medio Caura, Ll. Williams 11235 (US).

COLOMBIA: SanTanpeER: Vicinity of Puerto Araujo on Carare River, Haught
1705 (US). Between Narifio and El Tambor, Killip & Smith 14948 (US).

BRAZIL: Rro Branco: Furo do Cujubim, Kuhlmann 126 (US). AMAZONAS:
Madeira, ?Itapisuiua, Kuhlmann 243 (RB). Par4: Baiio, Tocantins, Kuhl-

GOLDBERG—GENUS MELOCHIA 257

mann 2093 (US). Rio Cupary, affl. do Tapajoz, Kuhlmann 1953 (US). Rio
Erepecuri, Sampaio 4991 (R, US), 5567 (R, US). Marabé, Pires 4522 (US).
Between S. Jofio de Araguaya, 8S. Anna, Burchell 9264 (P). Maranuio: Tur-
yassi, Snethlage 272 (US). Fazenda do Rosario, Lisboa 55 (US).

Perhaps this species approaches most closely the section Visenia
of Asia and the Pacific Islands in habit, leaves, inflorescence, and
fruit. It is characterized by having hairs at the base of the petal
blade, by open, branched, fairly large axillary inflorescences, by
large leaves sparsely hairy on the upper surface, and by dehiscence of
the fruit part way along the dorsal suture and the entire ventral suture.
It is one of the largest species of the section Mougeotia.

Schumann (47) in monographing Melochia of Brazil considered
Melochia ulmifolia St.-Hil. & Naud. and Melochia ulmifolia Benth.,
based on different types, to have been published in 1842. He retained
the name for the former on the basis of page priority and renamed
Bentham’s species Melochia Benthami. However, as explained by
Sprague (50), Bentham published his species in 1841 and therefore
the name should be applied to his species. R. E. Fries (15) pointed
out that Bentham’s species had been described under the names
Riedlera ramuliflora Miq. in 1847 and Riedleja dichotoma Turcz. in
1858. He proposed for it the new combination Melochia ramuliflora
(Miq.) R. E. Fries. Since he admits that Bentham was the first to
describe the species, his action is unacceptable.

18. Melochia gardneri Sprague, Kew Bull. 124. 1915. PuaTE 1
Melochia ulmifolia St.-Hil. & Naud. Ann. Sci. Nat. II, 18:36. 1842; non
Benth. 1841.

A branched, erect, slender shrub 1.8-2.4 meters high, the young
stems and petioles tomentulose, hairs stellate, average 0.2 mm.
(0.1-0.7 mm.) long, internodes 1.3-3.4 cm. long; stipules 3.2-4.0
mm. long, 0.6-1.2 mm. wide, deltoid-acuminate, tomentulose and
ciliate, hairs stellate and simple, 0.2-0.6 mm. long; petiole 0.7-2.4
(—3.5) em. long.

Lamina rather thin, 5.7-8.8 (-12) em. long, 2.8-5.0 (—6.8) em. wide,
ovate to ovate-lanceolate, the base rounded, the apex acute to acumi-
nate, the upper surface green, sericeous, majority of hairs simple,
fewer shorter, forked and stellate, 0.2-1.3 mm. long, the lower surface
lighter green, tomentulose, soft to the touch, hairs stellate, a few
simple, 0.2-0.7 mm. long, the margin irregularly crenate-serrate,
serrations rather high, 0.8-3.6 mm. wide, 0.5-2.6 mm. high, 10-12
pairs of primary veins, 2 are basal.

Inflorescences terminal, sometimes also axillary, spicate, dense,
extending 4—5 cm. beyond the first leaf, the axillary ones 1 cm. long,
the pedicels 1-2 mm. long; bracts 3.5-5.7 mm. long, 0.5-0.7 mm. wide
lanceolate, sericeous on dorsal surface and ciliate, longer hairs simple,
shorter ones stellate, 0.2-0.7 mm. long.

258 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Calyx 7-8.3 mm. long, 4.3-4.4 mm. wide, not accrescent, hairs most
abundant on the junctures of the sepals, and ciliate, stellate, 0.1 mm.,
forked and simple, 0.5-1.2 mm. long, the teeth 3.5-4.5 mm. long,
2-2.2 mm. wide, deltoid-acuminate, the sinus acute; petals ?purple
(dried), 11.6-12 mm. long, 2.5-3.1 mm. wide, up to 4 mm. wide
according to Schumann (47), oblanceolate, the claw rather long, 3-4
mm. long, attached to the stamens for 2.7 mm.

Longistylous form. Stamens 4.9-5.9 mm. long, the filaments united
up to the anthers, the anther 1.3-1.4 mm. long, 0.6-0.7 mm. wide,
oblong, emarginate at both ends; pistil 11.1-12 mm. long, the styles
free for 4-4.7 mm., united for 3.5-4.0 mm., the ovary globose, pen-
tagonal-ovoid, the ovary and half of the free portion of the styles
stellate tomentulose.

Brevistylous form not seen.

Fruit 7 mm. long, gradually aristate for about 1 mm., 4.6 mm. wide,
globular-ovoid, obtusely pentagonal, pointed at the apex, shallowly
sulcate between the carpels, tomentulose, stellate hairs 0.1-0.2 mm.,
forked hairs 0.4-0.7 mm. long, stipe none; dehiscence all along the
ventral suture and a third along the dorsal suture, slightly septicidal;
seeds (not ripe) 2.5-2.8 mm. long, 1.5-1.8 mm. wide, brown, chalazal
membrane 0.2-0.3 mm. long, almost winglike, 2 per cell.

Flowering and fruiting specimens collected in April.

Type Locauity: Brazil, Goids, in woods on the Serra de Santa
Brida. Type in Herbarium Delessert, collected by Gardner, no.
3608, isotype cited below.

DistrisuTion: Brazil, Goids, in woods; also Piauf according to
Schumann (47).

BRAZIL: Gors: Serra de Santa Brida, Gardner 3608 (P, isotype, also US
neg. 5665 ex P, and F neg. of type at G).

This species is characterized by its dense, unbranched terminal
spike, rather large, thin, broad leaves, and comparatively large fruit
for the section Mougeotia, dehiscent part way along the dorsal suture
in addition to the entire ventral suture. It appears to be most close-
ly related to Melochia pilosa (Mill.) Fawe. & Rendle.

As explained by Sprague (50), Bentham’s Melochia ulmifolia was
published prior to that of St.-Hilaire and Naudin. Sprague renamed
the latter species Melochia Gardnert Sprague. In accordance with
recommended practice, I have substituted the small ‘“g’”’ for the
capital ““G” in Gardner.

19. Melochia mollis (H.B.K.) Triana & Planch. Ann. Sci. Nat. IV, 17:343. 1862.
Mougeotia mollis H.B.K. Nov. Gen. & Sp. 5:328. 1823.
Riedleia mollis (H.B.K.) DC. Prodr. 1:491. 1824.
Visenia mollis (H.B.K.) Spreng. Syst. 3:30. 1826.
Riedleta multiflora Presl, Rel. Haenk. 2:147. 1835.

GOLDBERG—GENUS MELOCHIA 259

Anamorpha melochioides Triana & Karst. in Triana, Nuev. Jen. i Esp. FI.
Neo-Granad. 10. 1854.
Melochia globifera Triana & Planch. Ann. Sci, Nat. IV, 17:342. 1862.

Usually a shrub, sometimes herb, erect, 0.5-5.0 meters high, the
young stems and petioles densely tomentulose-villous, hairs brownish,
stellate and usually also forked and simple, 0.1-2.0 mm. long, woolly
to the touch, internodes 1.1-8.0 cm. long; stipules 3-8.5 (-10.0)
mm. long, 0.7-1.8 (-3.8) mm. wide, deltoid-acuminate or deltoid-
acute, lanceolate or ovate, tomentulose and sometimes ciliate, hairs
stellate (short), forked and sometimes also simple, 0.2-1.5 mm.
long; petiole 0.5-5.4 (-8.0) em. long.

Lamina 3.7-13.5 (-22) em. long, 2-8.8 (-10.5) em. wide, broadly
ovate to lanceolate-ovate, the base rounded to cordate, the apex acute
to slightly rounded or slightly acuminate, the upper surface usually
greenish, sometimes brownish due to the hairs, tomentulose, sericeous
or with scattered hairs, velvety to the touch, hairs stellate (short)
and sometimes also forked and simple, 0.2-1.5 mm. long, sometimes
appressed, the lower surface tomentulose, velvety or woolly to the
touch, hairs stellate, sometimes a few forked and simple, 0.2-1.2
mm. long, lighter green, or yellowish-tan canescent, the margin
irregularly crenate-serrate, serrations 0.6-5.5 mm. wide, 0.4-3.0
mm. high, 7-11 (-15) pairs of primary veins, 2 (-3) are basal.

Inflorescences 2-10.5 cm. long, axillary in the upper 2-11 or more
axils, rarely also terminal, in subcapitate umbels of 2-5 flowers, 4-6
umbels in an inflorescence, sometimes corymbiform or paniculate, the
peduncle relatively thick, (0.2—-) 0.5-8.0 cm. long, 0.5-1.5 mm. wide,
sometimes with a primary branch 0.8-3.5 cm. long, and rarely a
secondary one 1-1.5 cm. long, sometimes naked branches with pe-
duncles at 4-5 nodes, the pedicels 1-4 (-8) mm. long; bracts 1-6.2 mm.
long, 0.2—2.2 mm. wide, deltoid-acuminate to linear, tomentulose and
sometimes ciliate, hairs stellate, forked, and simple, 0.1-1.4 mm. long.

Calyx 4-6.8 mm. long, 3.3-6.0 (-6.5) mm. wide, the teeth 1.7-4.6
mm. long, 1.2-2.7 mm. wide, deltoid to subovate, acute to acuminate,
the sinus acute, sometimes accrescent, becoming 7-9.5 mm. long,
7.7-10.0 mm. wide, the teeth 4.5-5.5 mm. long, 3.5-5.5 mm. wide,
densely tomentulose, canescent, hairs stellate (short) and some-
times also forked and simple, 0.2-1.0 mm. long; petals white to pale
yellow or white distally and yellow proximally, 4.2-6.8 mm. long,
1.4-2.7 mm. wide, obovate, obovate-oblong, obovate-lanceolate,
or oval, the claw 0.4-1.0 mm. long, free from the staminal tube or
attached to it for 0.5 mm., sometimes 2 or 3 simple and forked hairs
0.1 mm. long just above the claw.

Longistylous form. Stamens 3.2-4.2 mm. long, the filaments
united up to the anthers, sometimes tufts of hairs 0.4-0.5 mm. long

260 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

at the base of the staminal tube, alternate with the filaments, hairs
0.1 mm. long, the anther 1.0-1.3 mm. long, 0.5-1.0 mm. wide, oblong,
emarginate at both ends, nearly to the middle from the apex, sacs
slightly divergent at the apex; pistil 4.1-6.3 mm. long, the styles
2.6-4.0 mm. long, free to the ovary, filiform, slightly clavate and
papillose at the apex for 0.7-1.0 mm., the styles and ovary densely
tomentulose, hairs stellate and sometimes also simple, 0.2-0.3 mm.
long, the ovary globose.

Brevistylous form. Stamens 4.3-5.4 mm. long, the filaments free
for 1.5-2.5 mm., the anther 0.8-1.3 mm. long, 0.4-0.7 mm. wide,
shape same as that above, staminode teeth 0.1-+).2 mm. long some-
times present; pistil 2.8-3.8 mm. long, the styles 1.2-2.2 mm. long,
free to the ovary, papillose for 0.6 mm., base of the styles and the
ovary densely tomentulose, hairs stellate, 0.1-0.3 mm. long.

Fruit 2.7-4.0 mm. long, 3.3-4.8 mm. wide, globose, obtusely
pentagonal, slightly carinate along the midrib of the upper half of
the carpels, the sulci between the carpels extending a fifth to a third
the way to the center of the fruit, hirsute-tomentulose, hairs stellate,
forked and simple, 0.1-0.3 (-0.5) mm. long, the stipe 0-0.6 mm. long;
dehiscence septicidal into cocci open along the ventral suture, some-
times also a third to half way along the dorsal suture; seeds 1.7—2.4 mm.
long, 0.9-1.5 mm. wide, trigonal, brown, 1 per cell.

Flowering specimens collected in January, March, May to October,
and December; fruiting ones, in March, June to October, and in
December.

Type tocauity: Honda and Santanna, Tolima, Colombia, at
234-720 meters elevation, isotype cited below.

DistriIBUTION: Western Venezuela, Colombia, Ecuador, Peru, and
western Bolivia, on open, wooded hillsides, in moist copse, along road-
sides, desert stream bottom, dry grassy mountains, loam clearing,
in full sun or semishade, at 234—2,100 meters elevation.

VENEZUELA: Zutia: Perija, Gines 2148 (US).

COLOMBIA: Valle del Magdalena, Karsten s.n. (W). MaGpaLena: Santa
Marta, H. H. Smith 733 (A, K, L, US); Vista Nieve, Santa Marta, Viereck 9 (US).
NorTE DE SANTANDER: REGION DEL SARARE: Hoya del rio Chitagd, sobre La
Cabuya, Cuatrecasas et al. 12198 (US). Cunprnamarca: On the rio Bogota
in the vicinity of the bridge of San Antonio de Tena, Cuatrecasas 8280 (US).
“Guayabetal” to Monte Redondo SE. of Quetame, Pennell 1753 (US), 1794
(US). Villeta, Karsten s.n. (W). Road between Villeta and Guaduas, Ariste-
Joseph, July 1923 (US). El Colegio, Artste-Joseph 1046 (US). Sasaima, A.
Marie 21K (US). Toumma: Honda, Bonpland 1701 (P, isotype, presumably of
Mougeotia mollis H.B.K.), Andre s.n. (K). F neg. 9606 ex B, Humboldt &
Bonpland, presumably isotype of Mougeotia mollis H.B.K. (US). El Moral,
Quindio, Mariquita, Triana (P, type of Melochia globifera Tr. & Pl., also US neg.
5663). “La Trinidad,” Libano, Pennell 3249 (NY, US). Totima?: La Vibora,
Central Andes, Lehmann 4751 (US). Hutza: Cordillera Oriental, E. of Neiva,

GOLDBERG—GENUS MELOCHIA 261

Rusby & Pennell 445 (NY). ANtroquta: Bello, Bro. Thomas 1011 (US); Archer
184 (US). VauLLEe DEL Cauca: Rio Bolo, Dryander 752 (US). Rio Cali in the
vicinity of Cali, Hwan 15853 (US). Quebrada Rio Yumbo, above Yumbo,
Ewan 15839 (US). Espinal, below Dagua, Killip & Hazen 11063 (US). Nanrifto:
Guaitara, Karstens.n. (W). Pasto, Karsten s.n. (W).

ECUADOR: Eastern Cordillera, Rimbach 271 (US). Tuncauranvua: Bafios,
Tate 623 (US).

PERU: In valleys of the Andes, Haenkes.n. (PR 174402 a, b, type of Riedleta
multiflora Pres], also US neg. 5552). Pillao circa Chacahuassi, F. neg. 9603 ex B
of Ruiz, collected in 1787 det.? HuAnuco: Cuchero, Poeppig 1269 in part
(L, P, W). Near Cuchero, ?Poeppig 1379, September 1829 (W). Muna,
Macbride 4005 (US) det.? Junfn: Huacapistana, Sandeman 4511 (K, also US
neg. 5418), 4511A (K). San Ramén, Killip & Smith 24786 (W). AyYacucHo:
Cearrapa, between Huanta and Rio Apurimac, Killip & Smith 22323 (US).
Cusco: coNVENCION: Hda. Sahuayaco, Vargas C 1685 (GH).

BOLIVIA: South Yungas, Sirupaya at Yanacachi, Buchtien 511 (US). Near
Yungas, Rusby 1848 (NY, US).

This species is similar to M. lupulina Sw., but it is more robust, has
longer and thicker peduncles and hairier stems and leaves, and the

hairs are often brownish.
20. Melochia colombiana Cuatr. Rev. Acad. Colomb. Cienc. 8(32):486. 1952.

A branched, erect suffrutex at least 0.4 meter high, the young
stems and petioles densely tomentose, hairs stellate 0.5-1.3 mm. long,
light brown, internodes 0.4-0.7 cm. long; stipules 3.5-5.2 mm. long,
0.6-1.0 mm. wide, lanceolate, densely tomentose, hairs 0.3-1.0 mm.
long; petiole 0.2-0.5 cm. long.

Lamina 1.1-3.8 cm. long, 0.7-2.0 cm. wide, ovate to nearly oval,
the base rounded, the apex acute to slightly rounded, both surfaces
densely tomentose, lanate, woolly to the touch, hairs tan, stellate
and simple, 0.3-1.0 mm. long, the margin irregularly crenate-serrate,
serrations 1-2.7 mm. wide, 0.5-2.0 mm. high, 8 or 9 pairs of primary
veins, 2 are basal.

Inflorescences axillary, in upper 2-4 axils, 2.1-3.1 cm. long, the
peduncle 1.2-1.6 cm. long, with branch 0.8 cm. long, the ultimate
divisions umbellate, 5-10 flowers in a cluster, the pedicels 1.5-3.0
mm. long; bracts 1.5-3.0 mm. long, 0.2 mm. wide, lanceolate to
nearly linear, densely tomentose, hairs 0.3-1.0 mm. long.

Calyx 5-5.7 mm. long, 4.8-5.3 mm. wide, not accrescent, densely
tomentose on the outside, hairs stellate, 0.3-0.7 mm. long, the teeth
2.5 mm. long, 1.5-2.2 mm. wide, deltoid-acute, the sinus acute;
petals yellow, 5 mm. long, 1.1-1.3 mm. wide, oblanceolate, the claw
0.5-0.6 mm. long, apparently free from the staminal tube.

Longistylous form not seen.

Brevistylous form. Stamens 4.5-4.9 mm. long, the filaments free
for 1.5 mm., the anther 1.1-1.2 mm. long, 0.5-0.6 mm. wide, oblong,
emarginate at both ends, nearly to the middle from the apex; pistil

262 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

3-3.5 mm. long, the styles filiform, stigmatic for 0.5 mm., 1.5 mm.
long, free to the ovary, the ovary globose, densely tomentulose, hairs
0.3-0.5 mm. long.

Fruit 4 mm. long, 4.5 mm. wide, globose, pentagonal, the distal
half of carpels somewhat carinate along the midrib, sometimes a
slight hump about the middle of the dorsal surface of the carpels, the
sulci extending a third to a fourth the way to the center of the fruit,
densely tomentulose, hairs stellate, 0.3-0.6 mm. long, the stipe 0-0.3
mm. long; dehiscence septicidal into cocci open along the ventral
suture, and sometimes a third along the dorsal suture; seeds 2.2-2.3
mm. long, 1.3 mm. wide, brown, 1 per cell.

Flowering and fruiting specimens collected in September.

Tyrer Locatity: Colombia, Dept. del Valle, western slope of the
Cordillera Occidental. Lobo Guerrero in xerophytic habitats, at
610-650 meters elevation. The type was collected by Cuatrecasas,
no. 17804, and is cited below.

Distripurion: The species is known only from the type locality.

COLOMBIA: VaLLe pEL Cauca: CORDILLERA OCCIDENTAL: Lobo Guerrero,
Cuatrecasas 17804 (F, type, also US neg. 5670).

This species is close to Melochia mollis (H.B.K.) Triana & Planch.,
but it has smaller laminae, shorter petioles, and shorter internodes.

21. Melochia leucantha Macb. Field Mus. Nat. Hist. Bot. Ser. vol. 13, pt. 3A
No, 2:625. 1956. PLaTE 2

A branched, procumbent shrub 0.4-0.8 meter high, the young
stems and petioles cinereous, tomentulose, hairs stellate, 0.1-0.3 mm.
long, internodes 1-2.8 em. long; stipules 2.8-6.0 mm. long, 0.7-1.5
mm. wide, deltoid-acuminate, tomentulose, hairs stellate, sometimes
a few simple cilia, 0.1-0.7 mm. long; petiole 0.3-0.9 cm. long.

Lamina 1.7-4.0 em. long, 1.0-3.2 em. wide, broadly ovate, the
base rounded to subcordate, the apex acute to rounded, velvety to
the touch, the upper surface greenish-canescent-cinerous, tomentulose,
hairs stellate, 0.1-0.3 (-0.4) mm. long, the lower surface canescent-
cinereous, tomentulose, hairs stellate, 0.1-0.3 mm. long, the margin
irregularly crenate-serrate, serrations 1.0-3.6 mm. wide, 0.3-1.8 mm.
high, 5-9 pairs of primary veins, 2 or 3 are basal.

Inflorescences corymbiform to paniculate, the ultimate divisions
umbellate, 3-10 flowers per umbel, in the axils of the upper 1-3
leaves, 2.5-6.5 cm. long, and terminal, extending 2.2-4.4 em. beyond
the first leaf, the peduncle 0.72.8 cm. long, the pedicels 1.5-3.0 mm.
long; bracts 1-4.5 mm. long, 0.2-1.0 mm. wide, lanceolate to linear,
tomentulose and sometimes ciliate, hairs stellate, 0.1-0.5 mm. long.

Calyx 5.9-6.3 mm. long, to 6.6 mm. long in the fruiting stage,
4,5-5.2 mm. wide, cinereous, densely tomentulose, hairs stellate, a few

GOLDBERG—GENUS MELOCHIA 263

forked, 0.1-0.5 mm. long, the teeth 3.1-4.0 mm. long, 1.5-2.3 mm.
wide, deltoid-acuminate, the sinus acute; petals white, 6.8-7.2 mm.
long, 2.0-2.8 mm. wide, obovate to nearly elliptic, cuneate, the claw
0.3-1.0 mm. long, free from the staminal tube or attached to it for
0.5 mm.

Longistylous form. Stamens 4.5-4.9 mm. long, the filaments
united up to the anthers, the anther 1.4-1.6 mm. long, 0.8-0.9 mm.
wide, oblong, emarginate at both ends, nearly to the middle from the
apex; pistil 5.1-5.6 mm. long, the styles filiform, clavate and stig-
matic at the apex for 0.7 mm., 3.1-3.4 mm. long, free to the ovary,
the base of styles and ovary tomentulose, hairs stellate, few glandular,
0.1-0.2 mm. long, the ovary globose, pentagonal.

Brevistylous form. Stamens 5.5-6.0 mm. long, the filaments free
for 2-2.3 mm., the base of staminal tube tomentulose for 1.3 mm. in
tufts alternate with the filaments, hairs stellate 0.1-0.2 mm. long,
the anther 1.2-1.4 mm. long, 0.5-0.6 mm. wide, oblong, slightly
emarginate at both ends; pistil 3.7-4.2 mm. long, the styles filiform,
papillose for 0.5-0.7 mm. at the apex, 2.1-2.5 mm. long, free to the
ovary, the base of styles and ovary tomentulose, hairs stellate,
0.1-0.2 mm. long.

Fruit 3.8 mm. long, 3.9 mm. wide, globose, pentagonal, the angles
somewhat acute, the sulci between the carpels extending a third of
way to the center of the fruit, tomentulose, hairs mostly stellate, a
few forked and simple, 0.1-0.8 mm. long, the stipe 0.5 mm. long;
dehiscence septicidal into cocci open along the ventral suture; seeds
not quite ripe, 2.4 mm. long, 1.5 mm. wide, trigonal, brown, 1 per cell.

Flowering and fruiting specimens collected in March.

TypE Locauity: Eight kilometers east of Mejorada, Huancavelica,
Peru, in gravelly shrublands at 2,400 meters elevation. The type is
Goodspeed Expedition No. 10908, collected by Stork & Horton,
cited below.

DisrrisuTioN: Common, though not abundant, in shrubland from
Cusco, Peru, westward, at 2,000-2,400 meters elevation.

PERU: HvancavELica: HUANCAVELICA: Eight km. E. of Mejorada, Stork &
Horton 10908 (UC, type, also US neg. 5616). Tayscasa: Valley of the Mantaro
River below Colcabamba, Weberbawer 6438 (GH, US). Ayacucno: Soukup
4014 (US). Macbride (30) also reports the species from Rfo Pachuchaca,
Apurimac, Peru.

This species is close to M. mollis (H.B.K.) Triana & Planch., but
differs by having a loose corymbiform inflorescence, rather than
subcapitate clusters; the leaves of A. mollis are covered with longer,
often simple and forked as well as stellate hairs, that are brownish
rather than cinereous-canescent.

264 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

22. Melochia lupulina Sw. Prodr. Veg. Ind. Occ. 97. 1788.
Mougeotia inflata H.B.K. Nov. Gen. & Sp. 5:330, pl. 484. 1823.
Riedleia inflata (H.B.K.) DC. Prodr. 1:491. 1824.
Visenia inflata (H.B.K.) Spreng. Syst. 3:30. 1826.
Visenia lupulina (Sw.) Spreng. Syst. 3:31. 1826.
Riedleia melissaefolia Presl, Rel. Haenk. 2:149. 1835.
Melochia inflata (H.B.K.) Benth. Bot. Voy. Sulph. 71. 1844.
Riedleia lupulina (Sw.) Beurl, Vet. Akad. Handl. Stockh. 115. 1854 (1856).
Anamorpha waltherioides Triana & Karst. in Triana, Neuv. Jen. i Esp. Fl.
Neo-Granad. 10. 1854.
Physocodon macrobotrys Turez. Bull. Nat. Mosc. 31(1):213. 1858.

An erect herb to trailing suffrutex, often with several slender stems
from the base, 0.45-3.6 meters long, the young stems hirsute-tomen-
tose, hairs stellate, forked, simple, and sometimes glandular, 0.2-2 mm.
long, internodes 2.5-7.0 em. long; stipules 3.4-6.0 mm. long, 0.3-1.3
mm. wide, deltoid-acuminate, mostly ciliate, hairs 0.1-1.0 mm. long;
petiole 0.7-6.7 cm. long, tomentose, hairs 1-2 mm. long.

Lamina 3-11.8 cm. long, 1.5-8.5 em. wide, ovate, the base rounded
to cordate, the apex acute to subacuminate, the upper surface green,
hairs mostly simple, appressed, not much overlapping, sometimes a
few stellate and forked, 0.2-1.5 mm. long, more numerous on the
lower surface, which is subtomentose, hairs simple, forked, stellate,
and sometimes glandular, 0.3-1.5 mm. long, the margin irregularly
crenate-serrate, serrations 1-3 mm. wide, 0.6-2.0 mm. high, 7-11
pairs of primary veins, 2 are basal.

Inflorescences axillary, umbellate, clusters at upper 4-16 nodes,
usually 2 or 3 clusters per axil, 2—9 flowers in a cluster, the peduncle
short and slender, 0.1—-0.7 cm. long, 0.3-0.5 mm. wide, the pedicel
1-5 mm. long; bracts 1.5-4.0 mm. long, 0.1-0.7 mm. wide, deltoid-
acuminate to sublinear, mostly ciliate, hairs 0.1-1.5 mm. long.

Calyx membranous, greenish white, at anthesis 5.0-5.6 mm. long,
4.5-5.5 mm. wide, accrescent, becoming 8-12.3 mm. long, 12 mm. wide,
densely tomentulose, hairs stellate, forked, and sometimes glandular,
0.2-0.5 mm. long, the teeth at anthesis 2.5-3.2 mm. long, 1.7—-2.3 mm.
wide, becoming 4.3-6.5 mm. long, 3.6-4.6 mm. wide, deltoid-acute
to deltoid-acuminate, the sinus acute; petals usually white, sometimes
yellowish white or white distally and yellow proximally, 4.5-7.2 mm.
long, 1.6-2.1 mm. wide, obovate or oblanceolate, the blade rather
sharply demarcated from the claw, claw 0.5-0.7 mm. long, apparently
not attached to the staminal tube.

Longistylous form. Stamens 3.3-4.0 mm. long, the filaments
united up to the anthers, the anther 1.0-1.2 mm. long, 0.6 mm. wide,
emarginate at both ends, nearly to the middle from the apex, sacs
divergent at the apex; pistil 4.5-5.8 mm. long, the styles 3.2-4.2 mm.
long, free to the ovary, tomentose at the base, papillose at the apex

GOLDBERG—GENUS MELOCHIA 265

for 0.9 mm., the ovary subglobose, shallowly pentagonal, tomentulose,
hairs 0.1 mm. long, the stipe 0.5 mm. long.

Brevistylous form. Stamens 4.2 mm. long, the filaments free for 2
mm., the anther 0.9-1.2 mm. long, 0.5-0.7 mm. wide; pistil occasionally
of 6 carpels, 2.5 mm. long, the styles 1.5 mm. long, free to the ovary,
papillose for 0.5 mm.

Fruit 2.5-3.2 mm. long, 2.9-4.3 mm. wide, globose, pentagonal, the
sulci shallow, the cocci compressed, carinate, tomentulose, sometimes
also glandular, hairs 0.1 mm. long, the stipe 0.5-0.7 mm. long; dehis-
cence septicidal into cocci, open along the ventral suture; seeds 1.7—2.2
mm. long, 1.1-1.3 mm. wide, trigonal, the sides flat, the dorsum
rounded, brown.

Flowering and fruiting specimens collected throughout the year, but
flowering most commonly from October to February.

TYPE LOCALITY: The type was collected by Swartz in Jamaica and
is cited below.

Distripution: Mexico, Central America, Jamaica, Venezuela, Co-
lombia, Ecuador, and Peru, usually in moist thickets on shaded river
banks, also in llanos, campo, savannas, and weedy fields, in loam or
clay soil, at 0-3,150 meters elevation.

MEXICO: “In the land of western Mexico,’”’ Haenke (PR 174401 a, b, type of
Riedleia melissaefolia Presl, also US neg. 5551). Curapas: Vol. Tacana, Union,
Matuda 2788 (MICH).

GUATEMALA: Atta VerapPas: von Tiirckheim II 2201 (US). Finca Mocca,
H. Johnson 86 (US). Chicacoa, J. D. Smith 1764 (US). Near the Finca
Sepacuite, Cook & Griggs 292 (US). Between Cahabon and Secanquim, Goll 129
(US).

BRITISH HONDURAS: El Cayo District, Bartlett 12888 (MICH, $), 11426
(MICH). El Cayo and vicinity, Chanek 206 (MICH). Vaca, Gentle 2267
(MICH).

HONDURAS: Yoro: Quebrada Seca, Standley 53896 (US). Et Parafso:
Quebrada del Ingenio de los Angeles, Standley 29417 (US).

COSTA RICA: ?Salamanca, Tonduz 8613 (US). Between Barba and Santa
Barbara, Pittier 1684 (US). Santiago, Stork 1246 (MICH). ?ALAJUELA:
Alajuelita, Tonduz 1479 (US). San José: Between Aserri and Tarbaca, Standley
34054 (US), 41343 (US). Las Vueltas, Tucurrique, Tonduz 12871 (GH, US, W).
Vicinity of San José, Tonduz 10901 (US). Vicinity of El General, Skutch 2269
(S, US). Carraco: Vicinity of Pejivalle, Standley & Valerio 46802 (US). Rio
Turrialba, J. D. Smith 6456 (US).

PANAMA: Porto-Bello, Billberg 292 (S). Andersson s.n. (8S). Almerante,
Skutch (US). Changuinola Valley along R. R., Dunlap 334 (US). Nature
trail at Camp No. 1 on Madden Dam Highway, Elmore 58 (MICH). Bocas DEL
Toro: Carleton 183 (US). Water Valley vicinity of Chiriqui Lagoon, von Wedel
1671 (US), 1848 (US). Canan Zone: Gamboa, Standley 28329 (US), 28501
(US). Vicinity of Summit, Standley 25817 (US), 26932 (US), 29630 (US),
30059 (US). Hills W. of the Canal, near Gatdn, Standley 27273 (S, US). Gatun,
Bro. Heriberto 22 (US). Frijoles, Standley 27640 (US). Darien Station, Stand-
ley 31524 (US). Obispo, Standley 31772 (US). Empire to Mandinga, Piper

266 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

5460 (US). Pedro Miguel, Piper 5498 (US). Rio Grande, near Culebra,
Pittter 2109 (US). Vicinity of Monkey Hill, Cowell 39 (US). Panam: N. of
Panama City, Bro. Paul 6138 (MICH). Rio Tapia, Standley 28053 (US). Be-
tween Matias Hernindez and Juan Diaz, Standley 32063 (US). Matfas
Hernindez, Pittier 6900 (US). Darisn: Along Sambi River, Pittier 5537 (US).

JAMAICA: Swartz s.n. (8, type, also US neg. 5615). Kempshot, Harris
10828 (US). Near Holland Bay, St. Thomas, Harris 11955 (S, US). Phillips-
field near Bowden, Harrts & Britton 10783 (US). Along trail from Bath to
Cuna Cuna Pass, Mazon 1691 (US). Sulphur River, near the Fountain, 1 mi.
N. of Bath, Mrs. K. L. Harts.n. (lJ). Cedar Valley, Mrs. K, L. Harts.n. (IJ).
Between Whitehall and Big Hill, Proctor 7674 (IJ). Gorge of Plantain Garden
River NNW. of Whitehall, St. Thomas, Proctor 11783 (IJ); Stearn 485 (S).

VENEZUELA: Miranpa: Las Mostazas on R.R. beyond Los Teques, Allart
206 (US), 285 (US). (These two specimens lead toward M. mollis [H.B.K.]
Triana & Planch.)

COLOMBIA: Dawe 851 (US). Borfvar: Rio Sint, Cartagena, Humboldt &
Bonpland 1369 (P, isotype of Mougeotia inflata H.B.K., F. neg. 35380 ex P
[US], also US neg. 5660). Sincelejo, Pennell 4074 (US). Occidente de Monteria,
near Rio Sind, Cervantes, Molina & Barkley 19 Bo 053 (US). SANTANDER:
Vicinity of Barranca Bermeja, Magdalena Valley, Haught 1533 (US). Cunptr-
NAMARCA: Route de Villavicencio, Guyabetal, Humbert et al. 27146 (P, US).
Antioquia: Vicinity of Medellin, Barkley 18 C 663 (US). Caxpas: Salamina,
Bro. Daniel 2785 (US). VALLE DEL Cauca: Guanabanal, Killip 6222 (US).
“Potredillo” to ‘Miraflores,’ Palmira, Pennell & Killip 6063 (US). Cisneros,
Killip 5368 (US), 35521 (US). Cauca: Karsten s.n. (W, ?isotype of Anamorpha
walthertoides Triana & Karst.). Piedra de Moler et Cartago, Triana s.n. (P,
?type of Anamorpha waltherioides Triana & Karst., also US neg. 5662).

EQUADOR: Balao, Eggers 14066 (L, US). Terecita, Stevens 143 (US).
21km. SW. of Guayaquil, Worth et al. 8933 (US). Vicinity of Portovelo, J. N. &
G. Rose 23438 (US). 8 km. N. of Guayaquil, Stork et al. 8974 (US), Guayas:
Chongén, Asplund 5860 (R), 7683 (US). Rio Grande near Hacienda Tengucl,
Holmgren 30 (S, US). Guayaquil and vicinity Fagerlind & Wibom 105 bis (S);
Hitchcock 19934 (US); Hartweg 643 (W); Jameson 361 (LE, also US neg. 5270,
isotype of Physocodon macrobotrys Turcz.). Oil camp between Guayaquil and
Salinas, Hitchcock 20044 (US). Milagro, Hitchcock 20191 (US). Daule, Fager-
lind & Wibom 371 (S). Neara tributary of Rio Daule 1.5° S. lat., Boecher et al.
39 (8S). Los Rios: Near Quevedo, Canton Vinces, Mezia s.n. (US). Cuim-
BORAZO: Cafion of the Rio Chanchan near Huigra, Camp E-2943 (US). Cafiar:
Valley of Rio de Cafar near Rosario, Prieto CP-21 (NY). Ex Oro: Near
junction of Rio Luis and Rio Ambocas, 10 km. S. of Portovelo, Wiggins 10892
(US).

PERU: Haenke s.n. (PR). San Martin: Pucayacu 11 km. S. of Tarapoto,
Ferreyra 7759 (US). Loreto: Maynas: Near Yurimaguas, Poeppig 2425 (P,
W). Near Yurimaguas, at the River Huallaga, Spruce 3898 (W). LipEeRTap:
Trujillo, Killip & Smith 21501 (US). Chicama Valley, Smyth 80 (US). Lima:
Lima and vicinity, Savatier 1658 (K), 1659 (KK); Née s.n. (US); Mathews 1054
(W). Callao, insula 8. Lorenzo, Andersson, Mar. 18, 1852 (S, US). Santa Clara,
Mr. & Mrs. J. N. Rose 18735 (NY, US). Chosica, Soukup 1840 (US); Macbride
& Featherstone 496 (US, W); Asplund 10897 (US). Ricardo Palma, Reitz 6016
(US). cuancay: Hda. Ana Marfa, Ferreyra 7174 (US). HuAnuco: 15 mi.
Sk. of Hudnuco, Macbride & Featherstone 2060 (US). Junin: Colonia Perene,
Killip & Smith 25064 (US).

GOLDBERG—GENUS MELOCHIA 267

Melochia lupulina Sw. can be distinguished from most members of
the section Mougeotia by its markedly accrescent calyx. Melochia
mollis (H.B.K.) Triana & Planch. may also have a markedly accres-
cent calyx, but it is more robust and usually taller than M. lupulina,
0.5-5 versus 0.45-3.6 meters high; AZ. mollis has longer and thicker
peduncles, 0.5-8.0 cm. versus 0.1-0.7 cm. long, and it has hairier
stems and leaves.

23. Melochia thymifolia (Presl) A. Goldberg, comb. nov. PLATE 3
Riedleia thymifolia Presl, Rel. Haenk. 2:149. 1835.
Melochia nervosa (should be venosa) Sw. « typica Schum. var. purpurea O.
Ktze. Rev. Gen. 3(2):25. 1898.

Melochia yungasensis Rusby, Bull. New York Bot. Gard. 4:331. 1907.
Melochia pseudonodiflora Wochr. Ann. Cons. Jard. Geneve 21:430. 1920.

A branched erect or decumbent subshrub 0.2-0.9 meter high,
sometimes with numerous (to 10) slender stems from the base, the
taproot woody, thicker than the stem, 0.4-0.8 cm. wide, sometimes
with secondary branches, the young stems and petioles tomentulose,
canescent, hairs stellate, simple and forked, sometimes also glandular,
0.1-0.2 (-2.0) mm. long, internodes 0.5-5.0 cm. long; stipules 2.1-6.0
mm. long, 0.5-1.8 mm. wide, lanceolate or deltoid-acute or deltoid-
acuminate, ciliate, hirsute or tomentulose, hairs simple, forked,
stellate, and sometimes glandular, 0.1-0.2 (-1.5) mm. long, petiole
0.2-1.6 (-3.1) cm. long.

Lamina (0.9-) 1.5-4.5 (-7.5) em. long, (0.5-) 1.3-3.0 (4.5) cm.
wide, usually broadly to narrowly ovate, sometimes oblong-ovate,
the base usually rounded, sometimes subcordate or truncate, the
apex acute to rounded, the upper surface green, with slightly over-
lapping, simple, appressed hairs, or sericeous-tomentose, a few hairs
stellate and forked, sometimes also glandular, or only stellate and
forked, 0.1-0.2 (—1.5) mm. long, the lower surface lighter green,
with scattered hairs or tomentulose and slightly canescent, hairs
mostly stellate, some forked, usually also simple, sometimes also
glandular, 0.1-0.2 (—1.0) mm. long, velvety or slightly scabrous
to the touch, the margin irregularly crenate-serrate, serrations 0.7-3.7
(-6.9) mm. wide, 0.4-2.0 (-3.1) mm. high, (4—) 5-9 pairs of primary
veins, (1—-) 2 are basal.

Inflorescences on axillary branches 0.7-9.5 cm. long, leafless or
with reduced leaves, the flowers at 1-4 nodes, in the axils of the upper
1-6 (-9) leaves of the main stem and 1-3 leaves of the branches, also
terminal, extending 1-6 cm. beyond the first leaf, racemose, corymbi-
form or spicate, the ultimate divisions subcapitate umbels of (1-) 3-5
(-10) flowers, peduncles absent, especially from the inflorescences of
the branches, or 0.2-4.0 cm. long, slender 0.4—0.7 mm. wide, the

268 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

pedicel 0.5-3.0 (4.0) mm. long; bracts 1.0-5.5 mm. long, 0.2-1.1
mm. wide, linear, lanceolate, lanceolate-ovate, or deltoid-acute,
ciliate and hirsute or tomentulose, hairs simple, forked, and stellate,
0.1-0.6 (-1.5) mm. long.

Calyx sometimes reddish or purplish in the fruiting stage, (3.5-)
4.0-4.9 (-6.0) mm. long, 3.0-5.2 mm. wide, sometimes slightly
accrescent, becoming 4.5-6.8 mm. long and 4.8-5.7 mm. wide, the
external surface with scattered hairs or tomentulose, hairs simple,
forked, and stellate, sometimes predominantly glandular or only
stellate, minutely canescent at the apex of the internal surface, hairs
0.1-0.3 (-1.0) mm. long, the teeth (1.5-) 2.2-2.9 (4.0) mm. long,
1,2-2.0 (-2.9) mm. wide, deltoid- to ovate-acute or slightly ovate-
acuminate, the sinus usually acute, sometimes rounded; petals rose
or purple, sometimes white, 4.2-7.5 mm. long, 1.5-2.7 mm. wide,
obovate, elliptic-oblong, or oblanceolate, cuneate, free from the
stamens, the claw 0.3-1.2 mm. long.

Longistylous form. Stamens 2.4-3.5 mm. long, the filaments united
up to the anthers, the anther 0.8-1.1 mm. long, 0.5-0.8 mm. wide,
oblong, emarginate at both ends, nearly to the middle from the apex,
sacs divergent at the apex; pistil 2.6-5.8 mm. long, the styles 1.3-
3.4 mm. long, free to the ovary, papillose for 0.7-0.8 mm. at the
apex, the ovary globular-ovoid, pentagonal, densely sericeous, hairs
0.1-0.8 mm. long.

Brevistylous form. Stamens 4.1-5.1 mm. long, the filaments free
for 2.0-2.3 mm., the anther 0.8-1.1 mm. long, 0.5-0.6 mm. wide,
shape same as that above; pistil 2.3-2.6 mm. long, the styles 1.3-1.7
mm. long, free to the ovary, the ovary globular, tomentulose-seri-
ceous, or only hirsute at the apex, hairs 0.1-0.3 mm. long.

Fruit 2.6-4.5 mm. long, 3.0-4.5 mm. wide, globular, pentagonal,
the carpels narrowing toward the apex, middorsal surface of carpels
rounded at the base, keeled in the upper half, the sulci between the
carpels extending one-fourth (one-fifth at the middle and one-third
at the apex) of way to the center of the fruit, with scattered hairs or
tomentulose, hairs simple, forked, and stellate, sometimes also glandu-
lar, 0.1-0.5 mm. long, the stipe 0-0.4 mm. long; dehiscence septicidal
into cocci open along the ventral suture, and sometimes readily
separated for a fourth to half way along the dorsal suture; seeds
trigonal, 1.8-2.6 mm. long, 1.1-1.7 mm. wide, rarely the testa of the
chalazal end is loosened for 0.1-0.2 mm. and almost winglike, brown,
1 per cell.

Flowering specimens collected in February, April, June, August,
September, and November; fruiting ones, in June, September, and
November.

GOLDBERG—GENUS MELOCHIA 269

Type Locauity: The lectotype was collected by Haenke, PR
174403a, in Hudnuco, Peru, and is cited below.

DistrisuTion: In the Andes of Ecuador to Bolivia, on open hill-
side, on schistose rocks or red, sandy soil, usually at 2,400-2,700
meters elevation, but extending downward toward the Amazon basin.

ECUADOR: Cuimporazo: Vicinity of Huigra, mostly on the Hda. de Licay,
J.N. & G. Rose 22546 (GH).

PERU: “Peru et Chili,” Ruiz s.n. (G, type of Melochia pseudonodiflora Hochr.,
also US neg. 5416). Casamarca: Tambillo, de Jelski 282 (W, also US neg. 5546).
Amazonas: Chachapoyas, Mathews 2115 (K), 3233 (K). HuAnuco: F neg. 9602
ex B of Ruiz, collected 1787 (US). “Peruvian Andes at Huanocco,” Haenke
(PR 174403 a and b, lectotype and isotype, respectively, also US neg. 5553).
AMBO: Arriba de Quicacan, Ferreyra 6631 (US). Junfn: Carpapata, above
Huacapistana, Killip & Smith 24331 (US).

BOLIVIA: Yungas, Bang 1977 (NY, type; US, isotype of Melochia yungasensis
Rusby). La Paz: Bang 2132 (GH). Cotafia at Illimani, Buchtien 254 (GH,
NY), 3222 (NY, 8, US). Larecasa: Vicinity of Sorata, Mandon 819 (K, S, US).
CocHABAMBA: Tunari, Kuntze, April 1892 (NY).

This species is close to Melochia nodiflora Sw., but the young stems
and leaves are generally hairier; the number of nodes with inflores-
cences are fewer, 1-6 (—9) versus up to 20; a peduncle is often present,
it is usually absent in MM. nodiflora, and shorter when present; the
pedicels are generally slightly longer, the terminal inflorescence is
more frequent and longer, and the sulci between the carpels are
shallower than in M. nodiflora.

This species is also related to Melochia nudiflora Standl. & L. Wms.,
but the leaves have fewer primary veins, 5-9 versus 9-14. The
bracts are narrow, to 1.1 mm. wide, in M. nudiflora the majority are
broad, to 3.9 mm. wide. The petals are free of the staminal tube
but attached to it for 1-2 mm. in M. nudiflora.

This species is also close to Melochia lupulina Sw., but the calyx is
not accrescent or slightly so, becoming 4.5-6.8 mm. long and 4.8—5.7
mm. wide, in M. lupulina it becomes 8-12.3 mm. long and 12 mm.
wide. There is only one flower cluster per axil, frequently 2 or 3
in M. lupulina. The pedicels are generally shorter and the peduncle,
when present, is usually longer than in M. lupulina.

24. Melochia bernoulliana Donn. Smith, Bot. Gaz. 35:2. 1903.

A branched, slender shrub 1-4 meters high, the young stems and
petioles covered with stellate hairs less than 0.1 mm. long, internodes
2-6 cm. long; stipules deciduous, 3.5—-5.0 mm. long, 0.6-0.8 mm. wide,
lanceolate, with silky hairs up to 0.3 mm. long; petiole slender, 0.3-
3.6 cm. long.

Lamina thin, 3-11.5 (-13) em. long, 1.1-7.5 cm. wide, usually
ovate, sometimes also lanceolate, the base usually rounded, some-

227-051—67__6

270 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

times truncate or subcordate, the apex usually acuminate, sometimes
acute, the upper surface of young leaves with appressed, simple,
deciduous hairs 0.1-0.5 mm. long on the blade, and stellate hairs
less than 0.1 mm. long on the veins, the lower surface with stellate
hairs less than 0.1 mm. long on the veins, the margin irregularly
crenate-serrate, serrations high, 1-5 mm. wide, 0.5-3.5 mm. high,
7-11 pairs of primary veins, 2 are basal.

Inflorescence slender, somewhat paniculate, the ultimate divisions
usually umbellate, 2-5 flowers per umbel, the terminal inflorescence
extending 3-5 cm. beyond the first leaf, the axillary inflorescences
2-13 cm. long, in upper 5-10 axils, slender axillary flowering branches
9-13 cm. long, the peduncle 0-2.0 cm. long, very slender, the pedicels
relatively long, slender, 3-7 mm. long; bracts 1-3.7 mm. long, 0.2-
1.5 mm, wide, ovate to nearly linear, with simple, forked, and stellate
hairs 0.1-0.4 mm. long.

Calyx 3.5-5.0 mm. long, 4-6.5 mm. wide, not accrescent, covered
with stellate, simple and glandular hairs up to 0.1 mm. long, the
teeth 2.1-3.8 mm. long, 1.4-2.2 mm. wide, deltoid- to ovate-acute to
ovate-acuminate, the sinus acute to narrowly rounded; petals pink
with white at the tips, or purple, 5-7.3 mm. long, 2.2-2.9 mm. wide,
obovate to oblanceolate, attached to staminal tube for 1 mm., the
claw 0-0.5 mm. long.

Longistylous form. Stamens 3.4 mm. long, the filaments united
up to the anthers, the anther 1.0-1.1 mm. long, 0.7 mm. wide, oblong,
emarginate at both ends, almost to the middle from the apex; pistil
4.8-5.0 mm. long, the styles 3.5-3.7 mm. long, free to the ovary,
papillose at the apex for 0.8-1.0 mm., the ovary ovoid, hirsute-
sericeous, hairs 0.5 mm. long, stipe none.

Brevistylous form. Stamens 2.8-4.0 mm, long, the filaments
free for 1.3-2.7 mm., filamentous staminodes 0.4 mm. long sometimes
present; pistil 2.7-3.1 mm. long, the styles 1.3-1.6 mm. long, free to
the ovary, the ovary ovoid, hirsute, hairs forked, 0.3-0.5 mm. long.

Fruit obtusely pentagonal, depressed-globose, 3.5-4.1 mm. long,
4.4-4.8 mm. wide, with forked and simple hairs 0.1-0.5 mm. long
and stellate hairs less than 0.1 mm. long, deeply sulcate between the
carpels, the sulci extending half way to the center of the fruit, stipe
none; dehiscence septicidal into cocci open along the ventral suture;
seeds 2.1-2.4 mm. long, 1.4—-1.6 mm. wide, red brown.

Flowering and fruiting specimens collected from October to
December (old fruit sometimes still present in March).

Type Locauity: Retalhuleu (Bernoulli & Cario 3112) and Santa
Rosa (Heyde & Lux 4319), Guatemala; lectotype and paratype
cited below.

GOLDBERG—GENUS MELOCHIA 271

DistrisutTion: Chiapas, Mexico, to San Vicente, Salvador, in
moist or dry thickets, sometimes on cliffs or along streams, at 350-800
meters elevation.

MEXICO: Cutapas: Escuintla, Matuda 485 (MICH, US), 2129 (MICH).
GUATEMALA: ReratuuLeu: Bernoulli & Cario 3112 (S, paratype). Santa
Rosa: Mouth of the Rio De Los Esclavos, Heyde & Lux 4319 (US, lectotype).
SALVADOR: Sonsonate: Near headwaters of Rio Acachapa, Hda. Las
Tablos, Balsam Range, Tucker 1324 (US). La Lipertap: Ateos, Calderén
1858 (GH, US). San Vicente: Vicinity of San Vicente, Standley 21676 (US).

This species is characterized by its slender, open, somewhat pan-
iculate inflorescences, long slender pedicels, and thin lamina with
very short pubescence and high serrations. It is perhaps most closely
related to Melochia lupulina Sw. but does not have an accrescent
calyx.

25. Melochia nodiflora Sw. Prodr. Veg. Ind. Oce. 97. 1788.
Melochia carpinifolia Wendl. Bot. Beobacht. 52. 1798.
Mougeotia nodiflora (Sw.) H.B.K. Nov. Gen. & Sp. 5:330. 1823.
Riedleia nodiflora (Sw.) DC. Prodr. 1:491. 1824.
Visenia nodiflora (Sw.) Spreng. Syst. 3:30. 1826.
Riedleja urticaefolia Turcz. Bull. Soc. Nat. Mose. 31(1):209. 1858.
Melochia conglobata Sessé & Moc. Pl. Nov. Hispan. ed. I, 106. 1889.
Melochia urticaefolia (Turcz.) Standl. Contr. U.S. Nat. Herb. 23:804. 1923.

An erect or spreading herb to slender, branched shrub 0.5-4.0
meters high, the taproot somewhat thicker than the stem, the young
stems puberulous, hairs simple and forked, 0.1-0.2 (-0.6) mm. long,
internodes 1.3-3.2 em. long; stipules soon deciduous, 1.7-5.2 mm.
long, 0.7-2.9 mm. wide, deltoid- to ovate-acute, hirsute-ciliate, hairs
scattered, appressed, 0.2-0.8 mm. long; petiole 0.6-4.3 cm. long,
hirsute-tomentose, hairs simple and forked, 0.4-1.0 mm. long.

Lamina 3.5-7.5 (2-11.9) em. long, 1.5-5.3 (1.0-7.7) cm. wide,
usually ovate, sometimes also ovate-lanceolate, the base rounded to
cordate, or subtruncate, the apex acute, the upper surface green,
subglabrate, a few scattered, simple, appressed hairs 0.1-0.7 mm.
long, the lower surface green, subglabrate or puberulous, a few scat-
tered simple, rarely also forked and stellate hairs, appressed or erect,
0.1-0.7 mm. long, the margin irregularly crenate-serrate, serrations
1.5-4.8 mm. wide, 0.7-2.0 mm. high, 7-10 pairs of primary veins,
usually 2 are basal, the second pair thicker than the first.

Inflorescences axillary, in heads of 2-10 flowers at the upper 15-20
nodes, peduncle usually none, rarely 0.4-0.6 cm. long, sometimes
naked axillary branches up to 7.5 cm. long with flowers at several
nodes, the pedicel 0-2 (-3) mm. long; bracts 1.7-3.0 mm. long, 0.2-
1.1 mm. wide, rarely wider and 2 united, deltoid-acute to deltoid-

272 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

acuminate or nearly linear, hirsute-ciliate, hairs simple, 0.3-0.7 mm.
long.

Calyx sometimes dull red in the fruiting stage, 3.2-7.0 mm. long,
3-3.8 mm. wide, not accrescent, hirsute-ciliate, hairs simple and
forked, 0.1-1.0 mm. long, the teeth 2-3 mm. long, 0.5-1.5 mm. wide,
deltoid-acute to deltoid-acuminate or ovate-acuminate, the sinus
acute; petals most commonly pink, also purplish, white with red or
pink veins, pale pink with purplish-red veins, white, rarely yellow,
3.7-4.9 (-7.0) mm. long, 1.4-2.1 mm. wide, obovate, oval or elliptic,
the claw 0-0.5 mm. long, base of petals shortly and loosely united to
the staminal tube.

Longistylous form. Stamens 2.7-3.0 mm. long, the filaments
united up to the anthers, the anther 0.7-0.9 mm. long, 0.4-0.6 mm.
wide, oblong, emarginate at both ends, nearly to the middle from the
apex, sacs slightly divergent at the apex; pistil 3.2-3.5 mm. long, the
styles 1.3-2.0 mm. long, free to the ovary, papillose at the apex for
0.5 mm., the ovary ovoid or orbicular, hirsute, hairs simple, to 0.5
mm. long.

Brevistylous form not seen.

Fruit rarely of 6 carpels, 3.0-3.5 mm. long, 3.9-4.5 mm. wide,
oblate-spheroid, pentagonal, the sulci between the carpels extending
half way to the center of the fruit, hirsute, hairs simple, 0.4-1.0 mm.
long, stipe none; dehiscence septicidal into cocci open along the ventral
suture; seeds 2—2.2 mm. long, 1.1-1.4 mm. wide, trigonal, brown or tan.

Flowering specimens collected from August to April; fruiting ones,
from July to May, both most commonly from November to February
in the Northern Hemisphere.

Tyrer LocALiry: The type was collected by Swartz in Jamaica and
is cited below.

DistrisutTion: Mexico to northern and eastern Brazil and the
West Indies, in moist thickets, along the roadside, in cultivated
ground, in waste ground, in sandy or open woods, in pastures, in wet
or dry places on rocky, calcareous or sandy soil, at 0-1,800 meters
elevation.

MEXICO: Sessé et al. (MA, type of Melochia conglobata Sessé & Moc.). F
neg. 30510 ex MA of drawing of M. congloblata Sessé & Moc. (F). SrNnauoa:
Imala, Gentry 4968 (GH). Jantsco: San Sebastian, Sierra Madre Mts., Mexia
1475 (A). Vera Cruz: Orcutt 3168 (K, US). Zacuapan, Barranca de Santa
Maria, Purpus 2358 (US). Rio de la Ternera, Purpus 8276 (US). Santa
Lucrecia, Mell 667 (US). Cameron, Purpus 8630 (UC). Mexico: TEMASCAL-
tererEc: Hinton 2072 (US). Chorrera, Hinton 5368 (US). Pefién, Hinton 2144
(US). Ixtapan, Hinton 2950 (MICH, 8). MicuoacaAn: zirAcuaro: Zitdcuaro-
La Florida, Hinton 13407 (MICH, US). W. of Jiquilpan along highway to
Colima, McVaugh 13207 (MICH, US). Gurrrero: Acapulco and vicinity, EF.

Palmer 261 in part (MICH, US). coyuca: Coyuca-Chacamerito, Hinton 5689
(US). mina: Manchon, Hinton 9640 (K, US). GaLeana: Plan de Carrizo,

GOLDBERG—GENUS MELOCHIA 273

Hinton 11024 (US). Oaxaca: Galeotti 4091 (W, isotype of Riedleja urticaefolia
Turez., also US neg. 5547). Vicinity of Cafetal Concordia, Morton & Makrinius
2706 (US), 2321 (US). Pochutla, Conzatti et al. 3021 (US), 3193 (US). Yucatan:
Izamal, Gaumer 948 (US). Chichankanab, Gawmer 2080 (MICH, US). Xnoéaé,
Gaumer & sons 23507 (S, US).

GUATEMALA: Hyde 631 (US). Gualan, Deam 267 (MICH). Santa Rosa:
Volcan Jumaytepeque, Heyde & Lux 4317 (US).

BRITISH HONDURAS: Prospecto, Northern River, Gentle 922 (MICH).
Little Cocquericot, Belize River, Lundell 4235 (MICH).

HONDURAS: Vicinity of Amapala, Isla Tigre, Standley 20696 (US).

SALVADOR: Cerro del Guayabal, Calderén 1958 (US). Zacatecoluca, Cal-
derén 329 (US). AnuacHapdn: Padilla 473 (US). Vicinity of Ahuachapdn,
Standley 20216 (US). San Vicents: Voledin de San Vicente, Iglesias 1 (US).
San Miavuex: Hacienda San Antonio, Calderén 2121 (US). Moraz4n: Eastern-
most peak, Montes de Cacaquatique, near summit on N. side, Tucker 630 (US).
La Union: Grant 720 (MICH).

NICARAGUA: C. Wright sn. (US). Granada, Baker 582 (US). Ometepe
Island, Lake Nicaragua, Shimek & C. L. Smith 110 (US). Manaaua: Chaves 31
(US). CuontTazes: Friedrichsthal 725 (W).

COSTA RICA: Around Nicoya, Tonduz 13489 (GH, US). Near Rfo Terraba,
vicinity of Palmar Norte de Osa, Allen 5451 (MICH). GuANAcaAsTE: Vicinity
of Libano, Standley & Valerio 44906 (US). Vicinity of Tilardn, Standley &
Valerio 45690 (US). Bebedero, Standley & Valerio 46685 (US).

PANAMA: Seemann 473 (8S). Pearl Islands, Johansen 116 (US). Isthmus
of Panama, Hayes 154 (GH). Cutriquf: Vicinity of San Carlos, Allen 1140
(US). Cana Zone: Balboa, Standley 25545 (US), 27126 (US), 29260 (US),
32112 (US). Near Summit, Standley 29649 (US). Vicinity of Miraflores Lake,
White 260 (US). Along the old Las Cruces Trail, between Fort Clayton and
Corozal, Standley 29149 (US). Panama: Vicinity of Panama, Macbride 2618
(US). Vicinity of Matias Herndndez, Standley 28870 (US); Pittier 6905 (US).
Between Matias Hernandez and Juan Dfaz, Standley 32092 (US). Between
Las Sabanas and Matfas Hernandez, Standley 31847 (US). Bella Vista, Stand-
ley 25307 (S, US). Near Chepo, Hunter & Allen 61 (S).

BAHAMAS: New Provipencs: Nassau, Curtiss 26 (L, US); Britton & Mills-
paugh 2087 (US).

CUBA: C. Wright 39 (US). de la Sagra 69 (W). Merats.n. (W). Yumuny,
Rugel 27 (L). Tumbadero, Otto 77 (W). Pinar pet Rfo: Sierra de Anafe,
Wilson 11490 (US). Hapana: Anafe near Lago de Ariguanabo, Ekman 1041
(S). Santiago de las Vegas, Wilson 3642 (UC, US). Las Vituas: Limones,
Soledad, Cienfuegos, Pringle 84 (R, US); Jack 4622 (US), 6649 <5), 6895 (S, US).
Camacuey: Vicinity of La Gloria, Shafer 46 (US). OnrienTE: Vicinity of Bar-
acoa, Pollard et al. 12 (US). Baracoa, at margin of Rio Toa, Ekman 4063 (S).
Santiago de Cuba, Ekman 8467 (S), 8988 (S). Bayate, Ekman 4534 (S).

HAITI: Port-au-Prince, Eggers 3312 (8); Holdridge 1027 (MICH, US);
Leonard 2801 (US). Vicinity of Port de Paix, ridge W. of La Coup River, E. C. &
G. M. Leonard 11128 (US). Massif de la Hotte, Fond-des-Négres, Ekman
H7200 (8S, US). Vicinity of Jean Rabel, E. C. & G. M. Leonard 12805 (US),
12632 (US). Dept. du Nord, vicinity of Plaisance, Leonard 9247 (US), 9286 (US).
Vicinity of St. Michel de l’Atalaye, Leonard 7077 (US), 7729 (US), 8019 (US).
Morne de l’H6pital, ?Ticanda 1202 (S). Vicinity of St. Louis du Nord, E. C. &
G. M. Leonard 14176 (US).

DOMINICAN REPUBLIC: C. Wright et al. 105 (US). von Tiirckheim 2579
(L, S, US). San Pedro de Macoris, Rose et al. 3713 (US). Azua, Rose et al.

274 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

4057 (US). Santo Dominao: Vicinity of Ciudad Trujillo, Allard 13378 (US),
13400 (S, US), 13688 (US), 13696 (US), 14622 (US). Santo Domingo City,
Ekman 10941 (8S, US). Baranona: Riocito, Fuertes 199 (US).

JAMAICA: Swartz s.n. (8, type, also US neg. 5608). Upper slopes of Mount
Diablo, Mazon & Killip 423 (US). Spanish Town, Clute 36 (US). Near Ewar-
ton, Harris 6615 (US). Above Kingston, Lehmann B. T. 896 (L). Granp
Cayman: Proctor 9565 (IJ).

PUERTO RICO: Krebs s.n. (8S). Sabana Grande, Sargent 470 (US). Rio
Piedras, Johnson & Stevenson 1133 (US). Juana Diaz, Sintenis 2894 (L, US).
Mayagiiez, Sintenis 729 (L, S, US). Guayama, Jobas Road, Goll 508 (US).
Mannabo-Guayama road, Dale s.n. (MICH). Bayamon, Stahl 850 (L, 8). Be-
tween Cabo Rojo and San German, Britton 4307 (US). Arecibo, Heller 342 (US).
Ponce, Goll 825 (US). Near Ponce on Adjuntas road, Heller 6059 (L, US).
Aguadilla, Heller 4527 (L, US). Yauco, Underwood & Griggs 651 (US). VIEQUES
Isuanp: Vicinity of Isabel Segunda, Shafer 2414 (US). Cunmpra IsLanp:
Britton & Wheeler 41 (US).

LESSER ANTILLES: U.S. Virain Istanps: st. crotx: Christiansted, Rose
et al. 3566 (US). Bassin, Ricksecker 84 (L, US). ST. THOMAS: Friedrichsthal 29
(W); Kuntze ? 546 (US); Eggers s.n. (8), 98 (US), 97 (MICH), 261 (L, US);
Oersted 30 (MICH, 8, US). st. soun: Morrow 160 (US). BrittsH VircIn
IsLANDS: ToRTOLA: Fishlock 15 (US). Sv. Barrufiemy: Forsstrém s.n. (S).
Anticua: Bor 1249 dupl. (US). Rose et al. 3490 (US). Montserrat: Shafer
646 (US). GuapxELoupeE: Forsstrém s.n. (5). Duss 2670 (US). Moule, Questel
807 (US), 808 (US). Morne Miquel, Questel 448 (US). Ste. Marguerite, Questel
4280 ? (US). Marrinraue: Sieber 395 (L, PR, W). Duss 1361 (US). Fort de
France, H. & M. Stehle 5734 (US).

TOBAGO: Scarborough, Broadway 2956 (L).

VENEZUELA: Miranpa: Santa Luefa along Rfo Guaine, Killip & Tamayo
37015 (US).

COLOMBIA: Maapaena: Santa Marta, H. H. Smith 465 (L, 8, US). AriAn-
rico: Barranquilla and vicinity, Bro. Elias 332 (US); Dugand 5430 (US). Los
Pendales at border of Dept. of Bolivar, Dugand & Jaramillo 4107 (US). Between
Palmar de Varela and Ponedera on the banks of the Magdalena River, Dugand
4702 (US). Usiacuri, Dugand & Barriga 2367 (US). Borfvar: Cartagena and
vicinity, Billberg 35 (S), 202 (S); Dahlin s.n. (8); Bro. Heriberto 330 (US), 273
(US). Ternera, near Cartagena, Bro. Angel 772 (US). Vicinity of Turbaco,
Bro. Heriberto 427 (US). Buenavista, FE. of Since, Pennell 3986 (US).

BRAZIL: Creark: Alleméo & de Cysneiros 132 (R, US). Maranguape, Drouet
2259 (MICH, US). Serra de Maranguape, Ule 9067 (L). Maracanat, Ducke
2607 (US). Rro pe Janerro: Cabo Frio, Restinga, Glaziou 10327 (P).

This species is characterized by its dense subsessile clusters of
flowers in the axils of numerous (15-20) leaves, and by its subglabrous
leaves.

26. Melochia nudiflora Standl. & L. Wms. Ceiba 3:122. 1952,

A slender, branched, erect or subscandent shrub 0.9-4.0 meters high,
the young stems and petioles tomentose, hairs yellowish brown,
stellate, mostly 0.1-0.2 mm. long, forked and simple, 0.5-1.2 mm. long,
internodes 1.7-9.0 cm. long; stipules soon deciduous, probably broad
judging from the scar; petiole 0.2-3.0 cm. long.

GOLDBERG—GENUS MELOCHIA 275

Lamina 3-11 cm. long, 1.0-6.7 cm. wide, ovate to lanceolate, the
base rounded to cordate, the apex acuminate to acute, the upper
surface subsericeous, covered with appressed simple, and a few forked
and stellate, nonoverlapping hairs 0.2-1.3 mm. long, the lower surface
subcanescent, tomentulose-sericeous, hairs stellate, simple, and forked,
0.1-1.0 mm. long, both surfaces velvety to the touch, the margin
irregularly crenate-serrate, serrations 0.8-3.2 mm. wide, 0.5-2.6 mm.
high, 9-14 pairs of primary veins, 2 are basal.

Inflorescences axillary and terminal in subcapitate clusters of 2-15
flowers, the terminal inflorescence extending 1-14.5 cm. beyond the
first leaf, with flowers at 1-6 nodes, the axillary inflorescences in the
axils of 3-9 leaves, sessile, 1 cm. long, or on branches to about 8 cm.
long, the pedicel 0-2 mm. long; bracts 2-6 mm. long, 0.6-3.9 mm. wide,
the majority rather broad, suborbicular, ovate to lanceolate, the apex
acute to rounded, several at the base of each flower, apparently 2 or 3
sometimes united nearly to the apex, hirsute and ciliate, hairs forked,
simple, and stellate, 0.1-1.3 mm. long.

Calyx 4.5-6.4 mm. long, 3.5-5.0 mm. wide, not accrescent, tomen-
tulose-sericeous on the external surface, hairs simple, forked, and
stellate, 0.1-1.3 mm. long, tomentulose and canescent at the apex of
the internal surface, the teeth 2.4-4.6 mm. long, 1.3-1.9 mm. wide,
deltoid- to ovate-acuminate, the sinus acute to narrowly rounded;
petals pink, 5.5-7.0 mm. long, 1.5-3.5 mm. wide, obovate-cuneate,
attached to the staminal tube for 1-2 mm., the claw 0.7-2.0 mm. long.

Longistylous form. Stamens 3.5-4.0 mm. long, the filaments
united up to the anthers, the anther 1.1-1.2 mm. long, 0.7-0.8 mm.
wide, oblong, emarginate at both ends, nearly to the middle from the
apex; pistil 5.1-5.7 mm. long, the styles 3.2-4.0 mm. long, free to the
ovary, papillose at the apex for 0.7-1.0 mm., the ovary densely seri-
ceous, hairs mostly simple, a few forked, 0.7-1.5 mm. long.

Brevistylous form. Stamens 5-5.5 mm. long, the filaments free
for 1.7-3.0 mm., the anther 1.1-1.4 mm. long, 0.6—-0.9 mm. wide, shape
same as that above; pistil 3.0-3.5 mm. long, the styles 1.5-2.2 mm.
long, free to the ovary, the ovary globular-ovoid, densely sericeous,
hairs simple, 0.5-1.0 mm. long.

Fruit 3.2-4.4 mm. long, 4.5-5.1 mm. wide, obtusely pentagonal,
oblate-spheroid, the sulci between the carpels extending a fourth to a
third of way to the center of the fruit, hirsute- to sericeous-tomentose,
hairs stellate (short), forked, and simple, 0.1-1.0 mm. long, stipe none;
dehiscence septicidal into cocci open along the ventral suture; seeds
2.32.6 mm. long, 1.6-2.0 mm. wide, trigonal, brown, 1 per cell.

Flowering specimens collected in September, October, December,
January, and February; fruiting ones, in October, December, and
February.

276 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Typr Locauiry: The type, cited below, was collected by Williams
and Molina, no. 11234, at Quebrada de Suyapa, Morazan, Honduras,
at 1,200 meters elevation.

Disrripution: Mexico to Costa Rica, in thickets on rocky soil, on
limestone hills, in dry forest, or in fields, at 750-1,700 meters elevation.

MEXICO: Morenos: Cuernavaca, Pringle 7342 (MICH). MIcHOACAN:
ARTEAGA: Fresnal, Hinton 15232 (US). GuErRERo: Near Iguala, Pringle 13509
(L, MICH, 8). Mountains above Iguala, Pringle 8404 (US, W). Iguala Canyon,
Pringle 10335 (R, 8, W). Los Ametes Station, Pringle 13818 (MICH, US).
Curapas: Tuxtla Gutierrez, Purpus 9226 (GH, UC, US). Tuxtla, Hac. Petapa,
C. & E. Seler 1953 (US).

GUATEMALA: Jacaltenanjo, Nelson 3587 (US). GuaremaLa: Near Ama-
titlin, Standley 61394 (MICH).

HONDURAS: Moraz&n: Barranco de Suyapa, L. O. Williams & Molina R.
11234 (US, type and isotype).

COSTA RICA: Rodeo de Pacaca toward the River Yarris, Pittier 3228b (US).

This species is close to Melochia nodiflora Sw., but it has considerably
hairier leaves and stems, generally broader bracts, and inflorescences
on naked axillary branches much more commonly than M. nodiflora.

The petals are usually larger than those of M. nodiflora.
27. Melochia tragiifolia Stand]. Contr. U.S. Nat. Herb. 23:804. 1923.

A sparsely branched woody plant at least 0.5 meter high, the young
stems densely covered with glandular hairs 0.2-0.5 mm. long, a few
hairs simple and stellate, 0.2 mm. long, internodes 1.2-6.0 cm. long;
stipules deciduous, 2.5-4.5 mm. long, 1.5-3.5 mm. wide, ovate, ciliate,
hairs simple to 1.2 mm. long, and simple, forked, stellate, and glandu-
lar hairs 0.2-0.3 mm. long on the abaxial surface; petiole slender, 1-2
em. long, pubescence similar to that of the young stems.

Lamina 3.3-5.2 em. long, 2.0-3.2 cm. wide, ovate, the base cordate,
the apex acute, the upper and lower surfaces green, covered with
suberect mostly simple, and a few forked, stellate, and glandular hairs
mostly 0.2 (0.1-0.5) mm. long, not overlapping, also ciliate, velvety to
the touch, the margin irregularly crenate-serrate to dentate, the teeth
1.8-5.0 mm. wide, 0.5-2.5 mm. high, 7-9 pairs of primary veins, 2
are basal.

Inflorescences on slender axillary branches which are up to 7 cm.
long, the flowers in the axils of 1-3 leaves or at up to 3 naked nodes,
3 or 4 flowers in subcapitate umbels at each node, the pedicels 0-3 mm.
long; bracts similar to the stipules in size, shape, and pubescence, but
sometimes only 1.0 mm. wide.

Calyx 5-6 mm. long, 3.2-4.2 mm. wide, not accrescent, hirsute-
tomentose, hairs simple, forked, stellate, and glandular, 0.2-0.7 mm.
long, the teeth 2.9-3.2 mm. long, 1.7 mm. wide, ovate, acute to sub-

GOLDBERG—GENUS MELOCHIA 277

acuminate, the sinus acute; petals reddish (dried), 8.0-8.2 mm. long,
2.4-3.2 mm. wide, obovate to oblanceolate, the claw 1 mm. long.

Longistylous form. Stamens 4.2 mm. long, the filaments united
up to the anthers, the anther 1.3-1.4 mm. long, 0.6-0.9 mm. wide,
oblong, emarginate at both ends; pistil sometimes consisting of 6
carpels, 6.1 mm. long, the styles 4.1 mm. long, filiform, clavate and
papillose at the apex for 0.7 mm., free to the ovary, hirsute at the base,
hairs 0.2 mm, long, the ovary broadly ellipsoid, pentagonal, hirsute,
hairs forked and stellate, 0.2-0.6 mm. long.

Brevistylous form not seen.

Fruit 4.5 mm. long, 5.3 mm, wide, globose, obtusely pentagonal,
the sulci extending a third of way to the center of the fruit, the
midrib subcarinate above the middle of each carpel, covered with
simple, forked, and glandular hairs 0.2-0.7 mm. long, the stipe 0.2
mm. long; dehiscence septicidal into cocci open along the ventral
suture; seeds not quite ripe, 2.7-3.0 mm. long, 1.7-1.9 mm. wide,
red brown, 1 per cell.

Flowering and fruiting specimen collected in March.

Type Locauiry: La Salada, Michoacfén, Mexico. The type was
collected by Nelson, no. 6927, and is cited below.

Distripution: The species is known only from the type locality.

MEXICO: Micuoac4n: La Salada, Nelson 6927 (US, type).

This species is similar to M. nodiflora Sw. and M. nudiflora Stand].
& L. Wms., but it differs in having an abundance of glandular hairs
on the calyx and young stems and some on the leaves; the hairs of the
latter two are not glandular. The petals are larger than those of
M. nodiflora and the leaves are not as hairy as those of M. nudiflora.

28. Melochia tomentella (Presl) Hemsl. Biol. Centr. Amer. Bot. 1:132. 1879.
PuaTE 4
Riedleia tomentella Presl, Rel. Haenk. 2:148. 1835.

A simple or branched, erect, perennial herb to shrub 0.6-2.0 meters
high, the young stems and petioles densely tomentulose, hairs stellate,
forked, simple, and sometimes ‘also glandular, 0.1-0.3 mm. long,
internodes 1.1-3.5 cm. long; stipules soon deciduous, 3.0-5.5 mm.
long, 1.0-1.7 mm. wide, lanceolate, hirsute and ciliate, hairs simple,
forked, and stellate, 0.1-0.5 mm. long; petiole 0.3-1.8 em. long, very
slender.

Lamina 3.0-9.0 cm. long, 1.2-3.9 em. wide, lanceolate to ovate, the
base rounded, the apex acute to long-acuminate, the upper surface
covered with slightly overlapping simple and forked appressed hairs
0.1-0.8 mm. long, the lower surface covered, especially on the veins,
with slightly overlapping stellate, forked, and simple hairs 0.1-0.6

278 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

mm. long, the margin irregularly crenate-serrate to crenate-dentate,
serrations 1.5-6.8 mm, wide, 0.5-2.6 mm. high, 5-10 pairs of primary
viens, 2 are basal.

Inflorescences in the axils of 4-12 upper leaves, the clusters sessile or
subsessile, subcapitate, 0.7-2.3 cm. long, the peduncle 0-1 cm. long,
the pedicel 0-2 mm. long, 2-5 flowers per umbel or head, sometimes
also a terminal inflorescence extending 3.5 cm. beyond the first leaf;
bracts numerous, 0.5-3.7 mm. long, 0.2-1.0 mm. wide, deltoid-acute
to nearly linear, hirsute and ciliate, hairs mostly simple and forked,
some stellate and fewer glandular, 0.1-0.7 mm. long.

Calyx 3.1-5.0 mm. long, 3.1-4.0 mm. wide, not accrescent, tomen-
tulose, hairs stellate, forked, simple, and glandular, 0.2 (0.1-0.6) mm.
long, the teeth 1.3-3.4 mm, long, 1.2-1.6 mm. wide, deltoid-acute to
deltoid-acuminate, the sinus acute to slightly rounded; petals pink or
white with pink veins, 5.5-7.8 mm. long, 2.1-3.0 mm. wide, obovate-
cuneate, attached to the staminal tube for 1.7 mm., the claw 1.0-1.5
mm. long.

Longistylous form. Stamens 3.8 mm, long, the filaments united
up to the anthers, the anther 1.3 mm. long, 0.7—-0.8 mm. wide, oblong,
emarginate at both ends, nearly to the middle from the apex; pistil
5.3-6.1 mm. long, the styles filiform, slightly clavate at the apex,
papillose for 0.8 mm., 4.0-4.5 mm. long, free to the ovary, the ovary
ovoid, densely sericeous, hairs 0.3-0.7 mm. long.

Brevistylous form, Stamens 4.8-5.0 mm. long, the filaments free
for 2.0 mm., the anther 1.2-1.3 mm. long, 0.9 mm. wide, shape same
as that above; pistil 3.6-3.8 mm. long, the styles filiform, papillose
for 0.5 mm., 1.7-2.0 mm. long, free to the ovary, the ovary ovoid,
densely sericeous-tomentose, hairs up to 0.8 mm. long.

Fruit 3.0-3.8 mm. long, 4.6 mm. wide, depressed-globose, obtusely
pentagonal, the sulci between the carpels shallow, extending a third
of way to the center of the fruit, densely tomentulose, hairs stellate,
forked and simple, 0.1-0.7 mm. long, stipe none; dehiscence septicidal
into cocci open along the ventral suture; seeds 2.2-2.6 mm. long,
1.3-1.5 mm. wide.

Flowering and fruiting specimens collected from September to
December, and in February.

Type Locauity: The isotypes, cited below, were collected by
Haenke at Acapulco, Mexico.

Distripution: Mexico, in shady, short-tree forest bottoms, associ-
ated with mesquite, palms, and oaks, at 10-600 meters elevation.

MEXICO: Sonora: atAmMos: Arroyo Cuchuhaqui, Gentry 862 M (MICH).
Quiricoba, Gentry 833 M (MICH). Srnazoa: Vicinity of Culiacan, Brandegee
s.n. (US). Falda del C. del Jacalito, San Ignacio, Montes & Salazar 565 (US).

El Norote, Ortega 5884 (US). Imala, Gentry 4968 (MICH). Rio Fuerte, Gentry
943M (MICH). Sr1natoa?: Ymala, EF. Palmer 1669 in 1891 (MICH, 8, US).

GOLDBERG—GENUS MELOCHIA 279

Duraneo: La Bajada, Tamazula, Ortega 4322 (US). GurRrERo: Acapulco,
Haenke s.n. (PR 174404 a-e, isotypes, also US neg. 5548); EH. Palmer 261 in part
1894-95 (MICH, US). Ganzana: Atoyac, Hinton 10908 (MICH, US). Ba-
gueta, Langlassé 495 (US). Sierra Madre del Sur, N. of Rio Balsac, Adama,
Temisco, Barranca del Consuelo, Meria 8782 (S). OAXACA: JAMILTEPEC: From
Collantes to Para Verde, Conzatti 4427 (US). From Jamiltepec to Rio Verde,
Nelson 2375 a (US). Huatulco, Liebmann 588-0 (L, US), 11881 fragment (US).

This species is close to Melochia nodiflora Sw., but the leaves are
usually lanceolate rather than ovate, and acuminate rather than acute,

and the stems and leaves are hairier.

Section III. Melochia
Section Riedleia Griseb. Fl. Brit. W. Ind. 94. 1859.

Perennial or annual herbs to shrubs, 0.2-2 meters high, the taproot
not thicker than the base of the stem or 2-3 times thicker, and 0.8-2.0
cm. wide, the young stems subglabrous to cinereous, hirsute, pilose,
sericeous or tomentose, hairs stellate, sometimes also forked, simple
or glandular, 0.1-3.2 mm. long; internodes 0.3-10.0 em. long; stipules
1.5-10.0 mm. long, 0.2-3.2 mm. wide, usually lanceolate, deltoid-
acuminate or linear, often ciliate, subglabrous to tomentulose, hairs
simple, sometimes stellate, forked, and glandular, 0.1-2.7 mm. long;
petiole 0.07-6.2 cm. long.

Lamina 0.7-10.5 cm. long, 0.2-8.5 cm. wide, ovate, lanceolate,
linear, oblong-ovate, or suborbicular, the base usually rounded, some-
times subcordate, truncate or cuneate, the apex usually acute,
sometimes acuminate or rounded, both surfaces glabrous to canescent,
hairs simple, appressed, or stellate, forked, and glandular, 0.1-1.0
(-2.2) mm. long, the margin irregularly crenate-serrate, serrations
0.3-12 mm. wide, 0.2-2.5 mm. high, 4-13 pairs of primary veins,
usually 2 are basal.

Inflorescences terminal, extending 1-34 cm. beyond the first leaf
and axillary, 0.7-19 em. long, or only terminal or axillary, subcapitate,
spicate or paniculate, rarely corymbiform, the peduncle short, 0-1.9
(4.1) em. long, the pedicels usually short, 0-9 (-14) mm. long;
bracts often long, 0.5-12.2 mm. long, 0.1-1.9 mm. wide, frequently 3
immediately subtending each flower, sometimes 2 united at the base,
similar in shape and pubescence to the stipules.

Calyx usually short, 1.4-6.0 mm. long, 2.0-5.0 mm. wide, sometimes
membranous, hirsute, sericeous or tomentulose, hairs simple, forked,
stellate, or glandular, 0.1-2.3 mm. long, the teeth short, 0.4-2.7 mm.
long, 0.2-2.2 mm. wide, the sinus usually rounded to nearly truncate,
rarely acute, the teeth often distant; petals white to pink, blue, lilac,
violet, purple distally and yellow proximally, red, red striped, yellow
or orange, 2.7-12.7 mm. long, 0.8-5.2 mm. wide, usually obovate or

280 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

oblanceolate, cuneate, free of the staminal tube or attached to it for
0.3-2.0 (-3.0) mm., the claw 0.2-1.5 (-4.0) mm. long; anther 0.4-1.7
mm. long, 0.3-0.9 mm. wide, oblong, emarginate at both ends, nearly
to the middle from the apex, staminodes rarely present, 0.2-0.4 mm.
long.

Longistylous form. Stamens 2.6-6.2 mm. long, the filaments
united up to the anthers; pistil 3.6-9.0 mm. long, the styles free for
1.5-6.0 mm., to the ovary, or united for 0.5-3.0 mm., papillose for
0.7-2.5 mm. at the apex.

Brevistylous form. Stamens 4.7-8.0 mm. long, the filaments free
for 1.7-4.3 mm.; pistil 2.3-6.2 mm. long, the styles free for 1.0-3.8
mm., to the ovary, or united for 0.3-1.8 mm., papillose for 0.7-0.8
mm. at the apex; the flowers of several species not dimorphic, stamens
and pistils 1.7-3.2 mm. long, the filaments free for 0-1.2 mm., the
styles free for 0.6-1.2 mm., to the ovary, or united for 0.1-0.3 (-0.7)
mm.

Fruit 2.4-5.0 mm. long, 3.2-5.5 mm. wide, globular, pentagonal,
the sulci between the carpels shallow, extending a fifth to a fourth of
way to center of the fruit, often hirsute-tomentulose, hairs simple,
sometimes also forked, stellate, or glandular, 0.1-1.5 mm. long, stipe
usually none, rarely 0.2-1.0 mm. long; dehiscence all along the dorsal
and ventral sutures, tardily septicidal, the fruit falling apart; seeds
1.4-3.2 mm. long, 1.0-1.9 mm. wide, trigonal, brown to black,
usually 1 per cell.

The flowering and fruiting periods are usually long, sometimes
throughout the year.

Tyre species: Melochia corchorifolia L.

Disrrisution: United States to Argentina, West Indies, Africa,
Asia, Pacific Islands, adventive elsewhere in savannas, thickets, or
fields, in wet or dry habitats, at 0-2,300 meters elevation.

29. Melochia villosa (Mill.) Fawe. & Rendle, Fl. Jamaica 5:165. 1926. PLATE 5

Usually a perennial, sometimes annual, erect, spreading, decumbent
or procumbent, usually branched herb, sometimes suffrutex, rarely a
small frutex, sometimes with two to several slender shoots from near
the base, 0.3-2.0 meters high, the root as thick as the base of the
stem, sometimes 2-3 times thicker and 0.8-2.0 cm. wide, the young
stems with a large pith, sometimes hollow, hirsute, villous, lanate,
sericeous or tomentose, hairs sometimes ferruginous, tan or golden,
simple, forked, glandular, sometimes stellate in longitudinal lines,
sometimes predominantly stellate all around the stem, 0.1-2.0 (-2.5)
mm. long, internodes (0.3-) 0.6-6.5 cm. long; stipules (2-) 4.2-9.5
mm. long, (0.3—-) 0.6-1.7 (-3.1) mm. wide, deltoid- or ovate-acuminate
or lanceolate, rarely sublinear or oblong, hirsute-tomentose or seri-

GOLDBERG—GENUS MELOCHIA 281

ceous, ciliate, hairs simple, sometimes also forked, stellate, and glan-
dular, 0.1-2.0 mm. long; petiole 0.1-6.2 cm. long, pubescence similar
to that of the young stems.

Lamina 1.5-10.5 cm. long, 0.9-8.5 em. wide, the basal laminae
sometimes smaller, usually ovate, sometimes oblong-ovate, lanceolate
or suborbicular, the base usually rounded to subcordate, sometimes
cordate or truncate, the apex usually acute, sometimes rounded, the
upper surface green and subglabrous to hirsute, villous, sericeous, or
tomentose and tan-canescent, hairs simple, sometimes also a few
forked and stellate, or entirely stellate, rarely as many as 10-12 rays
per hair, 0.2-2.0 mm. long, the lower surface green and subglabrous
or tan- or ferruginous-canescent, and sericeous or tomentose, hairs
simple, glandular, sometimes also forked and stellate or entirely
stellate, 0.1-2.0 mm. long, sometimes both surfaces velvety or woolly
to the touch, the margin irregularly crenate-serrate, serrations
0.6-4.1 mm. wide, 0.2-2.0 mm. high, (4-) 5-12 pairs of primary
veins, usually 2 (-3) are basal, the second pair stronger, veins promi-
nent beneath.

Inflorescences predominantly terminal, usually spicate, sometimes
subcapitate, usually coarctate, sometimes lax, extending 1.0-33.5
cm. beyond the first leaf, the flowers at 2-28 nodes, usually 1-5 (-15)
flowers at a node, the internodes 0.1-3.0 em. long, also axillary, 0.7-1.0
cm. long, at 1-6 nodes, rarely flowers at nodes not subtended by
leaves on the middle portion of the stem or the terminal inflorescence
interrupted by leaves at about 3 nodes, peduncle usually none,
sometimes 0.1 cm. long, pedicels none or very short, 0-1.6 mm.
long; bracts usually long, 3.5-11.0 mm. long, 0.3-1.0 (-2.5) mm.
wide, deltoid-acute to deltoid-acuminate, ovate-lanceolate or lanceo-
late to sublinear, sometimes subfalcate, 2 or 3 subtending the abaxial
side of each flower, merely touching or united at their very base,
usually one more below, ciliate and hirsute, sericeous or tomentose,
hairs simple and sometimes glandular, forked, or stellate, 0.22.0
mm. long.

Calyx 3-6 mm. long, 3-5 mm. wide, hirsute, sericeous or tomentose,
sometimes ciliate, hairs simple, sometimes also forked, stellate, or
glandular, 0.1-2.0 mm. long, the teeth 0.8-2.5 mm. long, 0.4-2.1 mm.
wide, deltoid- to ovate-acute or ovate-acuminate, the sinus acute to
rounded and the teeth then rather distant; petals usually pink or
purple, sometimes violet, blue, rarely yellow, white, white distally
and yellow proximally, or white with pink stripes, 6.0-12.7 mm. long,
1.9-4.5 mm. wide, usually obovate-cuneate, sometimes oblanceolate-
cuneate, attached to the stamens for 1.5-3.0 mm., the claw 1-3 mm.
long.

282 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Longistylous form, Stamens 3.5-6.2 mm. long, the filaments
united up to the anthers, scattered simple, sometimes also forked,
hairs 0.1-0.3 mm. long on the tube, the anther 1.1-1.7 mm. long,
0.5-0.9 mm, wide, oblong, emarginate at both ends, nearly to the
middle from the apex, sacs slightly spreading at the apex; pistil 5.7—
9.0 mm. long, the styles free for 3-5 mm., united for 0.7-1.7 (-2.7)
mm., papillose at the apex for 1.8-2.5 mm., the ovary ellipsoid, ovoid
or globular, the ovary, particularly the apex, and base of styles rather
densely sericeous-tomentose, hairs simple, sometimes also forked and
stellate, 0.4-1.5 mm. long, stipe none.

Brevistylous form. Stamens 6-8 mm. long, the filaments free
for 2.0-3.4 mm., glabrous or with scattered hairs 0.1-0.2 mm. long,
the anther 1.0-1.7 mm. long, 0.5-0.9 mm. wide, shape similar to that
above; pistil sometimes consisting of 6 carpels, 3.8-5.2 mm. long, the
styles free for 1.5-2.2 mm., united for 0.8-1.5 mm., papillose for 0.8
mm., the ovary ovoid or globular, densely sericeous, hairs 0.5-1.5 mm.
long.

Fruit 3.0-3.3 mm. long, sometimes a rostrum 1 mm. long present,
3 4-3.9 mm, wide, globular, slightly depressed, obtusely pentagonal,
the sulci between the carpels very shallow, extending a fifth to a
fourth of way to the center of the fruit, hirsute-tomentulose, mostly
near the apex, hairs simple, sometimes also forked and stellate, 0.1-
1.5 mm. long, stipe none; dehiscence loculicidal, first all along the
dorsal, then along the ventral suture, tardily septicidal, the fruit
falling apart; seeds trigonal, 1.7-2.2 mm. long, 1.1-1.7 mm. wide,
reddish brown, usually 1 per cell.

Flowering and fruiting specimens collected throughout the year.
According to Gray (20) anthesis is matutinal.

Key to Varieties of M. villosa

1. Leaves and stems tomentose or villous-tomentose, tan- or ferruginous-
canescent, hairs predominantly or exclusively stellate.
29c. M. villosa var. tomentosa
1. Leaves and stems hirsute-sericeous to subglabrous, hairs predominantly or
exclusively simple.
2. Internodes of terminal inflorescence usually short; bracts typically 1.5 mm.
wide; stems not pubescent in 2 longitudinal lines.
29a. M. villosa var. villosa
2. Internodes of terminal inflorescence long; bracts often up to 2.5 mm. wide;
stem pubescent in 2 longitudinal lines (Minas Gerais, Brazil).
29b. M. villosa var. regnellii.

29a. Melochia villosa (Mill.) Fawe. & Rendle var. villosa
Sida villosa Mill. Gard. Dict. ed. 8. 1768.
Melochia hirsuta Cav. Diss. 6:323, pl. 175, fig. 1. 1788.
Riedlea serrata Vent. Choix. 37, pl. 37, 1803.
Mougeotia hirsuta (Cav.) H.B.K. Nov. Gen. & Sp. 5:331. 1823.

GOLDBERG—GENUS MELOCHIA 283

Kiedleta hirsuta (Cav.) DC. Prodr. 1:492. 1824,

Melochia lilacina St.-Hil. Fl. Bras. Merid. 1:162. 1825.

Visenia hirsuta (Cav.) Spreng. Syst. 3:30. 1826.

Visenia serrata (Vent.) Spreng. Syst. 3:30. 1826.

Riedleia serrata Vent. 8 glabrescens Presl, Rel. Haenk. 2:147. 1835.

Riedleia elongata Presl, Rel. Haenk. 2:148. 1835.

Melochia vestita Benth. in Hook. Journ. Bot. 4:130. 1841.

Melochia serrata (Vent.) St.-Hil. & Naud. Ann, Sci. Nat. II, 18:36. 1842.

Melochia Clinopodium St.-Hil. & Naud. Ann. Sci. Nat. II, 18:36. 1842.

Melochia densiflora Mig. Linnaea 22:800. 1849.

Kiedleja Cubensis Turez. Bull. Soc. Nat. Mosc. 31(1):209. 1858.

fiiedleja Jurgensenit Turcz. Bull. Soc. Nat. Mose. 31(1):211. 1858.

Riedleja heterotricha Turcz. Bull. Soc. Nat. Mosc. 31(1):211. 1858.

Riedleja tenella Turez. Bull. Soc. Nat. Mose. 31(1):212. 1858.

?Riedleja scutellarioides Turez. Bull. Soc. Nat. Mose. 31(1):210. 1858.

Melochia jurgensenii (Turcz.) Hemsl. Biol. Centr. Amer. Bot. 1:131. 1879.

? Melochia scutellarioides (Turez.) Hemsl. Biol. Centr. Amer. Bot. 1:131. 1879.

Melochia tenella (Turcz.) Hemsl. Biol. Centr. Amer. Bot. 1:131. 1879.

Melochia cephalodes Schum. in Mart. Fl. Bras. 12(3):45. 1886.

Melochia hirsuta Cav. var. grandiflora Schum. in Mart. Fl. Bras. 12(3) :46.
1886.

Melochia hirsuta Cav. var. rotundifolia Schum. in Mart. Fl. Bras. 12(3) :46.
1886.

Melochia hirsuta Cav. var. glabrata Schum. in Mart. Fl. Bras. 12(3) :47.
1886.

Melochia hirsuta Cav. var. glabrescens (Presl) Gray Syn. Fl. N. Am. 1(1):340.
1897.

Melochia hirsuta Cav. var. paraguayensis Chod. Bull. Herb. Boiss. II, 1:403.
1901 (as var. paraguagensis).

Riedlea glabrescens (Presl) Small. Fl. SE. U.S. 780. 1903.

The leaves of the typical variety are hirsute-sericeous to sub-
glabrous and not canescent.

Type Locauity: Jamaica, Méiller’s type is in the herbarium of
the British Museum.

Disrrisution: Southern Florida, Jalisco and Veracruz, Mexico
to Paraguay and Sao Paulo, Brazil, the Greater Antilles, primarily
in savanna and pineland, but also in various other habitats, in wet or
dry clay, sandy, loam, or serpentine soil, at 0-2,300 meters elevation.

UNITED STATES: “California ad Monte-Rey” Haenke s.n. (PR 24355, type
of Riedleia elongata Presl, also US neg. 5549). The locality is probably incorrect;
it is probably somewhere in Mexico. Fioripa: Near Trabue?, Simpson 393 (US).
South Florida, A. W. Chapman s.n. ?754 (US). ex: Fort Myers and vicinity,
Hitchcock 26 (US); P. C. Standley 12615 (US), 12961 (US), 18964 (US); J. P.
Standley 235 (US), 300 (US); Buswell 1358a (S). Near Okaloacoochee Slough,
Big Cypress, Small 8292 (US). Alva, Francis 96 (US). Caloosa River, Garber
47 (8S, US); Curtiss 370 A (US), A (S, US). Caloosahatchee, D. Chapman s.n.
(US). HENDRY: La Belle, Brass 15468 (US). niaHLANDs: Bootheel Creek,
Venus, Brass 20562 (US).

MEXICO: Haenke (W). La Laja, Liebmann 588 K (US), 11875 (US), 11877
(US). Sierra San Pedro Nolasco, Talea, etc., Jurgensen 506 (K, isotype of
Riedleja tenella Turez., also US neg. 5414). Passo-Majo, Wawra 816 (W).

284. CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Nayarit: About 21 mi. SE. of Tepic, Gentry 18343 (US). Jaisco: Guadalajara,
Tequila, E. Palmer 287 in 1886 (US). Lowlands near Guadalajara, Pringle 4452
(MICH, 8S, US, W). Veracruz: Mirador, Sartorius s.n. (W). Zacuapan, Purpus
12071 (US), 2028 (US), 10877 (US). Xalapa, Coulter 793 (GH). Cuitlahuac,
Matuda 1443 (MICIL). san anpr&s TuxrLa: Between Los Mangos and San
Juan de Azicar, Dressler & Jones 205 (US). Mexico: TEMASCALTEPEC: Cerro
Mufieca, Hinton 1372 (US). Ixtapan, Hinton 1643 (S, US). Micnoacdn:
Coalcoman, Hinton 12206 (US). GUERRERO: GALEANA: Moreno, Hinton 14343
(MICH, US). Montes de Oca: Vallecitos, Hinton 11371 (US). Oaxaca: Near
Santo Domingo, Nelson 2706 (US). Rincén Antonio, Orcutt 3271 (GH, US).
From Cordoba to Santa Lucrecia, Conzatti 1716 (US). About 2 km. E. of Zante-
pec, King 1877 (US). Curtapas: Chicomuselo, Matuda 4430 (US). Acapetagua,
Matuda 16684 (MICH). Near the junction of Rio Perlas and Rio Jataté at San
Quintin and near Laguna Miramar, Sohns 1647 (US).

GUATEMALA: Los Amates, Deam 121 (MICH). Perrin: Big Fall Pine Ridge,
Lundell 4185 (8). La Libertad and vicinity, Lundell 2301 (MICH), 2389 (MICH),
2612 (MICH); Aguilar H. 172 (MICII). Sabana Zotz, Lundell 3582 (MICH).
HvEnvETENANGO: Cerro Victoria, across the river from Finca San Rafael, Sierra
de los Cuchumatanes, Steyermark 49569 (US). ALTA VERAPAZ: Cubilgiiitz,
Tiirkheim 7615 (US), IL 49 (S). Cajabon near Finca Sepacuite, Cook & Griggs
330 (US). Between Cajabon and Secanquim, Goll 108 (US). Chamd, Johnson
836 (US). Ritzul, between Cajabon and Lanquin, Goll 120 (US), 131 (US), 1382
(US). IzaBau: Cristina, Blake 7568 (US). Sanra Rosa: Las Vifias, Heyde &
Lua 6074 (US). Rio de las Caiias, Heyde & Lua 3953 (US).

BRITISH HONDURAS: Record s.n. (US). Honey Camp, Coastal Region,
Lundell 585 (US). Maskall Pine Ridge, Gentle 1044 (MICH, 8). Butcher Burn,
Sibun River, Bartlett 11413 (MICH, S), 11884 (MICH). All Pines, Schipp 549
(MICH, 8). Belize River, Lundell 4185 (MIC). Near Manatee Lagoon, Peck
297 (GH). Beruze: Baker’s Pine Ridge, Lundell 3793 (MICH). ‘ToLrEpo:
Swasey Branch, Monkey River, Gentle 3771 (MICH).

HONDURAS: Corn: Hac. Espiritu Santo to Quebrada Majanales, Blake
7463 (US). Comayaaua: Vicinity of Siguatepeque, Standley 56272 (US);
Yunker et al. 5540 (MICH, 8).

SALVADOR: Moraz&n: About 1.6 km. SE. of Divisadero, Tucker 580 (US).

NICARAGUA: Slopes of Mt. Mombacho, near Granada, Grant 758 (GH).

COSTA RICA: Hac. ‘La Argentina,” Grecia, Valerio 600 (US). Buenos
Aires, Tonduz 4028 (US). Infiernillo cerca de Juan Vitas, Tonduz 10416 (US).
Boruca, Pittier 4583 (US). Rodeo, Pittier 1647 (US). Las Céncavas, near
Cartago, Cooper 13 (US); Lankester 323 (8); Stork 2145 (MICH). Paraiso
Cartago, Torres 101 (US), 191 (US).

PANAMA: Curriguf: San Carlos, Roy 4 (MICH, US). Sabana de la
Tortuga, between El Boquete and Caldera, Pittier 3304 (US). Vicinity of
Boquete, W. L. Stern et al. 1175 (US). Cocifi: Aguadulce, Pittier 4858 (US).
Cana Zone: Along the old Las Cruces Trail, between Ft. Clayton and Corozal,
Standley 29087 (US). Chiva-Chiva Trail, Red Tank to Pueblo Nuevo, Piper,
5757 (US). Panamd: Sabanas and vicinity, Bro. Paul 74 (US); Standley
25845 (US), 25934 (US); Bro. Gervais 159 (US); Killip 3122 (US). Rio Tapia,
Standley 28300 (US). Near Rfo Tapia, Juan Diaz, Maxon & Harvey 6632 (5,
US). Between Matias Herndndez and Juan Diaz, Standley 28897 (S, US),
31980 (US). Matias Hernandez, Pittier 6797 (US), 6875 (US). Sabana de
Juan Corso, near Chepo, Pittier 4510 (US). Along the road between Panama
and Chepo, Dodge et al. 16660 (S). Nuevo San Francisco, Standley 30748 (US).
Near the big swamp E. of the Rfo Tecumen, Standley 26506 (US).

GOLDBERG—GENUS MELOCHIA 285

CUBA: Poeppigs.n. (8). dela Sagras.n.(W). Laguna los Piedras, ?Poeppig,
collected 1824 (W). 5S. Cristobal, C. Wright 2086 (S, US), 2088 (S, US). Isuiz
or Pines: Taylor 94 (US). Near Nueva Gerona, Curtiss 218 (L, US);
W. Palmer & Riley 867 (US); Killip 42538 (US). Near Managua, W. Palmer
& Riley 1079 (US). Las Tunas, Britton & Wilson 14745 (US). Santa Ana,
Ekman 12206 (8). Pinar pet Rfo: Pinar del Rfo, Pinar del Cajalbana, Ekman
10488 (S). Arroyo del Sumidero, Shafer & Bro. Leon 13567 (US). Palm barrens
W. of Guane, Shafer 10418 (US), 10645 (US). N. of San Diego de los Bafios,
W. Palmer & Riley 630 (US). El Guama, W. Palmer & Riley 129 (US). S.
of Las Mangas, Ekman, 13060 (8). Between Candelaria and Artemisa, Wilson
1716 (UC). Las Vituas: Near Santa Clara, Howard 5600 (US). cIrENFUEGOS:
Cieneguita, Combs 217 (US). Camagunry: Sierra Cubitas to Santa Rosa, Shafer
548 (US). Orrente: Sierra Maestra at Pinar de Caridad, SE. of Yara, Ekman
14685 (8).

HAITI: Depr. pu Norp: Vicinity of Dondon, Leonard 8649 (US). Massir
pu Norp: Le Borgre, Poste Marie Congo, Ekman H. 4844 (8, US). Massif
de la Hotte, western group, Corail, at Dutreuil, Ekman H. 10745 (8).

DOMINICAN REPUBLIC: Loma Mata Clara, Hggers 2336 (L, US). La
Nassa, Faris 135 (US). La Leonor, J. S. Jiménez 2407 (US). Sanra Dominao:
Vicinity of Ciudad Trujillo, Allard 14237 (US), 14232 (US). Sabana de Santa
Rosa, Allard 15892 (US), 15942 (US), 16023 (US). Moca: Villa Altagracia,
Ekman H. 11230 (8, US). Duarte: 33 km. de la Carretera, J. Jiménez 821
(US). Valle de Cibao, Pimentel, Ekman H. 18239 (8S). La Veaa: Sierra Prieta,
J. S. Jiménez 1148 (US). Trusitio: Vicinity of Piedra Blanca, Allard 13503
(US), 13831 (S, US), 18896 (US). Paciricapor: Villa Riva (Almacén), Abbott
536 (US), 626 (US). Serso: Vicinity of Higiiey, R. A. & HE. S. Howard 9874
(US). Guarabo, SE. of Jovero, Abbott 2762 (R, 8S, US), 2820 (US).

JAMAICA: W. Wright s.n. (W). Oxford, Harris 9428 (US). CLARENDON:
Clarendon Park to Four Paths, Britton 3788 (US). James Hill Savanna, Harris
12842 (US). Kellits, Harris 11151 (US). Hollis’ Savanna, Harris 12251 (8, US).
Hanover: Orange Bay Morass, Macfadyen s.n. (S).

PUERTO RICO: Sintenis 5922 (US). Goll 43 (US). Stahl 350 (L). Near
Cabo-Rojo at Port Real, Sintenis 162 b (L, US, W). Near Santurce, Heller 251
(US), 4671 (US); Cowles 158 (US). Moca, Sargent 248 (US). Rio Piedras,
Stevenson 112 (US). Sabana Abajo, near Carolina, Britton & Brown 5712 (US).
Bayamon, Underwood & Griggs 865 (US). Along the road to Rosario, Las Mesas,
Theoslohn 219 (GH).

TRINIDAD: Piarco SAvaANNan: Broadway 2132 (US); Warming 262 (US).
S. of Arouca, Britton et al. 58 (US). 8. of Dabadie, Britton & Hazen 695 (US).

FRENCH GUIANA: Le Prieur 129? (L). Cayenne and vicinity, Broadway
152 (US), 608 (US); Sagot 55 (S, W). Maroni, Melinon 45 (US).

SURINAM: Bergendaal, Focke 359 (L). Epereroe, Pulle 495 (L, US).

BRITISH GUIANA: Schomburgk 133 (W, syntype of Melochia vestita Benth.,
also F neg. 35388 ex. P). Between Takutu River and Kanuku Mts., A. C. Smith
3267 (S, US). Rupununi: 45 mi. N. of Lethem, Jrwin 835 (US).

VENEZUELA: Sucre: Cumana, Guanaguana, Funck 170 (LE, isotype Of an
unnamed variety of Riedleja heterotricha Turez.). Los Altos, Tamayo 2163 (US).
AnzoAteGut: Bajos del Rio Tigre, in Cristovero, SE. of San Tomé, Pittier 14595
(US). Rio Caris near El Tigre, Pittier 14321 (US). Araaua: Colonia Tovar,
Fendler 19385 (GH). Above Guamitas, Li. Williams 10356 (US). Valle de
Guamitas, Parque Nacional, Pittier 15251 (US). Carasoso: Near San Joaquin,
Pittier 8007 (US). TrusttLo: Near La Ceiba, Reed 949 (US). Zuxta: Vicinity
of Mene Grande, Pittier 10575 (GH, US). Apursn: Puerto Paez, Velez 2724 (US).

227-051—67—_—_7

286 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Botfvar: Morichal Santa Isabel, Cardona 660 (US). Calcetas en la Pica de El
Cristo, Cardona 1020 (US). Rio Aprada afluente del Uriman, Caroni, Guayana,
Cardona 2002 (US). GRAN SABANA: Cercanias de Santa Elena, Tamayo 3215
(US). ?BoLfvar: Camarata Valley, Simpson 66 (US). Amazonas TrErR.:
Maypures, Spruce s.n. (Kx).

COLOMBIA: Linden 887 collected 1842 (W). MaapaLtena: La Jagua,
Haught 3648 (US). ?MagpaLEena: Queremal Jagua, Arbeldez 3010 (US). Norre
DE SANTANDER: Ocafia, Schlim 94 (P). REGION DEL SARARE: El Barco, Cuatre-
casas 18174 A (US). SANTANDER: SAN GIL: Hacienda de La Mesita, Langenheim
3312 (US). Cunpinamarca: Paime, Bro. Ariste-Joseph A802 (US). Melgar,
Pennell 2918 (US). Ibama, E. of Yacopi, Grant 9143 (US). El Colegio, Schneider
928 (S). Mera Inv.: Villavicencio and vicinity, Apiay, Humbert & Fernandez
27182 (P), 27193 (P); Jaramillo el al. 293 (US). E. Mico Airstrip, E. savanna
before Rio Guejar, Philipson et al. 13816 (US). Boca de Monte, Llanos de San
Martin, S. G. Smith & Idrobo 1438 (US). ARAUCA: RIO CASANARE: Esmeralda,
Cuatrecasas 3959 (US). (This specimen approaches M. villosa var. tomentosa).
Vaupks: Cano Cuduyari, Cuatrecasas 7225 (US). Rio Vaupés, at Mitu, Hum-
bert & Fernandez 27211 (P, US). Touma: Mariquita and vicinity, Garcia Barriga
8128 (US); Pennell 3659 (US). W. of San Lorenzo, Pennell 3459 (US). Huina:
Cordillera Oriental, E. of Neiva, Rusby & Pennell 476 (US). N. of Neiva, upper
basin of Rio Magdalena, S. G. Smith 1120 (US). San Augustin, Bro. Daniel
4132 (US). One km. SW. of San Antonio Fortalecillas, Fosberg 19833 (US).
Antioquia: Medellin, Archer & Aguilar 702 (US). VALLE DEL Cauca: Rio Bolo,
Dryander 337 (US). Palmira, Pennell & Killip 6045 (US). Timba, von Sneidern
1268 (8S, US), 1185 (S). Samaria, 2 km. W. of Timba, Fosberg 20522 (US).
Cauca: El Tambo, Jdrobo & Fernandez 123 (US). Jamundi, Cauca Valley,
Pittier 1508 (US). Aganche, Pennell & Killip 8318 (US).

ECUADOR: “Yapio cerca Quilichao,” Jameson s.n. (US). Cafiar, Rose et al.
23793 (US).

PERU: San Martin: San Roque, Ll. Williams 7102 (8, US); Melin 123 (S).
Tarapoto, Ll. Williams 6149 (8); Ule 6528 (L). Zepelacio, near Moyobamba,
Klug 3705 (8S, US). Rioja, Woytkowski 6084 (US).

BOLIVIA: Beni: Lake Rogagua, Cardenas 1628 (US). Santa Cruz: SARA:
Buena Vista, Steinbach 6943 (5S), 5104 (A).

BRAZIL: Martii Herb. Fl. Bras. 582 (L). Pohl s.n. (W). Pirapora, Independ-
encia, Burret et al. 10104 (R). Western part, Tamberlik s.n. (W). Rio Branco:
Serra Pellada, Ule ?7634 (L). Amazonas: Séo Gabriel, Rio Negro, Holt &
Blake 625 (RB, US). Rio Igana, Tunuf, Black 48-2597 (US). Par: Vicinity of
Santarem, Spruce June 1850 (W). Almeirim, Rio Arayollos, Ducke s.n. (RB).
Praut: Gardner 2488 (US, syntype of Melochia vestita Benth.). Banta: Glocker
425 d(S). Blanchet 1588 (W). Jacobina, Blanchet 3663 (P, US, W, also US neg.
5666 ex P). Serra do 8. Antonio, F neg. 9599 ex B of Sello 1935, syntype of
Melochia cephalodes Schum. (US). Minas Grrats: Lassance-Pirapora, Cochran
s.n. (R, US). Corinto, Mexia 5530 (S, US). Rio Arassuahy, Glaziou 12460 (R).
Morro de Caxambt, Glaziou 15849 (R). Sao Sebastifio do Paraiso, Brade &
Altamiro 45 (R). Lagoa Santa, Hoehne 6140 (R). 8. Jodo d’El Rey, Casa da
Pedra, Armond s.n. (R). Caldas, Mosén ?4023 (S), 4024 (8S). rruruTaBa:
S. Terezinha, Macedo 1643 (RB), 3177 (US). Certad, Caeduis, Tapeira, near S.
Francisco River, St.-Hilaire 271823 (P, isotype of Melochia lilacina St.-Hil., also
US neg. 5672). Gords: Gardner 3030 (US, W). Planalto, Glaziow 20734 (8).
Between Conceigéio and Natividade, Burchell 8168 (GH). At Megaponte, Rio
Uruht, Pohl 1096 (W). Twenty km. N. of Sao Joao da Alianga, region of the
Chapada dos Veadeiros, Dawson 14247 (US). Mato Grosso: Cuyabd, Malme

GOLDBERG—GENUS MELOCHIA 287

2633 (S). Coxipé da Ponte O. (Cuyabd), Hoehne 2982 (719) (R). Tapirapoan,
Hoehne 1280 (R). SAo Pavuno: Near Batataes, Riedel & Luschnath 2243 (LE).
Ita, Riedel & Luschnath 1992 in part (LE, US neg. 5274, det. as M. hirsuta Cav.
var. macrophylla Schum. by Schumann). Moji-Guagu, Eiten & Machado de
Campos 1534 (US).

PARAGUAY: In regione cursas superior flumenis Y-ac4, Cordillera Central
near Piribebuy, Hassler 6748 in part (P, S).

This species is related to Melochia splendens St.-Hil. & Naud., but
the inflorescence of the latter is primarily axillary rather than primarily
terminal. It is also close to Melochia savannarum Britt., but the latter
generally is a smaller, more prostrate plant with smaller leaves and a
shorter terminal inflorescence with fewer flowers at a node.

29b. Melochia villosa (Mill.) Fawe. & Rendle var. regnellii (Schum.) A. Goldberg,
comb. nov.
Melochia hirsuta Cav. var. Regnellit Schum. in Mart. Fl. Bras. 12(3):46, pl.
10. 1886.

The leaves of this variety have few, scattered, mostly simple hairs;
the stipules are broader than those of the other varieties; the stem is
pubescent in two longitudinal lines; and the internodes of the ter-
minal inflorescence are long.

Typs nocaLity: Caldas, Minas Gerais, Brazil. The lectotype,
Regnell III 278, is cited below.

DistriBuTion: Minas Gerais, Brazil.

BRAZIL: Minas Gerais: Widgren s.n. (R, 8). Caldas, Regnell III 278
(S, lectotype of the variety; R, US, isotypes of the variety); Lindberg 284 C(S).

29c. Melochia villosa (Mill.) Fawc. & Rendle var. tomentosa (Schum.) A.

Goldberg, comb. nov.

Melochia hirsuta Cav. var. macrophylla Schum. in Mart. Fl. Bras. 12(3):47.
1886.

Melochia hirsuta Cav. var. calophylla Schum. in Mart. Fl. Bras. 12(8) :47.
1886.

Melochia hirsuta Cav. var. tomentosa Schum, in Mart. Fl. Bras. 12(3):47.
1886.

Melochia subcordata Morong, Ann. New York Acad. Sci. 7:62. 1892.

Melochia hirsuta Cav. var. tomentosa Schum. f. ferruginea, f. albiflora, and
f. minus induta, all Schum. & Hassl. Bull. Herb. Boiss. II, 4:72. 1903.

The leaves and stem of this variety are much hairier than those of
the typical variety. They are tomentose or villous-tomentose, and
usually tan- or ferruginous-canescent. The hairs are predominantly
or exclusively stellate, in the other varieties they are predominantly
or exclusively simple.

TYPE tocauity: Goids and Sao Paulo, Brazil. The lectotype,
Riedel no. 1992 dupl. 16, is cited below.

DistripuTion: Venezuela to northern Argentina.

VENEZUELA: APURE: ALTO-APURE: Hato Matapalo, Jahn 212 (US).
BRAZIL: Riedel 1992 dupl. 16 (W, lectotype). Regnell III 277a (S). Villa

288 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Boa, Pohl 1232 (W). Western Brazil, Tamberlik sn. (W). ?R1o0 Branco:
Kuhlmann 417 (US). Minas Gerais: Claussen 96 (GH, P, 8). Pires & Black
3064 in part (US). Uberaba, Regnell III 277 in part (S, US). Bello Horizonte,
Barreto 7858 (R). Bello Horizonte and Bairro do Bomfim, Magalhdes 78 (R).
?Morro de Caxambt, Glaziou ?15849 (K). Lagoa Santa, Warming s.n. (S).
Porto Alegre, Regnell III 277 in part (8S). Before Rib. do Frada, Pohl 3281 (W).
sfrro: 3 km. from Sérro, L. O. Williams & Assis 6851 (US). Rio das Pedras
ao Valli, Glaziow 18901 (R). Gots: Mossamedes, Ule 487 (R, US). Near
city of Goids, Burchell 6641 (GH, syntype of M. hirsuta Cav. var. calophylla
Schum.), 6920 (GH). ?Gotds: F neg. 19636 of Pohl? dupl. 517 (US, syntype of
M. hirsuta Cav. var. calophylla Schum.). Maro Grosso: Bomfim, Diogo 268
(R, US), 241 (R). Triumpho, Rio $8. Laurenco, Hoehne 4561 (R). Cuiaba,
Hoehne 4881 (711) (R); Malme 1292 (S). SAo0 Pauto: Batataes, Regnell III 277
in part (8S). Itt, Riedel & Luschnath 1992 in part (LE, US neg. 5267).

PARAGUAY: Near Bellavista (Apa), Hassler 7758 (S, W). Villarrica, Jorgen-
sen 3809 (S, US). Cerro Noaga, Anisits 2846 (8). Between Rio Apa and Rio
Aquidaban, Centurion, Fiebrig 4276 (L). In region of River Alto Parand,
Fiebrig 6392 (US). Road to ‘'Yerbales’” montum ‘‘Sierra de Maracayt,”’ Hassler
5584 (W). Cordillera de Altos, Hassler 3599 (W). Sierra de Amambay, Rojas
in Hassler 10006 (W). Central Paraguay, Morong 292 (US, isotype of Melochia
subcordata Morong). Caazapa: Borja, West 8544 (UC). In regione cursus
superior flumenis Y—Acd, Cordillera Central near Piribebuy, Hassler 6748 in
part (W).

ARGENTINA: Formosa: Colonia Clorinda, Venturi 9098 (US). CorRIENTES:
MouRuCcUYA: Est. Santa Teresa, Tacuaral, Cristébal et al. 10 (UC). GENERAL
paz: Los Cubanos, Schwarz 275 (S). PASO DE LOS LIBRES: Pueblo Paso de los
Libres, Zbarrola 2051 (US). BERON DE AsTRADA: ?Chuicho, Schwarz 333 (US).
MisI1onrEs: Posadas, Loreto, bank of River Alto-Parand’, Ekman 187 (S). At
River Yabebiry, Ekman 1974 (8). canpELaria: Loreto-Rinconada de Yabebiry,
Montes 5?, February 1946 (UC),

30. Melochia savannarum Britt. Mem. Torr. Bot. Cl. 16:81. 1920.

A prostrate or erect herb to suffrutex with several branches from
the base and sometimes branched above, 0.2—-0.6 meter long, the root
woody, thickened, 0.9 cm. wide, the young stems tomentulose, hairs
stellate, 0.1-0.3 mm. long, usually also forked and simple hairs 0.5-1.7
mm. long, few glandular hairs 0.3—-0.6 mm. long, internodes 0.7—2.7 em.
long; stipules 2.2-3.6 mm. long, 0.5-0.7 mm. wide, lanceolate, ciliate,
hairs simple, 0.3-0.8 mm. long; petiole 0.07-0.5 (-0.8) em. long,
pubescence similar to that of the stems but with long hairs more
abundant.

Lamina 1.0—2.2 cm. long, 0.7-1.8 em. wide, ovate, rarely also sub-
oblong, the base rounded to truncate, rarely subcordate, the apex
rounded to subacute, hairs on upper surface simple, appressed, 0.3—1.3
mm. long, sometimes also stellate, forked, sometimes also ciliate, hairs
glandular and simple, 0.5 mm. long, sometimes glabrate, the lower
surface with few to numerous stellate hairs 0.1-0.3 mm. long, and
simple somewhat appressed hairs 0.3-1.0 mm. long, the latter type may
be absent or predominant, the margin irregularly dentate, serrate or

GOLDBERG—GENUS MELOCHIA 289

crenate-serrate, serrations 0.9-4.0 mm. wide, 0.5-2.1 mm. high,
extending to the base of the leaf, 4-6 pairs of primary veins, 1 or 2
are basal.

Inflorescences terminal and in the axils of the upper 1-6 leaves,
subcapitate, 2-6 flowers in a cluster, less than 1 cm. to 4 cm. long,
with flowers at 1-3 nodes, peduncle none, the pedicels 0.5-2.0 mm.
long; bracts 2.5-5.8 long, 0.2-1.0 mm. wide, lanceolate to linear,
usually 3 immediately subtending the flower, others sometimes present
below, rarely 2 united part way, mostly ciliate, hairs simple, 0.3-1.0
mm. long, sometimes also a few glandular and fewer forked and stellate
hairs, 0.3 mm. long.

Calyx 3.0-4.8 mm. long, 3.0-4.7 mm. wide, hirsute-tomentose, hairs
simple, forked, and stellate, 0.1-1.0 mm. long, the teeth 1.0-1.8 mm.
long, 0.8-1.8 mm. wide, deltoid-acute, the sinus rounded to acute;
petals blue, pinkish blue, pink and pale toward the base, nearly white
with a faint tinge of violet, 5.2-9.0 mm. long, 2.1-4.5 mm. wide,
obovate, obovate-oblong, or obovate-cuneate, the claw 0.5-1.0 mm.
long, the base free of staminal tube or attached to it for 0.5-1.5 mm.

Longistylous form. Stamens 3.5-4.4 mm. long, the filaments united
up to the anthers, occasionally a staminode 0.4 mm. long present, a
few simple hairs 0.1-0.2 mm. long on the filament tube, the anther
1.2-1.5 mm. long, 0.6-0.7 mm. wide, oblong, rather deeply emarginate
at both ends, nearly to the middle from the apex; pistil 5.2-6.2 mm.
long, the styles free for 2.0-4.0 mm., united for 0.5 mm., papillose at
the apex for 1.5-2.0 mm., the ovary ellipsoid to ovoid, tomentulose,
hairs 1.0 mm. long, stipe none.

Brevistylous form. Stamens 4.7-6.8 mm. long, the filaments free
for 2.5-2.7 mm., very few simple hairs 0.1 mm. long on the filaments
and filament tube, the anther 1.1-1.4 mm. long, 0.6-0.8 mm. wide,
shape similar to that above; pistil 2.3-4.0 mm. long, the styles filiform,
recurved, free for 1.0-1.5 mm., united for 0.6 mm., the ovary ovoid,
tomentulose, hairs 0.2-1.0 mm. long.

Fruit 3.0-3.2 mm. long, 3.4-3.9 mm. wide, globular, pentagonal, the
sulci between the carpels very shallow, extending about one-fifth of
way to the center of the fruit, hirsute-tomentulose, hairs simple and
forked, 0.1-1.0 mm. long, stipe none; dehiscence loculicidal along the
dorsal suture, eventually also septicidal; seeds 1.8 mm. long, 1.2 mm.
wide (unripe 2.0-2.5 by 1.1-1.9 mm.), trigonal, dark brown, 1 per cell.

Flowering specimens collected from December to April, and in June
and August; fruiting ones, from December to February and in June.

Type Locauity: Herradura, Pinar del Rio, Cuba, in savannas and
along streams; type collected by Earle, no. 639, cited below.

Distrisution: Isle of Pines and Pinar del Rio, Cuba, in white sand,

290 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

tall-grass savanna, in sandy pinelands, in wet or dry habitats, near
sea level and on uplands.

CUBA: IsLE or Prnes: Near Playa Nueva Gerona, Killip 44515 (US). Cerro
de Cunagua, Killip 43669 (US). San Francisco de las Piedras, Killip 43549 (US).
Along the road to Playa, Killip 42996 (US). San Pedro region, Killip 43697
(US). Columbia, Killip 43575 (US). Rfo Jicaro region, Killip 42941 (US).
Playa Roja, Killip 43185 (US). Pinar pet Rio: Herradura, Harle 639 (NY,
type) ; Britton et al. 6518 (US); Ekman 10730 (8S). El Sdbalo, Finca Sabanalamar,
Killip 32253 (US). Sabalo, Ekman 16753 (8S). Near Pinar del Rio, W. Palmer
& Riley 448 (US). Laguna Santa Maria, 8. of Rio Feo, Ekman 17267 (5).

This species is close to M. villosa (Mill.) Fawe. & Rendle, but it is
generally a smaller, more prostrate plant with fewer flowers at a node,
a shorter terminal inflorescence, and smaller leaves. Usually the
apex of the lamina is rounded, but usually acute in M. villosa.

31. Melochia splendens St.-Hil. & Naud. Ann. Sci. Nat. II, 18:36. 1842.

An erect subsimple suffrutex at least 0.45 meter high, the young
stems and petioles densely tomentose, tan-canescent, hairs 0.3-2.3 mm.
long, internodes 1-3.1 cm. long; stipules 5.2-10 mm. long, 1-3.2 mm.
wide, oblique-ovate or lanceolate, sericeous-tomentose, hairs simple,
1 mm. long, and stellate, 0.1 mm. long; petiole short, 0.1-0.4 cm.
long.

Lamina 2.3-5.7 cm. long, 1.1-2.6 cm. wide, lanceolate to ovate,
the base cuneate to rounded, the apex acute to somewhat rounded,
silky and woolly to the touch, both surfaces tan-canescent, densely
tomentose, and sericeous, hairs 0.2-1.5 mm. long, short ones stellate,
long ones simple, the margin irregularly crenate-serrate, serrations
0.6-2.8 mm. wide, 0.2-0.8 mm. high, 8 or 9 pairs of primary veins,
2 are basal.

Inflorescences mostly axillary in the axils of 4-15 leaves, sometimes
a few of the middle ones not subtended by a leaf, sometimes ap-
parently also terminal, 1-1.4 cm. long, subcapitate, in clusters of 2-5
flowers, penduncle none, the pedicels 0-1 mm. long; bracts 10.8—12.0
mm. long, 1.0-1.3 mm. wide, 4 immediately subtending a flower,
2 united at the very base, lanceolate-acuminate, densely tomentose
and sericeous, hairs 0.1—2.0 mm. long.

Calyx 5.0-5.2 mm. long, 2.9-3.2 mm. wide, densely sericeous, hairs
1-2.3 mm. long, some stellate, 0.1 mm. long, the teeth 1.5-2.3 mm.
long, 1-1.5 mm. wide, deltoid-acuminate with rounded sides, the
sinus acute to somewhat rounded; petals 9.6-10.0 mm. long, 1.7—2.0
mm. wide, oblanceolate, attached to the staminal tube for 2 mm.,
the claw 2-4 mm. long.

Longistylous form not seen.

Brevistylous form. Stamens 7-8 mm. long, the filaments free for

GOLDBERG—GENUS MELOCHIA 291

2 mm., a few hairs 0.1-0.2 mm. long on the filament tube, the anther
1.2-1.4 mm. long, 0.6 mm. wide, oblong, emarginate at both ends;
pistil 6-6.2 mm. long, the styles filiform, papillose for 0.7 mm. at the
apex, free for 2.7-3.0 mm., united for 1.7-1.8 mm., the ovary ovoid,
sericeous, hairs 1-2 mm. long.

Fruit (one present) with 4 carpels aborted, 2.5 mm. long, 2.1 mm.
wide, sericeous, hairs 1-1.5 mm. long, stipe none; dehiscence loculicidal
all along the dorsal suture; seeds 2 mm. long, 1.1 mm. wide, brown,
1 per cell.

TypE Locauity: Goids, Brazil. The type was collected by
Gardner, no. 3601, and is in Herbier Delessert.

DistrisuTion: Goids, Brazil, also Piauf, Brazil, according to
Schumann (47), and Arauca, Colombia.

COLOMBIA: Arauca: Rfo CASANARE: Esmeralda, Cuatrecasas 3855 (US).
BRAZIL: Goths: F neg. 9607 ex B of Gardner 3601, isotype (US); Burchell
8587 (GH).

This species is close to Melochia villosa (Mill.) Fawe. & Rendle,
but differs by having axillary rather than terminal inflorescences and
generally narrower leaves, often lanceolate, and cuneate at the base,
rather than ovate leaves rounded or cordate at the base.

32. Melochia bracteosa F. Hoffm. Beitr. Fl. Centr.-Ost-Afr. 13. 1889.
Melochia crinita R. Br. ms. in Hiern Welw. pl. 1:90. 1896.
Melochia Welwitschit Hiern, Welw. pl. 1:90. 1896.
Melochia melissifolia Benth. var. microphylla, var. brachyphylla, var. bracteosa
(F. Hoffm.), var. Welwitschii (Hiern), var. mollis, all K. Schum. in Engl.
Mon. Afr. Pfl. 5:42. 1900.
Melochia mollis Hutch. & J. M. Dalz. Kew Bull. 297, 1928.

An herb or undershrub about 1 meter high, the young stems and
petioles densely covered with simple hairs, sometimes also stellate,
forked, and glandular, 0.2-3.2 mm. long, the internodes 2-6.5 cm.
long; stipules 8-10 mm. long, 1-2.8 mm. wide, lanceolate, ciliate and
hirsute, hairs simple, sometimes also forked, 0.2-2.7 mm. long;
petiole 0.3-2.3 cm. long.

Lamina 3.5-8.5 em. long, 1.2-4.3 em. wide, lanceolate to ovate, the
base rounded to subcordate, the apex acute, both surfaces sericeous,
woolly or velvety to the touch, hairs simple, sometimes also forked
stellate, and glandular, 0.2-2.2 mm. long, the margin irregularly
crenate-serrate, serrations 0.5-3.6 mm. wide, 0.4-2.0 mm. high,
7-11 pairs of primary veins, 2 or 3 are basal.

Inflorescences 1—4.8 cm. long, in the axils of 10-14 or more leaves,
sometimes 2 clusters per axil, 2-10 flowers per cluster, the peduncle
0-3.0 cm. long, the pedicels 0.5-3.5 cm. long; bracts sometimes 2
united at the very base, (3.5-) 5.2-12.2 mm. long, 0.3-1.5 mm. wide,

292 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

lanceolate to sublinear, sometimes falcate, ciliate and hirsute, hairs
simple, sometimes also glandular, 0.2-3.2 mm. long.

Calyx membranous, 3.0-4.5 mm. long, 2.6-4.3 mm. wide, hirsute-
sericeous, hairs simple, 0.2-2.0 mm. long, mostly near the apex, the
teeth 1-2 mm. long, 0.8-1.7 mm. wide, deltoid, acute to acuminate,
the sinus broadly rounded, teeth distant; petals white, 7.5-10 mm.
long, 2-3.5 mm. wide, obovate to oblanceolate, cuneate, attached to
the stamens for 1-2 mm., the claw 1.5-2.0 mm. long.

Longistylous form. Stamens 5.9-6.8 mm. long, the filaments
united up to the anthers, sometimes very few scattered, simple hairs
on the staminal tube, the anthers 1.1-1.4 mm. long, 0.5-0.9 mm. wide,
oblong, emarginate at both ends, nearly to the middle from the apex;
pistil 6.4-8.1 mm. long, the styles filiform, free for 2-6 mm., to the
ovary, or united for 1-3 mm., papillose for 0.7 mm., the ovary globular-
ovoid, sericeous, hairs mostly at the apex, sericeous, simple, 0.5-1.7
mm. long, stipe none.

Brevistylous form not seen.

Fruit 3.0-3.5 mm. long, 3.3-4.0 mm. wide, globular, obtusely
pentagonal, the sulci between the carpels shallow, extending a fifth
to a fourth of way to the center of the fruit, hirsute-tomentose,
hairs simple, 0.2-1.5 mm. long; dehiscence first loculicidal all along
the dorsal sutures and then along the ventral sutures, finally septicidal,
the fruit falling apart; seeds brown, 1.7—-2.3 mm. long, 1.0-1.5 mm.
wide, 1 per cell.

Flowering specimens collected in January, May to July, October
and November; fruiting ones in January, May to July, and November.

Type Locauity: Ugalla River, W. Tanganyika, type F’. Hoffman 275.

DistriBuTion: Sierra Leone, Cameroon, Central African Republic,
Angola, The Congo, Uganda and Tanganyika, in forest, low grassland
swamp, and dry clay soil, at 30-1,216 meters elevation.

SIERRA LEONE: Thomas 8101 (US). Yonibana, Thomas 4191 (8).

CAMEROON: Bipinde, Zenker 1818 (L, 8), 4740 (L). Sourma Cameroon:
Bitye near R. Ja., Bates 604 (US).

CENTRAL AFRICAN REPUBLIC: Haut-Oubangui, between the Yambéré
and Krébedje, Chevalier 6044 (1).

ANGOLA: PorruGuEsE Conao: Sumba, Peco, near Zaire River, Gossweller
8714 (US).

THE CONGO: Kwango, Vanderyst 1418 (US). Bangala, Fr. Hens C 388 (L).
CoquiLHATVILLE: Tkenga, Vebrun 717 (8S). Yangambi, Louzs 16145 (US).

UGANDA: Kivuru, Diimmer 881 (US).

TANGANYIKA: Stromgebiect des Ruhudje, Landschaft Massagati, nordlich
des Flusses, ca 35°30’ E., Schlieben 1092 (S, US, US neg. 5611).

This species is close to M. villosa, M. mellissifolia, and M. splendens.
It differs from the first by having axillary inflorescences rather than
predominantly terminal ones. Its flowers are twice as large as the

GOLDBERG—GENUS MELOCHIA 293

second and its leaves are larger, often hairier and usually have rela-
tively shorter petioles. It differs from the third by having longer
petioles, usually broader leaves, rounded to subcordate at the base
rather than cuneate to rounded, higher serrations, longer pedicels,
a shorter calyx, and broader petals.

33. Melochia melissifolia Benth. in Hook. Journ. Bot. 4:129. 1841.
Melochia concinna Miq. Linnaea 22:467. 1849.
Riedleia concinna (Miq.) Walp. Ann, 2:166. 1852.
Riedleia sparsiflora Klotzsch ex Walp. Ann. 2:166. 1852.

An erect or spreading herb to suffrutex, blooming the first year,
sometimes with several slender stems from near the base, 0.08—1.0
meter high, the taproot as thick as the base of the stem, the young
stems pilose, hairs stellate and simple, occasionally a few glandular
and forked, 0.2-1.0 mm. long, internodes (0.8—) 1.3-3.7 cm. long;
stipules 2.6-5.3 mm. long, 0.6-1.0 mm. wide, lanceolate, mostly ciliate,
hairs simple, 0.3-1.3 mm. long; petiole 0.2-1.5 em. long, hirsute-
pilose, hairs simple and stellate, 0.2-1.5 mm. long.

Lamina 0.5-3.5 cm. long, 0.5-1.6 em. wide, ovate to lanceolate, the
base rounded, the apex usually acute, sometimes rounded, a few
scattered, simple, generally appressed hairs, sometimes a few glandu-
lar, 0.3-1.1 mm. long on both surfaces, especially on the veins of the
lower surface, the upper surface sometimes glabrate and sometimes
reddish, the margin irregularly crenate-serrate, serrations (0.5-)
0.9-2.3 mm. wide, 0.3-1.0 mm. high, 5-8 pairs of primary veins, 2 are
basal.

Inflorescences axillary, consisting of 1-5 flowers in the axils of each
of the upper 6-19 leaves, 0.5 cm. long, sometimes also terminal at 4
nodes, 4 cm. to first leaf, peduncle none, the pedicel 0-2 mm. long;
bracts sometimes united at the base, 3 immediately subtending the
flower, longer than the petals, (2-) 3-6.7 (-8.5) mm. long, 0.3-1.1
mm. wide, lanceolate to linear, mostly ciliate, hairs simple, 0.5-1.5
mm. long.

Calyx membranous, 1.4-2.4 mm. long, 1.7-2.8 mm. wide, with a
few scattered simple hairs 0.3-1.0 mm. long, the teeth 0.4-1.0 mm.
long, 0.4-0.8 mm. wide, deltoid, the sinus obtuse; petals white, pink,
or rose lilac, (1.7—) 2.7-4.0 mm. long, 0.5-1.4 mm. wide, obovate-
oblanceolate, the claw 0.3-0.5 mm. long, almost entirely free from
the staminal tube.

Androecium and gynoecium about equal in length. Stamens (1.2-)
1.7-2.4 mm. long, the filaments free for 0.3-1.2 mm., the anther
relatively small, oblong, 0.3-0.4 mm. long, 0.2-0.3 mm. wide, emar-
ginate at both ends, nearly to the middle from the apex; pistil 1.3-
2.3 mm. long, the styles free to the ovary, 0.6-1.1 mm. long, the
ovary ovoid, densely hirsute-sericeous, hairs 0.3-1.0 mm. long.

994 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Fruit 2.4-2.7 mm. long, 3.2-3.3 mm. wide, subglobose, pentagonal,
the sulci between the carpels very shallow, extending a fifth to a fourth
of way to the center of the fruit, hirsute, hairs simple, 0.1-1.0 mm.
long, stipe none; dehiscence loculicidal along the dorsal suture and
then along the ventral, eventually also septicidal, the fruit falling
apart; seeds 1.4-1.8 mm. long, 1-1.3 mm. wide, trigonal, dark brown,
1 per cell.

Flowering specimens collected from March to May, and July to
January; fruiting ones, from March to July, and September to
January.

Type Loca.ity: British Guiana, syntypes collected by Schomburgk,
no. 366, and French Guiana, syntypes collected by Leprieur, no. 122,
one cited below.

Disrripution: Central America, the Guianas, Brazil, Guinea,
Sierra Leone, and the Congo, usually in wet or moist thicket, road-
side, weedy fields, savanna, sometimes in dry forest, at 0-850 meters
elevation.

BRITISH HONDURAS: Schipp 1060 (MICH, §).

COSTA RICA: Boca Culebra, shore of the Pacific, Pittzer 12056 (US). Boruca,
Pittier 4573 (US). Mano de Tigre, between Boruca and Térraba, Tonduz 4592
(US). San Jos#: El General, Skutch 2881 (S, US).

PANAMA: Paitilla, Bro. Heriberto 235 (US). Coci&: Aguadulee, Pittier 4833
(US). CanaL ZonE: Gamboa, Standley 28364 (US). Corozal, Standley 27351
(S, US). Summit, Standley 26925 (US). Las Cascadas Plantation, near Summit,
Standley 29629 (US), 30131 (US). Balboa, Standley 25520 (US). Hills west of
the Canal, near Gattin, Standley 27271 (US). Cerro Gordo, near Culebra,
Standley 26028 (US). Coi6n: Between France Field, Canal Zone, and Catival,
Standley 30384 (US). PanamA: Vicinity of Juan Franco Race Track, near
Panamd, Standley 27756 (US). Corozal Road, near Panama, Standley 26863
(US). North of Panamd, Bro. Paul 586 (US). Rio Tecumen, Standley 29435
(S, US). Near the big swamp east of the Rio Tecumen, Standley 26496 (US).
Near Matias Herndndez, Standley 28992 (US). Nuevo San Francisco, Standley
30756 (US). Laguna de Portala, near Chepo, Pittier 4619 (US).

FRENCH GUIANA: Leprieur 122 (US, syntype). fles du Salut, Mana, Sagot
53 (P, 8, W), 54 (W). Cayenne, Aublet s.n. (W).

SURINAM: Focke 31 (LL). Hostmann 1575 (W). Tulleken 242 (L). Near
Paramaribo, Kappler 1575 (W). Fluv. Cappename Sup., Boon 1115 (US).
Para, Splitgerber 1158 (L).

BRITISH GUIANA: Roraima, Schomburgk 265 (W). Mazaruni Station,
Sandwith 1084 (8).

BRAZIL: Burchell 9027 (KK). Amazonas: Manaus, Schwacke 366 Coll. III
(R, US). Par&: da Costa 67A (R, US). Near Pard, Burchell 9566 (GH, K, L).
Between S. Jofo de Araguaya and 8. Anna, Burchell 9234 (I<). Vicinity of San-
tarem, Spruce May 1850 (RB, W). Marto Grosso: Cuiabd, Malme s.n. (8).

GUINEA: Near Mamou, Sweeney 4 (US).

SIERRA LEONE: Thomas 6117 (US). Freetown, W. H. & A. H. Brown 59
(US).

THE CONGO: Wompsatt: Casier St. Jean Mission, Vanderyst 1269 (8).

GOLDBERG—GENUS MELOCHIA 295

This species is close to Melochia manducata Wright, but the inflo-
rescences are not pedunculate, the fruit and leaves are smaller, and
the fruit is not covered with glandular hairs. It is also related to
M. villosa (Mill.) Fawe. & Rendle but differs in the position of the
inflorescences and size of the fruit, and its flowers are not dimorphic.

It is apparently indigenous to America and Africa.

It is illustrated by Schumann (47).

34. Melochia arenosa Benth. 7n Hook. Journ. Bot. 4:127. 1841.

Melochia cinerea St.-Hil. & Naud. Ann. Sci. Nat. IT, 18:35. 1842.

Riedleia littoralis Poepp. & Endl. Nov. Gen. & Sp. Pl. 3:73. 1845.

Melochia littoralis (Poepp. & Endl.) Schum. in Mart. Fl. Bras. 12(3):44.
1886.

Melochia corumbensis 8. Moore, Trans. Linn. Soc. II, Bot. 4:317. 1895.

?Melochia cinerea St.-Hil. & Naud. var. discolor Hassl. Addenda Pl. Hassl.
12. 1917 (without description).

Melochia llanorum Cuatr. Rev. Acad. Colomb. 8:487. 1952.

A low shrub, sometimes herb, 0.3-2 meters high, the young stems
and petioles densely cineraceous-tomentulose, hairs stellate, 0.1 mm.
long, also a very few glandular hairs, 0.1-0.2 mm. long, internodes
1-4.3 cm. long; stipules deciduous, 2.6-9.5 mm. long, 0.3-1.0 mm.
wide, usually linear-acute, densely tomentulose, hairs stellate, 0.1
mm. long, a few simple hairs, 0.2-1.0 mm. long; petiole 0.2-2.9 cm.
long.

Lamina 0.9-7.5 cm. long, 0.6-5.5 cm. wide, ovate, oval, oblong or
lanceolate, the base rounded to subcordate, the apex rounded to
acute, the upper surface green to canescent, the lower cinereous,
both densely tomentulose, velvety to the touch, hairs stellate, with
8-12 rays, 0.1 mm. long, sometimes a few simple hairs 0.2-0.8 mm.
long, the margin irregularly crenate-serrate, serrations 0.7-3.8 mm.
wide, 0.2-2.5 mm. high, 5-10 pairs of primary veins, 2 are basal.

Inflorescences in axils of upper 3-9 (—25) leaves, 1.3-2.5 em. long,
the ultimate divisions umbellate, 3-6 flowers per umbel, the peduncle
0.3-1.5 cm. long, 1-3 per axil, the pedicel 2-5 mm. long, to 8 mm.
long in the fruiting stage; bracts at the base of the pedicels, 1.8-4.3
(-8.0) mm. long, 0.2-1.1 mm. wide, usually linear, densely tomen-
tulose, hairs stellate, occasionally glandular, 0.1 mm. long.

Calyx 3.1-5.5 mm. long, 3.2-5.4 mm. wide, not accrescent, densely
tomentulose, hairs stellate, a few glandular, 0.1-0.2 mm. long, the
teeth 1.0-2.7 mm. long, 0.5-1.7 mm. wide at the base, nearly linear
distally, broadening at the base, the sinus rounded, the teeth distant;
petals rich pink, red, rose violet, lilac, lilac or violet with yellow
proximally, 6-10 mm. long, 1.8-4.0 mm. wide, obovate, oblanceolate,
or oblong, the claw 0.8-1.5 mm. long, the base free from the staminal
tube or attached to it for 0.5-1.5 mm.

296 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Longistylous form. Stamens 3.4-4.7 mm. long, the filaments
united up to the anthers, sometimes a few stellate hairs 0.1 mm.
long near the base of the staminal tube, the anther 1.0-1.3 mm. long,
0.5-0.7 mm. wide, oblong, emarginate at both ends, nearly to the
middle from the apex; pistil 4.8-7.0 mm. long, the styles filiform,
free for 2.7-4.0 mm., to the ovary or united for 0.6-1.0 mm., papillose
at the apex for about 1.2-2.0 mm., the ovary ellipsoid to globular-
ovoid, the ovary and base of styles tomentulose-hirsute, hairs 0.1-
1.0 mm. long, stipe none.

Brevistylous form. Stamens 5.6-7.7 mm. long, the filaments free
for 2.6—-4.3 mm., sometimes a few hairs at the base of the tube, some-
times staminodes present, 0.2-0.3 mm. long, the anther 1.1-1.7 mm.
long, 0.5-0.8 mm. wide, oblong, slightly emarginate at both ends;
pistil 4.8-6.2 mm. long, the styles filiform, free for 1.7-3.8 mm.,
united for 0.6-0.7 mm., the ovary ovoid, the ovary and base of styles
tomentulose, hairs 0.1-1.7 mm. long.

Fruit 3.2-3.5 mm. long, 3.5-4.5 mm. wide, obtusely pentagonal,
elobose, the sulci between the carpels shallow, extending a fourth to
a fifth of way to the center of the fruit, hirsute-tomentulose, hairs
simple, forked, and stellate, a very few glandular, 0.1-1.0 mm. long,
stipe none; dehiscence loculicidal along entire dorsal and ventral
sutures, later septicidal, the fruit falling apart; seeds 2.0-2.3 mm.
long, 1.2-1.5 mm. wide, trigonal, dark brown to black, 1 per cell.

Flowering specimens collected from April to July and September
to January; fruiting ones, from October to December, April, and
June.

TypE LOCALITY: Syntypes collected by Schomburgk, nos. 31 and
519, the latter cited below, on sandy banks of the Essequibo and
Rupununi Rivers, and by Gardner, no. 2489, in Piauf, Brazil, also
cited below.

DisTRIBUTION: Cuba and Venezuela to Paraguay, often in places
that are sometimes inundated and moist savanna and woods at
130-?1,300 meters elevation.

CUBA: Maranzas: N. of Matanzas, Hoyo Colorado, Bro. Alain 3951 (US).

BRITISH GUIANA: Essequibo and Rupununi Rivers, Schomburgk 519 (L,
syntype). Rupununt: Lethem, Jrwin 567 (US). Rupununi Savanna, Vesey-
Fitzgerald 745 in part (US). BASIN OF THE RUPUNUNI RIVER: Karenampbo, lat.
3°45’ N., A. C. Smith 2246 (A, K, P, 8, US).

VENEZUELA: GvuArico: Palo Seco, between La Encrucijada and the Mision
de Arriba, Calabozo, Pittier 14960 (US).

COLOMBIA: Castafial, Sonntag 18 (GH, also US neg. 5612). Aravuca:
Llanos de Casanare, Esmeralda, Cuatrecasas 3934 (US, isotype of Melochia
llanorum, Cuatr.).

BRAZIL: Regnell II 631 (P). Western part, Tamberlik s.n. (W). AMAZONAS:
Amazon Rio, F neg. 9605 ex B, probably an isotype of Melochia littoralis (Poepp. &
Endl.) Schum. Rro Branco: Sao Marcos, Ule 7869 (L). Pard: Boa Vista on

GOLDBERG—GENUS MELOCHIA 297

the Tapajés River, Dahlgren & Sella 65 (US). Jutahy de Almeirim, Ducke 18959
(RB). Medio Cumind, de Sampaio 5921 (R, US). Alto Cumina, de Sampaio
5696 (R, US). Rio Erepecurt, de Sampaio 5372 (R, US). Prauf: Gardner 2489
(US, syntype, also F neg. 9597 ex B in US). Minas Gerais: Claussen 95 (P,
isotype of Melochia cinerea St.-Hil. & Naud., also US neg. 5667). Maro Grosso:
Corumbd4, F neg. 9601 ex B of Moore 905, isotype of Melochia corumbensis Moore
(US); Malme s.n. (S).

PARAGUAY: Alto-Paraguay, Chaco, 21° S. lat., Piebrig 1836 (K, 8).

This species can be distinguished from other members of the section
by the abundant, very short, stellate hairs with numerous rays (8-12),
0.1 mm. long, on the leaves and by its axillary pedunculate
inflorescences.

In addition to the localities cited above, Schumann (47) reported
M, arenosa Benth. from Goids, M. littoralis (Poepp. & Endl.) Schum.
from the shores of Lake Ega, Amazonas, Brazil, and M. cinerea
St.-Hil. & Naud. from Bahia, between Victoria and Bahia, and from
upper Surinam. I consider the last two species to be synonyms of
M. arenosa Benth.

I have not selected a lectotype because I have not seen the syntypes
located at Kew.

35. Melochia simplex St.-Hil. Fl. Bras. Mer. 1:164. 1825.
Melochia oblonga Benth. in Hook. Journ. Bot. 4:128. 1841.
Melochia sorocabensis Schum. in Mart. Fl. Bras. 12(3):42, pl. 8. 1886.
Melochia stricta Schum. in Mart. Fl. Bras. 12(3):42. 1886.
Melochia Missionum Ekman, Arkiv. Bot. Stockh. 9(4):47, fig. 10. 1910.
Melochia stricta Schum. f. Missionum (Ekman) Hassl. Repert. Sp. Nov. Fedde
8:123. 1910.

An erect or subdecumbent-ascending herb to suffrutex 0.4-1.5 me-
ters high, simple or 2- to 3-branched from near the base, blooming
during the first year, the young stems and petioles tomentulose, hirsute
or subglabrous, hairs stellate, sometimes also forked, simple, and glan-
dular, 0.1-0.2 (-0.5) mm. long, internodes 0.8-6.9 cm. long; stipules
caducous, 2.1-8.0 mm. long, 0.3-1.7 mm. wide, lanceolate to nearly
linear, ciliate and sometimes tomentulose, hairs simple, stellate, or
forked, 0.1-0.5 mm. long; petiole short, 0.2-1.0 em. long.

Lamina 1.5-9.5 cm. long, 0.5-3.0 cm. wide, lanceolate to suboblong,
nearly linear near the apex of the stem, the base narrowly rounded to
slightly cuneate, the apex acute, acuminate, or slightly rounded, the
upper surface with few scattered stellate, forked, simple, and rarely
glandular hairs, 0.1-0.7 mm. long, the lower surface with scattered
hairs or slightly tomentulose, especially on the veins, hairs stellate, a
few forked and simple, most 0.1 mm., a few to 1 mm. long, the margin
irregularly crenate-serrate, serrations 0.3-6.0 mm. wide, 0.2-1.5 mm.
high, 5-12 pairs of primary veins, 1 or 2 are basal.

Inflorescences terminal, extending 1-18 cm. beyond the first leaf,
spicate- to paniculate-cymose, the ultimate divisions subcapitate um-

298 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

bels of 2-5 flowers, sometimes also or only axillary, 0.8-7.5 cm. long,
in the axils of 1-9 leaves, the peduncle 0.2—4.1 cm. long, the pedicel
0.3-3.6 mm. long; bracts 1-5.5 mm. long, 0.1-1.5 mm. wide, deltoid-
acuminate, lanceolate, or linear, 2 or 3 immediately subtending the
flower or on the pedicel, sometimes in pairs at the base of the pedicel
or peduncle, ciliate, a few forked and stellate hairs 0.1-0.5 mm. long
on the abaxial surface.

Calyx pale yellowish green, sometimes reddish purple, 2.4-4.5 mm.
long, 2.5-5.0 mm. wide, tomentulose or with scattered stellate and
sometimes also forked, simple, and glandular hairs 0.1-0.2 (-0.5) mm.
long, the teeth 0.7-1.9 mm. long, 0.6-2.2 mm. wide, deltoid at the
base to linear at the apex, distant, the sinus rounded to truncate;
petals violet, purple, rose, blue, purple distally and yellow proximally,
or orange, 5.6-10.2 mm. long, 1.3-4.7 mm. wide, oblanceolate- to
obovate-cuneate, free from the staminal tube or attached to it for 0.3-
2.0 mm., the claw 0.5-1.5 mm. long.

Longistylous form. Stamens 3.4-4.6 mm. long, the filaments
united up to the anthers, the anther 0.9-1.6 mm. long, 0.4-0.7 mm.
wide, oblong, emarginate at both ends, nearly to the middle from the
apex; pistil 4—-5.8 mm. long, the styles filiform, free for 1.6-4.3 mm.,
united for 0.5-2.8 mm., papillose and subclavate for 0.7-2.0 mm. at
the apex, the ovary globular to ovoid, sericeous or hirsute, especially
at the apex, hairs simple, 0.2-0.8 mm. long, the stipe 0-0.5 mm. long.

Brevistylous form. Stamens 4.5-6.6 mm. long, the filaments free
for 1.7-3.4 mm., the anther 0.9-1.4 mm. long, 0.4-0.7 mm. wide, sim-
ilar in shape to that above; pistil 3.4-3.6 mm. long, the styles filiform,
free for 1.2-1.5 mm., united for 0.3-0.9 mm., the ovary globular to
ovoid, sericeous or hirsute, hairs simple, 0.1—-1.0 mm. long.

Fruit 3.4-3.7 mm. long, 3.9-4.0 mm. wide, globular, obtusely
pentagonal, the sulci between the carpels shallow, extending a fifth
to a fourth of way to the center of the fruit, hirsute, hairs simple and
sometimes also forked, stellate, and glandular, 0.1-1.1 mm. long,
stipe none; dehiscence loculicidal all along the dorsal and ventral
sutures, also septicidal, the fruit falling apart; seeds 2.2-2.5 mm. long,
1.1-1.8 mm. wide, trigonal, dark brown, 1 per cell.

Flowering specimens collected from December to April, and in
June, August, and October; fruiting ones, from December to March,
and in August, and October.

Type Locauiry: The type was collected by St.-Hilaire near the
city Guaratingueta, in the northern part of the province of Sao
Paulo, Brazil. The isotype is cited below.

Distripution: British Guiana to southern Brazil, Bolivia, Para-
guay, and northern Argentina in humid campo, marshes that are often

GOLDBERG—GENUS MELOCHIA 299

inundated, among bushes at the edge of a swamp and in shrubby
fields, at 160-500 meters elevation.

BRITISH GUIANA: Savannas of the upper Rupununi, Schomburgk s.n. (K,
type of Melochia oblonga Benth., also US neg. 5419).

BOLIVIA: Santa Cruz: sara: Buenavista, Sternbach 5483 (GH), 14821 (8S).

BRAZIL: F neg. 9608 ex B of Pohl 421 (US, presumably the type of Melochia
stricta Schum.). AMAZONAS: Serra Tayana, Luetzelburg 20898 (R). GorAs:
Brejo do Riacho Fundo, Glaziou 20735 (P). Goids beyond Jaragua, Meia Ponte
to Cavalcante, Burchell 7821 (P). Gor4s?: Corgo-Corallinho, Pohl 1834 (W).
Mato Grosso: Coxim, Hoehne 2990 (R). corumBA: Fazenda Aguassuzinho,
Pereira et al. 401 (US). Fazenda MarilAndia, Pereira et al. 209 (US). Rio pE
JANEIRO OR SAo Pauvto: Coralfalza, Pohl 314 (W). So Pauto: Sellow 5288
in part (R). Near the city Guaratingueta in the northern part of the province,
St.-Hilaire Cat. No. 756d (P, isotype); F neg. 35385 of the type ex P (US).
Mooga, Brade 5973 (8). Near Sorocaba, Riedel & Luschnath 1994 (LE, isotype
of M. sorocabensis Schum., also US neg. 5271); Riedel 1994 dupl. 19 (P). Atibaia,
George & Liene Hiten 2404 (US). Parana: Féz do Iguagu, Hatschbach 9726
(US). Rio GranvE po Sut: § Leopoldo near P. Alegre, Theissen (PACA 7554).

PARAGUAY: Border of Rio Tapiracuai at Primavera, Woolston 1072 (US).
Est. Tagatiya, Anisits 2648 (S). Between Rio Apa and Rio Aquidaban, Fiebrig
4780=5207 (GH). Road to ‘Yerbales’” montium ‘Sierra de Maracayu,”’
Jejuigauzu River, Hassler 5700 (GH).

ARGENTINA: CorrIENTES: SANTO TOME: Pueblo, [barrolas.n. (8S). MIsto-
NES: Posadas, Vattuone & Bianchi 71 (US). Banks of the Alto-Parand River,
Ekman 186 (S, type of Melochia Missionum Ekman, also US neg. 5614).

This species is related to Melochia graminifolia St.-Hil., but the
leaves of the latter are linear, rather than lanceolate or oblong, with
obscure and considerably fewer serrations.

36. Melochia graminifolia St.-Hil. Fl. Bras. Mer. 1:160, pl. 31. 1825. Pare 6
Visenia gramintfolia (St.-Hil.) Spreng. Syst. 4:Cur. Post. 248. 1827.
Riedleia graminifolia (St.-Hil.) Steud. Nom. II, 2:460. 1841.

An erect perennial herb or suffrutex with 1-3 slender, branched
shoots from the base, 0.3-1.2 meters high, 2-4 mm. wide, the root
woody, thick, up to 2 cm. wide, the young stems and petioles with
scattered stellate hairs about 0.1 mm. long, internodes 0.5-4.7 cm.
long; stipules caducous, 2-4.5 mm. long, 0.2—0.4 mm. wide, deltoid-
acuminate to acicular, with few hairs about 0.1 mm. long; petiole very
short, 0.1-0.3 cm. long.

Lamina 2-9.7 cm. long, 0.2-0.8 cm. wide, nearly linear, the base
narrowed, rounded to cuneate, the apex acute, both surfaces green
and glabrous, rarely a few hairs on the veins beneath, the margin
entire at the base, irregularly and obscurely serrate at the apex,
serrations 2-12 mm. wide, 0.2-0.3 mm. high, 4-8 pairs of primary
veins, 1 is basal.

Inflorescence terminal, long, slender, branched, racemiform or
paniculate, extending 6-31 cm. beyond the first leaf, the flowers soli-

300 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

tary or in umbels of 2 or 3, sometimes also in the axils of the upper 1-3
leaves, 2.5-19 cm. long, the pedicels 2-9 mm. long; bracts 0.5-1.5
mm. long, 0.2-0.4 mm. wide, deltoid-acuminate to nearly linear, with
few hairs about 0.1 mm. long.

Calyx 1.8-3.0 mm. long, 2.7-4.1 mm. wide, stellate and glandular
puberulous, hairs 0.1-0.3 mm. long, the teeth 0.6-1.1 mm. long, 0.4-
0.5 mm. wide, deltoid-acute or deltoid-acuminate to nearly linear,
the sinus very obtuse, the teeth distant; petals pale lilac, blue, bluish
pink, rose, yellowish and red striped at the base, purple distally and
yellow with purple veins proximally, violet, yellow, or white, 6.5-10.5
mm. long, 3.0-5.2 mm. wide, obovate-cuneate, the claw 0-0.5 mm.
long, the base of petals attached to the staminal tube.

Longistylous form. Stamens 2.6-4.0 mm. long, the filaments
united up to the anthers, the anther 1.1-1.2 mm. long, 0.5-0.7 mm.
wide, oblong, emarginate at both ends, almost to the middle from the
apex, sacs somewhat divergent at the apex; pistil 3.6-4.7 mm. long,
the styles free for 1.5-3.1 mm., papillate at the apex for 1.5-2.3 mm.,
the ovary ellipsoid to ovoid, glabrous or with few hairs 0.1 mm. long,
the stipe 0.3-0.8 mm. long.

Brevistylous form. Stamens 5.8 mm. long, the filaments free for
3.0 mm., the anther 1.3-1.4 mm. long, 0.6 mm. wide; pistil 3 mm.
long, the styles free for 1.2 mm., the stipe 1 mm. long.

Fruit obtusely pentagonal, depressed-globose, very shallowly sulcate
between the carpels, 3.3-4.0 mm. long, 4.2-4.8 mm. wide, tometulose,
hairs simple and forked, 0.1—0.2 mm. long, the stipe 0.4-0.8 mm. long;
dehiscence loculicidal along the dorsal suture and later septicidal;
seeds 2.2-2.5 mm. long, 1.4-1.6 mm. wide, trigonal, black.

Flowering specimens collected from September to July; fruiting
ones, from October to January, and March to May.

Typr Locauiry: Marshes near the village of S. Domingos, Minas
Gerais, Brazil. Type collected by St.-Hilaire, isotype cited below.

Disrrrpution: Eastern Colombia to Paraguay, and southern
Venezuela and British Guiana to Sao Paulo, Brazil, usually in wet
argilaceous savannas, sometimes dry grassy Ilanos, and campo, at
70-500 meters elevation.

BRITISH GUIANA: On Rio Branco, R. Schomburgk 805 (LL, US).

VENEZUELA: Botfivar: La Paragua, L. Williams 12582 (US). Rio Paragua,

Potrerito, Cardona, 696 (US). Rio Paragua, near Chorrera de Aguacanta,
Killip 37552 (US).
COLOMBIA: Mera: Rio Meta, La Poyata, 65 km. E. of Villavicencio,

Cuatrecasas 4473 (US); Haught 2518 (US). Rio Guaviare, San José del Guaviare,
Cuatrecasas 7720 (US). San Juan de Arama, Idrobo & Schultes 699 (US).

BOLIVIA: Velasco, Kuntze (US).
BRAZIL: Rio Branco: Estrada Boa Vista-Caracaraf, Black 51-13509 (US).
Fazenda Bom Intento, Black 51-14047 (US). Minas Gerais: 8. Domingos,

GOLDBERG—GENUS MELOCHIA 301

St.-Hilaire 1589 (P, isotype, also US neg. 5664). Gordks: Gardner 3604 (US).
Mato Grosso: Caceres, Hoehne 1119 (R), 604 (R). Cuiaba, Malme Regnell II
3271 (8). Porto Murtinho, Malme s.n. (8S). Rio pp JANEIRO: Serra da Estrella,
Mandioca, Glaziow 12459 a (R). Sdo Pauto: Bank of Rio Grande at Melaniros
(?), Regnell III 279 (S).

PARAGUAY: Between Rio Apa and Rio Aquidaban, Centurion, Fiebrig
4123=4029 (L, US). Rio Apa, Hassler 7683 (8). In Quebrachal Beco. Branco,
Anisits 2257 (S).

This species is distinguished from all other species in the genus by
its nearly linear, obscurely serrate leaves. It appears to be most

closely related to Melochia simplex St.-Hil., which has broader leaves
and distinct serrations.

37. Melochia manducata Wright in Sauvalle, Anal. Acad. Ci. Habana 5:241.
1868.
Melochia glandulifera Standl. Contr. U.S. Nat. Herb. 23:803. 1923.

An erect, annual, simple or branched herb to suffrutex, the stem
hollow, 0.3-0.65 meter high, the primary root not as thick as the base
of the stem, secondary roots abundant, the young stems glabrous or
with a few glandular, stellate, forked, or simple curved hairs 0.1-0.5
mm. long in a narrow longitudinal line, internodes 1.5-6.2 cm. long;
stipules 3-7.4 mm. long, 0.7-1.3 mm. wide, lanceolate, ciliate, hairs
simple, 0.1-0.7 mm. long; petiole 0.2-5.0 cm. long, tomentulose in a
longitudinal line on the upper surface, hairs stellate, forked, a few
glandular, 0.1-0.5 mm. long.

Lamina 3-7.5 cm. long, 0.9-6.2 cm. wide, lanceolate to broadly
ovate, the base rounded, truncate or subcordate, the apex acute,
subglabrous, both surfaces with a few simple appressed hairs 0.1-0.5
mm. long, mostly on the veins, the margin irregularly crenate-
serrate, serrations 0.7-3.1 mm. wide, 0.3-1.6 mm. high, 9-12 pairs of
primary veins, 2 are basal.

Inflorescences axillary, 0.7-2.8 cm. long, in the axils of the upper
6-10 leaves and sometimes also terminal, extending 1-1.5 cm. beyond
the first leaf, umbellate and capitate as well as pedunculate and non-
pedunculate in the same axils, 2 or 3 flowers per peduncle, 5-8 flowers
per axil, the peduncle 0-1.6 cm. long, the pedicels 0-14.0 mm. long;
bracts 2 or 3 immediately subtending the flower, 1.5-5.0 mm. long,
0.3-0.9 mm. wide, lanceolate, ciliate, hairs simple, 0.1-0.5 mm. long.

Calyx membranous, 1.5-2.4 mm. long, 2.3-3.2 mm. wide, with
scattered simple and sometimes also a few glandular hairs 0.1-0.2
mm. long, the teeth distant, 0.5-0.8 mm. long, 0.2-0.6 mm. wide at
the base, nearly linear toward the apex, the sinus rounded to truncate;
petals rose, reddish lilac, or purple, 3.8-4.7 mm. long, 1.3-1.7 mm.
wide, obovate to obovate-oblong, free from the staminal tube or
attached to it for 0.5 mm., the claw 0.2-0.4 mm. long.

227-051—67 8

302 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Androecium and gynoecium nearly equal in length. Stamens 2.1-
2.4 mm. long, the filaments united up to the anthers, the anther
0.5-0.8 mm. long, 0.3-0.5 mm. wide, oblong, emarginate at both
ends, nearly to the middle from the apex; pistil 2.2-2.5 mm. long, the
styles filiform, free for 0.8-1.2 mm., united for 0.3 mm., the ovary
globular-ovoid, hirsute-tomentulose, hairs simple, 0.1-1.0 mm. long,
stipe none.

Fruit 2.7-4.0 mm. long, 3.8-4.8 mm. wide, globular, pentagonal, the
sulci between the carpels very shallow, extending a fourth to a fifth of
way to the center of the fruit, covered with glandular and simple
hairs 0.1—-0.7 mm. long, the stipe 0-0.2 mm. long; dehiscence loculicidal
along the dorsal and ventral sutures, later septicidal, the fruit falling
apart; seeds 1.7-2.3 mm. long, 1.0-1.4 mm. wide, blackish brown, 1
per cell.

Flowering and fruiting specimens collected from October to March,
and in August.

Type nocauity: In palm forest, Ilanos, and moist places around
Huaiquiba, San Cristébal, Cuba, probable isotypes cited below. I
select Wright 3510 (US) as lectotype.

DistriBuTION: Mexico, Guatemala, Panama, Cuba, Dominican
Republic, Colombia, Ecuador, and northern Brazil, in swamp
shrubbery, moist places on the road, marsh, llanos, in sandy soil, at
0-100 meters elevation.

MEXICO: Guerrero: Acapuleo and vicinity, E. Palmer 107 in 1984-95
(US). Cutapas: Tonali, Purpus 6925 (UC; US, type of Melochia glandulifera
Standl.).

GUATEMALA: Retaluleu, Bernoulli & Cario 3114 (K).

PANAMA: Panama: Near the big swamp E. of Rio Tecumen, Standley
26655 (US).

CUBA: C. Wright 3510 (GH, 8, US, probably isotypes).

DOMINICAN REPUBLIC: Pactricapor: Pimentel, Abbott 653 (US).
Santo Domingo, Duarte, Matanzas at Rio Boba, Ekman 15866 (S).

COLOMBIA: MagpaLtena: Santa Marta, H. H. Smith 724 (8, US).
SANTANDER: Along Rio Magdalena near Barranca Bermeja, Molina & Barkley
185205 (US). Puerto Wilches and vicinity, Killip & Smith 14790 (US).

ECUADOR: Los Rios: Hacienda Clementina on Rio Pita, Asplund 5573
(R, US).

BRAZIL: Amazonas: Manaus, Rio Negro, Ule 5393 (K, L). Parf: Below
Obidos, Kuhlmann 83 (RB).

This species is close to M. corchorifolia L., but differs in that its
inflorescences are principally axillary rather than terminal, and are
often evidently pedunculate, those of M. corchorifolia L. are usually
not pedunculate.

38. Melochia lanceolata Benth. in Hook. Journ. Bot. 4:128. 1841.
Melochia betsiliensis Baker, Journ. Linn. Soc. 22:451. 1887.

GOLDBERG—GENUS MELOCHIA 303

An erect, simple herb, the stem hollow, or branched shrub, the
young stems with a narrow longitudinal line of stellate hairs 0.1-0.4
mm. long, glabrous beyond the third internode, internodes (0.7-)
1.5-6.0 cm. long; stipules 1.5-2.2 (-5.0) mm. long, 0.4-0.5 (-1.5) mm.
wide, nearly linear or lanceolate, ciliate and with few scattered, simple
hairs 0.1-0.2 mm. long; petiole (0.2—-) 0.5-3.2 cm. long, slender, with
few simple, forked, and stellate hairs 0.2-0.5 mm. long.

Lamina (1.3—) 4.1-10.0 cm. long, 0.6-3.5 em. wide, lanceolate, the
base rounded, the apex acuminate or acute, sometimes obscurely
lobed as in M. corchorifolia L., subglabrous with very few scattered
simple hairs 0.1-0.3 mm. long on the midrib of the upper surface and
veins of the lower surface, the margin irregularly serrate, serrations
1-2.7 (-4) mm. wide, 0.3-0.9 mm. high, 9-13 pairs of primary veins,
1 or 2 are basal.

Inflorescence 1-4 cm. long, up to 11 cm. in the fruiting stage,
terminal and in the axil of the first leaf, coarctate, corymbiform, the
ultimate divisions umbellate, 2 or 3 flowers per umbel, the peduncle
0.5-1.0 em. long, 2 cm. according to Schumann (47), the pedicel
(0.3-) 1.5-4.7 mm. long, with glandular and simple hairs 0.1-0.3 mm.
long; bracts usually at the base of the pedicel, 0.5-3.5 mm. long,
0.1-1.0 mm. wide, lanceolate to linear, ciliate with few simple, forked
and sometimes glandular hairs 0.1-0.3 mm. long.

Calyx 1.5-2.0 mm. long, 2.0-3.5 mm. wide, with very few simple
hairs 0.1 mm. long and glandular ones 0.3-0.6 mm. long, the teeth
0.4-0.5 mm. long, 0.2 mm. wide, linear, distant, the sinus truncate;
petals flavescent or lilac, (3.7—) 5.6-6.5 mm. long, (1—) 3-4 mm. wide,
obovate-cuneate, attached to the staminal tube for 0.6 mm., the claw
0.2-0.3 mm. long.

Androecium and gynoecium nearly equal in length. Stamens
2.2-2.6 mm. long, the filaments free for 0.2-0.4 mm., the anther 0.6
mm. long, 0.4 mm. wide, oblong, emarginate at botk ends, nearly
to the middle from the apex; pistil 2.2-2.7 mm. long, the styles
filiform, 0.8-1.3 mm. long and free to the ovary, or united for 0.1-0.2
(-0.7) mm., the ovary globular, with few glandular and simple hairs
0.2-0.4 mm. long, stipe none.

Fruit 3.5-4.0 mm. long, 4.0-4.1 mm. wide, globose, very slightly
pentagonal, the sulci between the carpels very shallow, extending
one-fifth of way to the center of the fruit, with scattered simple and
glandular hairs 0.3-0.5 mm. long, the stipe 0.5 mm. long; dehiscence
loculicidal along the dorsal and ventral sutures, the fruit falling apart;
seeds not quite ripe, 2 mm. long, 1.1 mm. wide, brown, 1 per cell.

Flowering and fruiting specimens collected in May.

TYPE Locatity: The type was collected by Schomburgk, no. 362,
in British Guiana; an isotype is cited below.

304 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Disrrisution: British Guiana and Surinam, in shallow water, in
shade.

SURIMAN: Hostmann s.n. (L), 449 (W), 457 a (S). Hostmann & Kappler
46 (8), 657 BS). Schimper 657 (W). Para, Splitgerber 1001 (L).

BRITISH GUIANA: Schomburgk 362 (L, isotype).

MADAGASCAR: Baron 4757 (P), 4678 (P), syntypes of M.. betszliensis
Baker.

This species is close to M. corchorifolia L., but its inflorescence is
corymbiform rather than subcapitate. The leaves of AZ. corchorifolia
are most often ovate, those of AZ. lanceolata are lanceolate. Both
species are characterized by the nearly equal length of androecium
and gynoecium.

39. Melochia corchorifolia L. Sp. Pl. 675. 1753.

Melochia concatenata L. Sp. Pl. 675. 1753; Hook. Fl. Brit. Ind. 1:374.
1874, in synonymy.

Melochia supina L. Sp. Pl. 675. 1753; Hook. Fl. Brit. Ind. 1:374. 1874,
in synonymy.

Corchorus javanicus Burm. f. Fl. Ind. 123. 1768.

Melochia cordata Burm. f. Fl. Ind, 143. 1768.

Melochia erecta Burm. f. Fl. Ind. 143. 1768.

Mougeotia corchorifolia (L.) H.B.K. Nov. Gen. & Sp. 5:326, adnot. 1823.

Riedleia concatenata (L.) DC. Prodr. 1:492. 1824.

Riedleia corchorifolia (L.) DC. Prodr. 1:491. 1824.

Riedleia supina (L.) DC. Prodr. 1:491. 1824.

Riedleia radiata Blume, Bijdr. 86. 1825.

Visenia concatenata (L.) Spreng. Syst. 3:31. 1826.

Visenia corchorifolia (L.) Spreng. Syst. 3:30. 1826.

Visenia supina (L.) Spreng. Syst. 3:31. 1826.

Melochia affinis Wall. Cat. no. 1198. 1829.

Melochia pauciflora Wall. Cat. no. 1199. 1829.

Polychlaena ramosa G. Don, Gen. Syst. 1:488. 1831.

Polychlaena simplex G. Don, Gen. Syst. 1:488. 1831.

?Geruma subtriloba Blanco, Fl. Filip. 182. 1887.

Lochemia corchorifolia Arn. Ann. Sc. Nat. sér. 2. 11:172. 1839.

Hibiscus donii Walp. Rep. Bot. Syst. 1:304. 1842,

Hibiscus endlichert Walp. Rep. Bot. Syst. 1:304. 1842.

Melochia Burmanni Zoll. & Mor. Syst. Verz. Zoll. 27. 1846.

Melochia longebracteolata J. Ar. in Mém. Inst. Sci. Madag., sér. B. 7:66.
1956.

An annual herb or perennial subshrub, sometimes gregarious,
simple or branched, erect or decumbent, a few centimeters to 2.5
meters long, the taproot not thicker than the base of the stem, the
young stems sometimes hollow, with scattered stellate hairs some-
times also forked and simple, 0.1-0.7 mm. long, and also sometimes
hairs in longitudinal lines about 0.5 mm. wide, internodes 2-10 em.
long; stipules 3-9 mm. long, 0.5-1.5 mm. wide, deltoid-acuminate to
lanceolate-ovate, mostly ciliate, hairs mostly 0.3-1 mm. long; petiole
0.3-2.0 (-6.0) em. long.

GOLDBERG—GENUS MELOCHIA 305

Lamina green, sometimes purplish, (1.0-) 2-6 (-9.8) em. long,
(0.5-) 1-2.6 (-7.0) em. wide, ovate to lanceolate, the base usually
rounded, sometimes truncate, the apex acute, both surfaces sub-
glabrous, usually a few simple hairs 0.1-0.8 mm. long on the midrib
of the upper surface and on the midrib and veins of the lower surface,
the margin irregularly crenate-serrate, serrations 1-4.2 mm. wide,
0.5-1.0 mm. high, 6-10 pairs of primary veins, 2 are basal,

Inflorescence terminal, 1-2 cm. long, 0.5-3.5 cm. wide, subcapitate,
rarely radiately grown out to 3 cm., consisting of about 5-25 often
fragrant flowers, also sometimes at apices of branches, and rarely
1 or 2 flowers in the axils of the upper 3-5 leaves, the pedicels 0-3
mm. long; bracts 3.9-7.5 mm. long, 0.3-1.9 mm. wide, 3 or 4 immedi-
ately subtending the flower and usually 2 more below, ovate-lanceolate
to nearly linear, mostly ciliate, hairs 0.2-1.0 mm. long,

Calyx pale green, 2.0-2.7 mm. long, 2.4-3.3 mm. wide, hirsute,
hairs simple, 0.2-0.8 mm. long, the teeth 0.4-0.7 mm. long, 0.3-0.6
mm. wide, deltoid-acute, distant, the sinus obtuse; petals white to
pink, pink or white distally and yellow proximally, rarely yellow, pur-
ple or violet 4.7-6.5 mm. long, 1.2-2.5 mm. wide, obovate to oblance-
olate, almost entirely free from the staminal tube or attached to it
for 0.8-0.9 mm., the claw 0-0.5 mm. long.

Androecium and gynoecium about equal in length. Stamens
2.4-3.2 mm. long, the filaments white, yellow, or green, united up
to the anthers or free for 1 mm., the anther yellow, 0.6-1.0 mm.
long, 0.3-0.6 mm. wide, oblong, emarginate at both ends, nearly to
the middle from the apex; pistil 2.5-3.2 mm. long, the styles white
or green, 1.0—-1.5 mm. long, free for 0.9-1.5 mm., usually to the ovary,
sometimes united for 0.4-0.6 mm., papillose for most of their length,
the ovary ovoid, densely hirsute, hairs simple, 0.5-1.0 mm. long,
stipe none.

Fruit green, white, pink, black, or purplish, (3.3-) 4-5 mm. long,
(3.6-) 4.5-5.5 mm. wide, globular, pentagonal, the sulci between car-
pels very shallow, hirsute, hairs simple, 0.3-0.6 mm. long, stipe none;
dehiscence first loculicidal along the dorsal suture, then along the
ventral suture, eventually also septicidal, the fruit falling apart;
seeds 2.2-3.2 mm. long, 1.4-1.8 mm. wide, trigonal, blackish brown.

Flowering and fruiting specimens collected throughout the year
but especially in late summer and fall.

TypE Locatity: ‘“Cosmop. trop.” The type is in the Dillenius
herbarium at Oxford University. There are two specimens in the
Linnaean herbarium attributed to Melochia corchorifolia. The one
labeled by Sir James E. Smith is M. corchorifolia; the other, labeled
Melochia corchorifolia by Solander and M. nodiflora by Smith is
M. nodiflora.

306 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Distripution: Old World tropics and subtropics, introduced in
the New World. It occurs as a weed in cultivated ground, in open
pineland, meadows, and in waste ground, in wet or dry sandy or
clay soil, mostly at 0-250 (-1,000) meters elevation.

UNITED STATES: Sout CaroLINA: GEORGETOWN: Georgetown, God-
frey & Tryon, Jr. 1678 (US). BEAurorr: Bluffton, Mellichamp s.n. (US).
Grorata: Altamaha River, Curtiss 400** (MICH, US). mcinrosH: Sapelo
Island, Duncan 20477 (US); McAtee 3306 (US). FLorma: puvaL: Jackson-
ville, Curtiss 6511 (US). Lee: Vicinity of Fort Myers, J. P. Standley 385
(US). FRanxiin: Apalachicola, Chapman 4206 (US). ALABAMA: MOBILE:
Mobile, Curtiss 6512 (US). Mohr s.n. (US). Mississippi: HARRISON: Biloxi,
Tracy 4493 (MICH, US). Lovurstana: st. TAMMANY: Covington, Fairview,
Arstne 11428 (US). JEFFERSON Davis: Lake Arthur, Wise s.n. (US). TExas:
?CHAMBERS: Stowall (?Stowell), Hanson 8 (US). Harris: Houston, Thurow
14 (US). Live oax: George West, Shiller 814 (US).

PANAMA: San José Island, Pearl Archipelago, goat farm near Rio Mato
Puerco, Erlanson 189 (US).

BRAZIL: Rio pe JANErRO: Environs of Rio de Janeiro and D’Ouro Preto,
Glaziou 14518 (K). Rio GRANDE po Sut: Gravatai near P. Alegre, Capparelli
(PACA 48235).

CONGO: Kitobola, ?Flamigri 135 (8). Hala, Corbister 1053 (S)?. Wangota,
around Coquilhatville, Verbrun 921 (S). Dolo, Schlechter 12498 (L). Lutéte,
Fr. Hens A. 280 (L). Yangambi, Louzs 10626 (US).

KENYA: Kwate: Kivumoni, near Shimoni, Drummond & Hemsley 3928 (5).

TANGANYIKA: Dar-es-Salam, Stuhlmann 7582 (US). Lutamba See, 40
km. W. of Lindi, Schlieben 5372 (8).

ZANZIBAR: Hildebrandt 1132 (L).

NYASSALAND: Buchanan 73 (US). Nyassa highland, Station Kyimbila,
Stolz 2026 (L, S).

MADAGASCAR: D’Alleizette 818 in part (L). Nossi-Bé, Ambanja, Mahilaka,
Waterlot 248 (P) type of M. longebracteolata J. Ar.

INDIA: Wight 240 (L). Koenig (S). Malabar Concan Ve., Stocks (Herb.
Hooker & Thomson Ind. Or.) (L). Simla, ?D. J. Yaclar 21 (US). Jalor (Dscha-
lor), Vaughan 592 (L). Mysore, W. B. 601 (L). Maisur and Carnatic, G.
Thomson (S). Brneaat: Hooker & Thomson (L). Sulp. Kurz (US). Calcutta
and vic., Helfer 73 (L, 8, US); Wallich 1196/D (L); Voigt collectors in 1834-41
(US). Assam: Col. Jenkins (L, US). TIRAP RIVER VALLEY: 12.4 mi. mark on
Ledo Road, Steinacker U.S. Amer. Typhus Comm. 6 (8, US). TIRAP-NAMCHIK
DIVIDE: 15.4 mi. mark on Ledo Road, Juan U.S. Amer. Typhus Comm. 320 (US).
NAMCHIK RIVER VALLEY: 21 mi. mark on Ledo Road, Juan U.S. Amer. Typhus
Comm. 523 (US).

UPPER BURMA: Myitkyina ReGion: Camp Landis area, Juan U.S. Amer.
Typhus Comm. 120 (8, US).

BURMA and MALAY PENINSULA: Herb. of late East India Co. 620 (L).
Spiff 620 (S).

THAILAND: SE. Part: Sriracha, Mrs. Collins 1880 (US).

MALAYA: Srtancor: Kanching, Tin mines, Nur Singapore field no. 34419
(L, US). Matacca: Griffith s.n. (L); Fish Culture Research Station, Shah 12
(L). Jonore: Kg. Selantai, Mersing, Burkill & Shah 2503 (L). Sincaprore:
Nee Soon village, J. Sinclair s.n. (L).

INDOCHINA: Trian near Bienhoa, Hvrard 57 (S), 64 (S). CamsBopia:
Pnompenh, Godefry 54 (L). Tonkin: Long-Tchéou, Simond s.n. (8).

GOLDBERG—GENUS MELOCHIA 307

CHINA: Harnan: cuine Mal: Ku Tung village: Pak Shik Ling, C. I. Lei 750
(US), 798 (L). Near: Ko Leng, Lau 572 (US). KWANGsI: NORTHERN PART:
Kwei-lin: San-min village, Tsang 28029 (US); Ta-chiang-yuan village, Tsang
28258 (US). Kwanruna: Canton, Levine 1633 (US). YANG SHAN: Yang Shan
(8S. of Linchow), Tsui 624 (US). vrapu: Tung Kao Shan, Tsang 21732 (S).
Formosa: Oldham 53 (8S). Takow, Henry 2037 (US). KiIaNasI: KIENAN:
Sai Hang Cheung, Lau 4267 (S, US). Curxrane: Tien Mo Shan, Ching Herb.
U. Nanking 9081 (US). ANuwer: Wu Yuan, Ching Herb. U. Nanking 8942
(US). Huprn: Henry 2315 (US).

SUNDA ISLANDS: Sumarra: Junghuhn 568 (L). Atjeh, Kp. Sigleng, N.
van Troeman, Asdat 155 (L). Lampong Estate Wai Lima, ?Sloet 332 (L), 460
(L). Taloe, Biinnemeyer 192 (L). Djambi, S. Lesing, Posthumus 973 (L).
NORTHERN Sumatra: Medan, Lérzing 13045 (L); Balige to Toba Lake, Lérzing
14796 (L). PADANG AND BEDAGEI: Plantation Badjalinggi, Lérzing 15204 (L).
West Coast: Pajakoemboeh, Stomps s.n. (L); Koerintji, Buinnemeyer 7994 (L).
W. K. G, Lago, Bunnemeyer 3677 (L); Padang Si Dimpoean: Padang Lawas:
Tapianoeli, Toroes 4447 (US). East Coast: ASAHAN:! Tandjoeng Balei, Hamel
1097 (S); Padang boelan, Stomps s.n. (L); asAHaN: Rantau Parapat, Bila, Toroes
1765 (L), 1556 (US); Aer Kandis (formerly Radja Mas), Toroes 2424 (US);
Sigamata, Toroes 2879 (US). Sournern Sumatra: Palembang, Monod de
Froideville 554 (L); Pladjoe, Rutter-Kooistra 10 (L); BANKA: Soengei Liat, Biinne-
meyer 1943 (L). Linaca ArcHIPELAGO: Bunnemeyer 6818 (or 8187). Riouw
ARCHIPELAGO: Buinnemeyer 6033 (L). Java: Houwink 973 (L), 1098 (L).
Zollingero s.n. (L). Junghuhn s.n. in part (L). Tanch Soereal, Raap 476 (L),
Pasoeroean, Kobn ? s.n. (L). Madjinang, Backer 18702 (L). Kedoe Karangbo-
long, Koorders 29722 8 (L). Depok, Halliers.n. (L). Priok near Jakarta Canal,
Raap 376 (L). Bogor and vic., Hallier 33 (L), 35a, b, ¢ (L); Hasselt s.n. (L);
Kampong Nangrang, Schiffner 2223 (L). Boerlage, Tjiomas s.n. (L). Bantard-
jaja estate, 18 km. 8S. of Rangkasbitung, Buwalda 2868 (L). pREANGER: C.
Ploem, (L). Banjoemas and vic., Kient 1588 (L), 2694 (L), 3423 (L);
Brinkman 587 (lL); Koorders 22129 B (L). Semarang, Karta 218 (L); Koorders
24862 6 (L). Djapara, Koorders 29722 8 (L). Pekalongan Soeba, Koorders
27456 8 (L). Bawean Island, Coert 1407 (L). MADOERA: Bangkalan, Backer
19023 (L). Borneo: Poeloe-lampei, Korthals s.n. (L). Dachlan 2422 (L).
NortH Borneo: Yates 72 (US). Tamu Darat at mi. 7 on the Kota Belud,
Ranau Road, Darnton 55 (L). Sarawak: Purseglove & Lundu P. 4515 (L).
CrLeBEs: Lasao, Eurekang, Kjellberg 1177 (S), 3897 (S). Bonto-Parang,
Biinnemeyer 10556 (L). Pampanoea, Noerkas 78 (?75) (L), 74 (L). Menado,
Gorontalo, ?V. Gorontalo 33a (L).

PHILIPPINES: Minpanao: coraBato: Marbel, Ala Valley, Santos 4863
(L, US); Buluan, Buluan Marsh, Barrio Siling, Santos 5956 (L, US). BUKIDNON:
Vic. of Tanculan, Féniz B.S. 26003 (L, US); Bugeaon, Cid 24 (L). pavao:
Davao, De Vore & Hoover 188 (US), Copeland 541 (US); Santa Cruz, De Vore &
Hoover 238 (US), R. S. Williams 2755 (US). Sraraao: Ramos & Pascasio B.S.
34898 (L). Nrcros: Mt. Katugasan, NW. slope of Canlaon Vol., Edafio 7648
(L). Curion: Merrill 537 (US). Minporo: Vicinity of San José, Lambert &
Brunson 108 (US). Luzon: San Miguel, Hallier sn. (L). CAMARINES NORTE:
Paracale, Ramos & Edafio B.S. 33625 (L). Manila, Quezon City, and vic.,
Andersson s.n. (S), Rogerson 1026 (US), Merrill 3432 (US), Steiner 662 (L, US),
Layosa 71 (L). Lacuna: Los Bafios, U.S. Explor. Exped. under Capt. Wilkes
(US). RiIzaL: Ramos 1978 (L), B.S. 138554 (8). Bosoboso, Ramos B.S. 1469
(US), U.S. Cemetery Camp McKinley, Steiner 721 (L, US). unron: Bauang,
Elmer 5603 (US). BaTaan: Lamao River, Mt. Mariveles, Borden For. Bur.

308 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

1917 (US). quezon: Alabat Island, Ramos & Edafio B.S. 48000 (5). TAYABAS:
Atimonan, Whitford 635 (US). sorsogon: Irosin (Mt. Bulusan), Elmer 14495
(L, 8, US). Bunacan: Angat, Merrill Species Blancoanae 325 (US). MouN-
TAIN: Benguet, Loher 2029 (US). Central Luzon: Loher 154 (US).

MOLUCCAS: Werar: Bloembergen 3631 (L). MHatmanerra: Kan, Kp.
Koesoe, Nedi 462 (L). S. peninsula, Weda, Anang 442 (L). AMBOINA (AMBON):
Robinson 1764 (L, US).

NEW GUINEA: Stephansort, Nyman 130 (8), Lewandowsky 56 (L). Cyclops
Mts., along path to Netar, Dreis 142 (L). Merauke, Verstug 1926 (L). F'inseh-
hafen, Weinland 246 (L, US). Noord, Alasrip 202 (L). Papua: Koitaki Carr
12682 (L). Norraern Div.: Near Gwaiari village, Hoogland 3669 (L, US).
Western Div.: Dagwa, Oriomo River, Brass 5958 (US); DARU ISLAND: Brass
6340 (L). Crnrrat Div.: NAKEO: Baroka, Brass 3698 (L, US).

AUSTRALIA: NorrHerN QuEENSLAND: Townsville, White 8967 (US).

SOLOMON ISLANDS: Bovaarnvitte: Karngu, Buin, Kajewski 2279 (L,
S, US). GuapatcanaL: Vie. Tenam River and the sea, Riley 42 (US).

CAROLINE ISLANDS: Ponape, Hallier s.n. (Li).

MARIANAS ISLANDS: Guam: Thompson Guam Exp. Sta 240 (US).

JAPAN: Kyusuu: Kawaminami in Hyuga, Togasi TNS. 1429 (US).
ousuMI: Koneshime-mura, Kawagoe Nov. 5, 1917 (US).

This species is characterized by its subcapitate terminal inflorescence
and nondimorphic flowers.

I dissected a flower from the type specimen of AZ. longebracteolata
J. Ar. The petals were separate and some of the bracts were shorter
than the petals. I have seen a range of hairiness in AZ. corchorifolia.
Arénes considers union of petals, bracts longer than the petals, and
hairiness of the bracts to be distinguishing characters of his species.

Section IV. Pyramis A. Goldberg, nom. nov.
Section Humelochia Griseb. Fl. Brit. W. Ind. 98. 1859.

Perennial, sometimes annual, herbs to shrubs, 0.08-5.0 meters
high, the root sometimes 2-5 times thicker than the base of the stem
and 0.5-1.3 em. wide, the young stems often canescent, sometimes
glabrate, hairs usually stellate, sometimes simple or forked, sometimes
tan, 0.1-1.6 mm. long, internodes 0.2-10 em. long; stipules 1.5-9.0
mm. long, 0.2-3.7 mm. wide, deltoid-acute or deltoid-acuminate,
lanceolate or linear, usually tomentulose and ciliate, hairs simple and
usually also stellate and forked, 0.1-2.0 mm. long; petiole 0.1—-5.1
em. long.

Lamina 0.6-12.7 cm. long, 0.3-7.7 cm. wide, usually lanceolate
to ovate, obovate or oval, the base rounded to cordate, or cuneate,
the apex acute or rounded, rarely acuminate, the upper surface
glabrous, hirsute, or tomentose and canescent, hairs stellate, forked
or simple, 0.1-2.0 mm. long, the lower surface glabrous to tomentulose
and canescent and woolly, hairs usually stellate, sometimes also
forked and simple, rarely a few glandular, 0.1-1.6 mm. long, the
margin irregularly crenate-serrate, serrations 0.7-7.0 mm. wide,

GOLDBERG—GENUS MELOCHIA 309

0.2-2.6 mm. high, 4-12 pairs of primary veins, usually 2 are basal.

Inflorescences usually leaf-opposed, sometimes also axillary, terminal
or 90° from the leaf, 1-7 em. long, the flowers in umbels, sometimes
subcapitate, the peduncle (0-)0.1-4.0 em. long, rarely with a primary
branch 0.3-1.1 cm. long, the pedicel 1-7 mm. long; bracts 0.5-6.5
mm. long, 0.1-1.2 mm. wide, similar in shape and pubescence to the
stipules.

Calyx 3.7-10.2 mm. long, 2.1-7.0 mm. wide, often tomentulose and
canescent, sometimes ciliate and hirsute, hairs stellate, usually also
forked or simple, 0.1-2.0 mm. long, the teeth 2.0-7.8 mm. long,
0.6—-2.3 mm. wide, the sinus usually acute, sometimes narrowly rounded;
petals purple, purple distally and yellow orange proximally, rose, red,
violet, blue, white, or yellow, 6.0-15.5 (-17) mm. long, 1.5-5.2 (-5.7)
mm. wide, usually obovate or oblanceolate, cuneate or spatulate,
attached to the staminal tube for 1-5 mm., the claw 0.6-4.0 mm. long,
often with scattered glandular hairs 0.05 mm. long.

Anther 0.7-2.0 mm. long, 0.4-1.0 mm. wide, oblong, slightly emar-
ginate at both ends; staminodes rarely present, 0.1-0.2 mm. long.

Longistylous form. Stamens (2.6-) 3.5-7.0 (-9.0) mm. long, the
filaments free for 0.7—-3.0 mm. ; pistil 4.3-10.9 (-12) mm. long, the styles
free for 2-6.7 mm., united for 0.5-4.0 mm., papillose for 0.5-1.5 mm.
at the apex.

Brevistylous form. Stamen 5.7-11.3 mm. long, the filaments free
for 2.5-7.5 mm.; pistil 3.7-6.9 (-8.0) mm. long, the styles free for
1.4-4.0 mm., rarely free to the ovary, united for 0.5-3.0 mm., papillose
for 0.3-1.0 mm. at the apex.

Fruit large, 5-10.5 (-12) mm. long, 3.8-17.0 (-22) mm. wide,
pentapterous, usually pyramidal, rarely ovoid, usually rostrate and
with acute, acuminate, or rounded lateral angles, the sulci between the
carpels deep, extending (one-fourth to) one-third to half way to the
center of the fruit, sometimes the base narrowing into a stipe 0.5-1.5
mm. long, tomentulose or hirsute-tomentulose, sometimes canescent,
or subglabrous, hairs stellate, sometimes also forked, rarely also simple,
0.1-1.3 mm. long; dehiscence loculicidal all along the dorsal and ventral
sutures, eventually also septicidal, rarely tardily dehiscent; seeds
1.7-3.0 mm. long, 1.1-2.0 mm. wide, trigonal, brown, 1 or 2 per cell.

Flowering and fruiting period usually long, sometimes throughout
the year.

Lecroryre: Melochia pyramidata L.

Disrrisution: Texas and Florida to Argentina, West Indies, one
species adventive in tropic and subtropic regions throughout the world,
usually in open disturbed, usually dry, sometimes wet, areas, at
0-2,500 meters elevation.

310 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

40. Melochia hermannioides St.-Hil. Fl. Bras. Mer. 1:1638, pl. 32. 1825.

Melochia parvifolia H.B.K. f. roseiflora Schum. & Hassl. pro parte. (Hassler
no. 6402) Bull. Herb. Boiss. IT, 4:70. 1903.

Melochia lacinulata Schum. & Hassl. Bull. Herb. Boiss. II, 4:69. 1903.

Melochia hermannioides St.-Hil. var. lacinulata (Schum. & Hassl.) Hassl.
Repert. Sp. Nov. Fedde 8:122. 1910.

Melochia hermannioides St.-Hil. var. lacinulata (Schum. & Hassl.) Hassl. f.
heterophylla and f. typica Hassl. Repert. Sp. Nov. Fedde 8:122. 1910.

?Melochia hermannioides St.-Hil. var. lanceolata Hassl. Repert. Sp. Nov.
Fedde 8:122. 1910.

A procumbent or decumbent trailing subshrub or perennial herb
0.1-0.5 meter long with several (5-8) slender stems from the base, the
root 2% to 3 times thicker than the stem, the young stems and petioles
hirsute-tomentose with short stellate hairs and simple curved hairs
0.1-0.3 mm. long and forked, and straight simple hairs 1.0-1.6 mm.
long, internodes 0.5-2.8 em. long; stipules 2.2-6.5 mm. long, 0.3-1.0
mm. wide, lanceolate to linear, mostly ciliate, hairs simple, 0.2-1.2
mm. long; petiole 0.2-1.7 cm. long.

Lamina 0.9-2.1 em. long, 0.6-1.4 em. wide, ovate, sometimes also
obovate or nearly orbicular, the base rounded to cuneate, the apex
rounded, the upper surface with rather sparse simple, and sometimes
also forked and stellate appressed hairs 0.2-1.5 mm. long, the lower
surface similarly pubescent, stellate hairs sometimes predominant,
hairs 0.2-1.2 mm. long, the margin crenate or crenate-serrate, crena-
tions 1.0-2.6 mm. wide, 0.5-1.9 mm. high, the base entire, 4-6 pairs
of primary veins, 1 or 2 are basal.

Inflorescences terminal, rarely also leaf-opposed, 1-3.2 cm. long, the
ultimate divisions umbellate, 5-8 flowers per umbel, the peduncle
0-2.2 em. long, rather slender, the pedicel 1-3 mm. long; bracts
2.0-6.0 mm. long, 0.2-0.6 mm. wide, linear to lanceolate-linear,
mostely ciliate, hairs simple, 0.5-1.2 mm. long.

Calyx 5.5-8.2 mm. long, 3.1-4.8 mm. wide, hirsute and ciliate,
hairs mostly simple, sometimes also forked and stellate, 0.2-1.5 mm.
long, the teeth 4.2-5.6 mm. long, 1.2-2.0 mm. wide, deltoid-acuminate,
the sinus acute; petals rose, blue, reddish violet, purple, or violet,
rarely white, 8.4-12.8 mm. long, 2.0-4.9 mm. wide, oblanceolate-
obovate-cuneate, attached to the stamens for 2.0-3.5 mm., the claw
1-3 mm. long.

Longistylous form. Stamens 5.7-6.0 mm. long, the filaments free
for 1.4-2.0 mm., the anther 1.2-1.6 mm. long, 0.6-0.8 mm. wide,
oblong, slightly emarginate at both ends; pistil 8.6-9.4 mm. long,
the styles filiform, free for 4.2—-5.1 mm., united for 1.6—2 mm., papillose
for 1-1.5 mm., the ovary narrowly elliptic-ovoid, the ovary and
styles up to 1 mm. of the apex covered with stellate, forked, and simple
hairs 0.1-1.3 mm. long.

GOLDBERG—GENUS MELOCHIA 311

Brevistylous form. Stamens 7.5-8.7 mm. long, the filaments free
for 5.0-5.5 mm., the anther 1.3-1.6 mm. long, 0.7-0.8 mm. wide;
pistil 4.2-5.2 mm. long, the styles free for 1.5-1.8 mm., filiform,
papillose for 0.5-0.8 mm., the ovary narrowly ovoid, pentagonal,
hairs on the ovary and styles forked and stellate, 0.3-1.4 mm. long,
stipe none.

Fruit 5.0-7.0 mm. long, of which the rostrum is 1.0-2.0 mm. long,
5.0-6.0 mm. wide, pentapterous, pyramidal, the lateral angle of the
wings rounded to subacute, points above the middle of the fruit, the
sulci extending a fourth to half way to the center of the fruit, pubes-
cence stellate, forked, and simple, hairs 0.1-1.2 mm. long, the longer
ones forked and simple, stipe none; dehiscence tardily loculicidal
along the dorsal suture and somewhat septicidal; seeds not quite
ripe, 2.4-2.9 mm. long, 1.4-1.7 mm. wide, trigonal, obovate, brownish.

Flowering and fruiting specimens collected from September to
March.

Type Locatity: Province of Misiones, Argentina, in herbaceous
fields. Type collected by St.-Hilaire, Cat. Cz, no. 2643; isotype cited
below.

Distrisution: Bolivia to Argentina in dry clay campo, sandy
herbaceous fields, on hills, at 150-2,500 meters elevation.

BOLIVIA: Tarisa: Padcaya, Fiebrig 2536 (L, S, US).

PARAGUAY: Cordillera de Villa-Rica, Hassler 8738 (K). Vicinity of
Caaguazt, Hassler 8899 (K, 8). Near Arroyo Primero (Apa), Hassler 8450
(S, type of M. lacinulata Schum. & Hassl.). Loma noérdlich der Cordillera de
Altos, Fiebrig 935 (K, L, US). Capilla Duarte near Villa-Rica, Balansa 2657
(IX). Trinidad near Asuncion, Rojas 1789 (S). Est. Armonia, Anisits 1875
(S). Paraguarf, Grosse A3575 (S), A3845 (8). Hiaty, Jorgensen 3915 (S, US).
Near Tobaty, Hassler 6402 (5S). ? Itapé, Jorgensen 4477 in part (S). Centurion,
between Rio Apa and Rio Aquidaban, Fiebrig 4207 (L).

ARGENTINA: Jusuy: nuMAHUACA: La Soledad, Venturi 8784 (US). Saura:
Wall 12 (S). Salta City, N. Caseros, Sparre 1184 (8S). Formosa: Colonia
Clorinda, Venturi 9150 (US). CérpoBa: San Roque, Hunziker dupl. 7067 (S).
CorRRIENTES: MBURUCUYA: Est. Sta. Maria, Pedersen 510 (8, US). santo
Tomé: Gob. Virasoro, Ibarrola 1289 (S). mTE. caseros: Mocoretd4, Ibarrola
2394 (S). MERCEDES: Justino Solaricercainas, Ybarrola 2633 (UC). MIsIONEs:
St.-Hilaire Cat. C2, no. 2643 (P, isotype, also US neg. 5668). Posadas, Bon-
pland, Ekman 185 (8).

This species is characterized by its relatively small leaves with
relatively large crenations, and by its small capsules. It is similar
to M. parvifolia H.B.K., but the latter is an erect shrub 0.5-3 meters
high with somewhat canescent young stems, and the serrations of the
leaves are about a third to half as high as those of M. hermannioides.
It is also similar to M4. anomala Griseb., but the latter generally has
larger, more densely hairy leaves and shorter peduncles (0-0.3 cm.).

312 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Schumann (47) placed this species in the section Mougeotia, and
in his key to the species he claimed that the inflorescence is axillary.
However, the fruit is pyramidal and the inflorescence is in part
leaf-opposed as in the section Pyramis; in the section Mougeotia,
the fruit is subglobose and the inflorescence is axillary rather than
leaf-opposed.

Hassler (23) described two varieties of M. hermannioides: The
type of variety lanceolata Hassl. is Fiebrig 4207, collected at Centurion,
Paraguay; the type of variety lacinulata (Schum. & Hassl.) Hassl. is
Hassler 8450, collected near Arroyo Primero (Apa), and cited above.
The variety lanceolata differs from the type of the species in having
lanceolate leaves. I have not seen the type of this variety and with-
hold judgment as to whether it is distinct. I do not consider the type
of the other variety to differ appreciably from the isotype of the species.

41. Melochia parvifolia H. B. K. Nov. Gen. & Sp. 5:325. 1823.

An erect or spreading, branched herb to shrub 0.5-3.0 meters high,
the young stems and petioles somewhat canescent, hirsute-tomentu-
lose-villous, hairs stellate, 0.2-1.0 mm. long, internodes 1.0-7.5 em.
long; stipules 2-4 mm. long, 0.4-0.9 mm. wide, deltoid-acuminate,
lanceolate or nearly linear, hirsute to tomentulose, ciliate, hairs
stellate, forked, and simple, 0.1-1.0 mm. long; petiole 0.1-1.3 em. long.

Lamina relatively small, 0.6-4.5 em. long, 0.3-2.8 em. wide, obovate
to elliptic, rarely ovate, the base cuneate or rounded, the apex rounded,
the upper surface green, glabrous, subglabrous or tomentulose, hairs
simple, forked, and stellate, usually 0.1 (rarely a few to 0.8) mm. long,
mostly between the veins, the lower surface green, hirsute to tomen-
tulose, hairs mostly on the veins, stellate and forked, 0.1-0.6 mm.
long, puberulous between the veins, hairs 0.1 mm. long, the margin
irregularly crenate-serrate, serrations 0.7—3.0 mm. wide, 0.2—-1.0 mm.
high, 4-7 pairs of primary veins, usually one is basal.

Inflorescences leaf-opposed, 1.3-3.8 em. long, rarely also terminal,
clusters at the upper 2-8 nodes, 3-20 flowers per cluster, umbellate-
subcapitate, the peduncle 0.1-1.7 em. long, 0.6-1.0 mm. wide, rarely
branched, the pedicel 1-4 mm. long; bracts 1-4 mm. long, 0.2-0.5 mm.
wide, deltoid-acuminate, lanceolate or linear, hirsute to tomentulose,
ciliate, hairs simple and stellate, 0.2-0.8 mm. long.

Calyx 5.0-9.5 mm. long, 3.7-5.6 mm. wide, hirsute to tomentulose,
ciliate, hairs forked, simple, and stellate, 0.1-1.0 mm. long, the teeth
3.3-5.0 mm. long, 1.2-1.8 mm. wide, deltoid- to ovate-acuminate, the
sinus acute to narrowly rounded; petals usually white, sometimes
yellow or rose, rarely pale lilac, 7-11.5 mm. long, 2.5-4.8 mm. wide,
obovate- to oblanceolate-spatulate or oblanceolate-cuneate, sometimes

GOLDBERG—GENUS MELOCHIA 313

with scattered glandular hairs 0.05 mm. long, attached to the staminal
tube for 2-3 mm., the claw 1-3 mm. long.

Longistylous form. Stamens 4-5.8 mm. long, the filaments free for
1-2.5 mm., the anther 1.1-1.5 mm. long, 0.5-0.7 mm. wide, oblong,
slightly emarginate at both ends, more so at the base; pistil 7.5-9.5
mm. long, the styles free for 3.4-4.5 mm., united for 1.5—3.5 mm.,
papillose for 1 mm. at the apex, the ovary ellipsoid to elliptic-ovoid,
slightly pentagonal, the ovary and styles tomentulose, hairs stellate
and forked, 0.1-0.5 mm. long, the stipe 0-0.5 mm. long.

Brevistylous form. Stamens 8-9.5 mm. long, the filaments free for
9-6.8 mm., the anther 1.1-1.7 mm. long, 0.6-0.7 mm. wide, shape the
same as that above; pistil 5-5.7 mm. long, the styles free for 1.6-2.2
mm., united for 1.0-1.7 mm., papillose for 0.5-1.0 mm. at the apex,
the ovary ellipsoid, occasionally of 6 carpels, hirsute or sericeous-
tomentulose, the styles hirsute to their separation, hairs stellate,
0.2-0.8 mm. long, stipe none.

Fruit 5-7 mm. long of which 1-2 mm. is rostrum, 4.2-7.2 mm.
wide, pentapterous, pyramidal, the sulci extending one-third of way
to the center of the fruit, the angles of the wings near the middle of
the fruit, short, acute to rounded, hirsute-tomentulose, hairs simple,
forked, and stellate, 0.2-0.8 mm. long, stipe none; dehiscence loculi-
cidal along the dorsal and ventral sutures; seeds 2~2.7 mm. long,
1.2-1.7 mm. wide, trigonal, obliquely obovate, brown to black, one
per cell.

Flowering and fruiting specimens collected throughout the year.

Key to Varieties of M. parvifolia

1. Lamina subglabrous, hairs mostly simple, base usually cuneate,
41a. M. parvifolia var. parvifolia
1 Lamina hairier, sometimes lower surface tomentulose, hairs mostly stellate
and forked, base more frequently rounded than in the typical variety.
41b. M. parvifolia var. fasciculata

41a. Melochia parvifolia H. B. K. var. parvifolia

? Melochia parviflora Loud. Hort. Brit. 269. 1830.

Melochia scordiifolia Turez. Bull. Soc. Nat. Mose. 31(1):207. 1858.

Melochia parvifolia a normalis O. Ktze. Rev. Gen. 3(2):25. 1898.

Melochia parvifolia 8 Esperanzae O. Ktze. Rev. Gen. 3(2):25. 1898.

Melochia parvifolia H.B.K. var. genuina Hassl. Repert. Sp. Nov. Fedde
$:120. 1910.

Melochia parvifolia H.B.K. var. genuina Hassl. f. induta Hassl. Repert. Sp.
Nov. Fedde 8:120. 1910.

Melochia obovata Urb. Symb. Ant. 9:233. 1924,

The typical variety is sparsely hairy; the leaves are small, the
lamina generally not exceeding 2.5 cm. in length and 1.3 em. in
width, and the base is usually cuneate.

314 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Typr tocauiry: The type was collected by Humboldt in an arid
region near Caracas, Venezuela.

Disrrrpution: Cuba and northern South America to Argentina,
most commonly in savanna, also in campo and in cultivated and
fallow land, in dry or moist places, in sandy or serpentine soil, and
among granite rocks, at 0-1,400 meters elevation, mostly at lower
elevations.

CUBA: OrtentE: Santiago de Cuba, Siboney, Bro. Clemente 7005 (US).
Santiago de Cuba, Vista Alegre, Acufia & Morton 3896 (US). Near Santiago
de Cuba, Ekman 7694 (S, type of Melochia obovata Urb., also US neg. 5605),
8894 (S, isotype M. obovata Urb.). Siboney, Morton 3957 (US). Near El
Cobre, Bro. Clemente 6285 (US); Bro. Leén 16773 (US).

BRITISH GUIANA: Rupunvnt: Ireng River, 45 mi. N. of Lethem, Irwin
833 (US).

VENEZUELA: ?Cerro del Galipan, Eggers 13192 (L.) Lower Orinoco,
Sacupana, Rusby & Squires 36 (MICH, R, 8, US). Oronoco Delta, Cano del
Uricoa-San Antonio, Bond et al. 134 (US). Las Trincheras, Warming 267 or 266
(US). Cumanacoa, Mocquerys 764 (US). “Venezuela, ete.,”’ Funck & Schlim
564 (LE, and US neg. 5268, isotype of Melochia scordiifolia Turcz.). GuArico:
La Rubiera near Calabozo, Grisol 38 (US). Cordoncito, Gines 4121 (US).
Mrranpa: Los Chorros, Eggers 13092 (US). Rio Chico, Jahn 1223 (US).
Higuerote, Gines 4585 (US). FEDERAL District: Caracas, Humboldt, F neg.
9807 of the type (US). Near Caracas, Middle Cotiza, Pittier 7213 (US). Around
Caracas, San Liézara, Pittier 9746 (US). Around Caracas, Pittier 6171 (US).
Sabana de Conejo Blanco, El Valle, Fernandez 120 (US). Araaua: La Trinidad
de Maracay, Pittier 5796 (US), 5820 (US). Maracay, Vogl 337 (S). Near
colony of Tovar, Fendler 112 (GH). CaraBono: Puerto Cabello, Suringar Mar.
1885 (L). ?Carasoso: Puerto Cabello, Kuntze 1719 (US). Cosnpfs: Culebra
Lagoon near 8. Carlos, Pittier 11705 (US). Apure: Llanos del Alto-Apure,
Hato Metapalo, Jahn 205 (US). Puerto Pdez, Valez 2698 (US). Rio Meta,
La Ochovera, Cuatrecasas 4183 (US). Botfvar: Ciudad Bolivar, Holt & Gehriger
114 (US), 140 (US). Ciudad Bolivar and vicinity on the Orinoco, L. H. & E. Z.
Bailey 1626 (US). Between Ciudad Bolfvar and Rio Pao, Killip 37214 (US).
Morichal Sta. Isabel, Cardona 699 in part (US). El Palmar, L. Williams 12869
(US).

COLOMBIA: Artxdntico: Barranquilla and vicinity, Fosberg 21424 (US);
Bro. Elias 109 (US), 178 (US), 274 (US), 995 (US), 1324 (US); Dugand 5437
(US), 5437 B (US), 5437 C (US). Usiacuri, Dugand & Garcia Barriga 2313
(US). Between Palmar de Varela and Ponedera on the banks of the Rio Mag-
dalena, Dugand & Garcia Barriga 2541 (US). Between Sabanalarga and
Campeche, Dugand & Jaramillo 2762 (US). Vicinity of Galapa and Baranoa,
Dugand & Jaramillo 3269 (US). Vicuapa: Puerto Carrefio on the Rio Orinoco,
Cuatrecasas 4067 (US).

BRAZIL: Western part, Tamberlik (W). Rio Branco: Ule 7971 (L, US).
Sao Jorje, Rio Flexa, Frées 23129 (US). SURUMU: Serra do Mel, Ule 8226 (L).
Amazonas: 8. Marcos, Luetzelburg 20509 in part (R), 20510 in part (R). GotAs:
Porto Real to Santa Anna, Burchell 8856 (L), 9083 (L). Marto Grosso; Cuiabé4,
? Riedel 793 (K); Malme s.n. (8). Coxipé da Ponte, Cuiab4, Hoehne 3024 (R, US),
3025 (R). Porto Esperidiao, Hoehne 775 (R). Corumba4, Hoehne 4839 (R),
4875 (R).

GOLDBERG—GENUS MELOCHIA 315

PARAGUAY: Rio Paraguay, ? Tonciére, Sandeman 4873 (K). Alto-Paraguay,
Chaco 21°S. lat., Fiebrig 1215 (8). Near Concepcion, Hassler 7399 (8S). Road to
“Yerbales” montium “Sierra de Maracayt,” Hassler 4989 (S, W).

ARGENTINA: Tucum4n: monreros: Kilometro 1220, Herrera 161 in
part (5).

This species is close to Melochia hermannioides St.-Hil., but the
latter is procumbent or decumbent and small, whereas M. parvifolia
is generally erect and can be a tall shrub. The leaves of M. herman-
nioudes are broader, rounder, and have larger teeth, and the flowers
are rose, blue, purple or violet rather than generally white. The
inflorescences of M. hermannioides are more frequently terminal and
less frequently leaf-opposed. The species is also close to Melochia
morongit Britt., but the latter has longer and narrower leaves which
are hairier on the lower surface, and the apex is acutely pointed rather
than rounded; the penduncles of AM. morongii are longer. It is also
related to Melochia anomala Griseb., but the leaves of the latter are
generally larger, generally ovate rather than obovate, the serrations
are larger, and the base rounded rather than often cuneate. The
flowers are usually rose, blue, purple or violet rather than usually

white.

41b. Melochia parvifolia H.B.K. var. fasciculata (Benth.) Hassl. Repert. Sp. Nov.

Fedde 8:121 1910.

Melochia fasciculata Benth. in Hook. Journ. Bot. 4:127. 1841,

Melochia parvifolia H.B.K. f. albiflora Schum. & Hassl. Bull. Herb. Boiss.
II, 4:70. 1908.

Melochia parvifolia H.B.K. var. fasciculata (Benth.) Hassl. f. microphylla
Hassl. Repert. Sp. Nov. Fedde 8:121. 1910.

Melochia parvifolia H.B.K. var. fasciculata (Benth.) Hassl. f. microphylla
Hassl. subf. transiens Hassl. Repert. Sp. Nov. Fedde 8:121. 1910.

Melochia parvifolia H.B.K. var, fasciculata (Benth.) Hassl. f. typica Hassl.
Repert. Sp. Nov. Fedde 8:121. 1910.

This variety has hairier leaves, particularly on the lower surface,
and the hairs are generally stellate and forked rather than simple.
The lamina is often larger and the base is rounded rather than cuneate.

Type tocauity: The syntypes were collected by Schomburgk, no.
738, at Pirara, British Guiana, by Gardner, no. 2068 (?2060) in Piaul,
Brazil, and by Blanchet, no. 2750, at Utinga, Belém, Par4, Brazil.
Schomburgk no. 738 is cited below.

Disrrisution: British Guiana, northern Brazil, and Paraguay in
dry savannas, on rather humid cultivated areas, at about 200 meters
elevation.

BRITISH GUIANA: Pirara, Schomburgk 738 (L, US, syntype of Melochia
fasciculata Benth.), also F neg. 35379 ex P of Schomburgk 738 (US).

BRAZIL: Amazonas: Serra das Guaribas, Luetzelburg 21950 (R). Passarfo,
Rio Araricuera, Luetzelburg 20573 in part (R).

316 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

PARAGUAY: Colonia Risso near Rio Apa, Malme 1048 (8, US). Fuerte
Olimpo, Anisits 2107 (5).

42. Melochia morongii Britt. Ann. New York Acad. Sci. 7:62. 1890.

Melochia Morongii Britt. var. denudata Fries, Svensk. Vet. Akad. Handl.
42(12):9. 1908.

Melochia parvifolia T.B.K. var. Morongit (Britt.) Hassl. Repert. Sp. Nov.
Fedde $:121. 1910.

Melochia parvifolia H.B.K. var. Morongii (Britt.) Hassl. f. genuina Hassl.
Repert. Sp. Nov. Fedde 8:121. 1910.

Melochia parvifolia H.B.K. var. Morongii (Britt.) Hassl. f. genuina Hassl.
subf. denudata (R. E. Fries) Hassl. Repert. Sp. Nov. Fedde 8:121, 1910.

Melochia parvifolia H.B.K. var. Morongii (Britt.) Hassl. f. intermedia Hassl.
Repert. Sp. Nov. Fedde 8:121. 1910.

Melochia parvifolia H.B.K. var. Morongit (Britt.) Hassl. f. tomentosa Hassl.
Repert. Sp. Nov. Fedde 8:122. 1910.

A branched, erect subshrub 0.38-0.6 meter high with 2-5 stems from
the base, the root 0.8 cm. wide, 2-5 times as thick as the base of the
stems, the young stems and petioles canescent, tomentose, hairs
slender, stellate, 0.5-1.3 mm. long, internodes 2-6 cm. long; stipules
3.0-6.8 mm. long, 0.3-0.5 mm. wide, deltoid-acuminate, linear or
lanceolate, mostly ciliate, hairs simple, 0.3-1.2 mm. long, a few forked
and stellate, 0.2 mm. long; petiole 0.5-2.0 em. long.

Lamina tomentulose, sometimes glabrescent, 2.0-7.5 cm. long, 0.8-
3.0 em. wide, ovate to lanceolate, the base rounded, the apex acute to
rounded, the upper surface greenish or canescent, tomentulose, hairs
simple, appressed, forked, and stellate, 0.1-1.0 mm. long, the lower
surface canescent, tomentulose, hairs stellate, 0.3-0.7 mm. long,
woolly to the touch, the margin irregularly crenate-serrate, serrations
1-5 mm. wide, 0.3-1.2 mm. high, 5-10 pairs of primary veins, | or 2
are basal.

Inflorescences leaf-opposed, 2.5-7.0 cm. long, sometimes also
axillary, 1-1.5 cm. long, at 2-4 nodes, and terminal, extending 3-3.6
cm. beyond the first leaf, in subcapitate umbels, 5-12 flowers per
umbel, the peduncle usually fairly long, 0.7-4.0 em. long, 0.6-1.4 mm.
wide, sometimes with a branch 1 cm. long, the pedicel 2-5 mm. long;
bracts 1.5-5.0 mm. long, 0.1-0.3 mm. wide, linear, ciliate and tomen-
tulose, hairs simple, forked, and stellate, 0.2-1.1 mm. long.

Calyx 6.7-10.2 mm. long, 4-5 mm. wide, densely tomentulose, hairs
stellate and forked, 0.1-1.0 mm. long, the teeth 4.5-7.8 mm. long,
1.3-1.7 mm. wide, lanceolate- to deltoid-acuminate, the sinus acute to
slightly rounded; petals blue, purple, lilac, or dirty white, 9.5-11.5
mm. long, 2.0-3.8 mm. wide, obovate-oblanceolate or obovate-oblong,
attached to the stamens for 1-3.3 mm., the claw 1-3 mm. long, some-
times glandular hairs 0.05 mm. long scattered over both surfaces.

GOLDBERG—GENUS MELOCHIA 317

Longistylous form. Stamens 5.5 mm. long, the filaments free for
2.0-2.3 mm., the anther 1.3-1.4 mm. long, 0.7 mm. wide, oblong,
slightly emarginate at both ends; pistil 10.5 mm. long, the styles free
for 4.5-6.5 mm., united for 1.5-4.0 mm., filiform, papillose for 1.5 mm.
at the apex, the ovary and styles tomentulose, hairs stellate, 0.2-0.5
mm. long, the ovary ovoid, stipe none.

Brevistylous form. Stamens 8.1-10.5 mm. long, the filaments free
for 6.0-7.5 mm., the anther 1.3-1.8 mm. long, 0.5-0.8 mm. wide,
shape the same as that above, staminodes sometimes present, 0.1-0.2
mm. long; pistil 5.0-6.5 mm. long, the styles free for 1.5-2.5 mm.,
united for 1-2 mm., filiform, papillose for 0.7-1.0 mm. at the apex,
the ovary elliptic-ovoid, united parts of the styles and upper part
of the ovary hirsute-tomentulose, hairs stellate and forked, 0.2-1.0
mm. long.

Fruit 6-7 mm. long, of which the rostrum is 1.52.0 mm. long, 4.0—
5.8 mm. wide, pentapterous, pyramidal, the wings short, points near
the middle of the fruit, angle acute to rounded, the sulci extending
a fourth to half way to the center of the fruit, tomentulose, hairs
stellate, 0.1-0.6 mm. long, the stipe of glandular tissue 0-0.3 mm. long;
dehiscence loculicidal all along the dorsal suture ; seeds 2.4-2.6 mm.
long, 1.5-1.7 mm. wide, brown, one per cell.

Flowering and fruiting specimens collected from October to Febru-
ary.

TypE Locatity: The type was collected by Morong, no. 201a, at
Asuncién, Paraguay, and is cited below.

Disrrisution: Southwestern Brazil, Paraguay, and Misiones,
Argentina, in grassy fields, in a valley, on cultivated land, and on
calcareous campo, at 220-225 meters elevation.

BRAZIL: Minas Gerats: Ituiutaba, Macedo 1034 (US). Gotrks: Jataf,
Balsam, Macedo 1504 (US).

PARAGUAY: Anisits 2731 (8). Est. Santa Maria, Anisits 2529 (S, type of
Melochia morongii Britt. var. denudata R. E. Fries, also U.S. neg. 5613). Est.
Tagatiya, Anisits 2640 (S). Asuncién, Morong 201a (NY, type; US, isotype).
Along road between Caacupé and Tobaty, Archer & Rojas 4864 (US). Rio Apa,
Hassler 7744 (8S, W). Centurion, between Rio Apa and Rio Aquidaban, Fiebrig
4620 (L). canrera: Colonia Alborada, Montes 3254 (US). Gran Cuaco: On
the western bank of the River Paraguay, Hassler 2406 (K), 2406a (Ix).

ARGENTINA: Misiones: CANDELARIA: Loreto, Montes 24B (US); Ekman
178 (US).

Fries (15) made the variety denudata on the basis of the glabrescent
nature of the leaves of some specimens. There is considerable varia-
tion in the vestiture of the leaves, but I consider the transition from
one type to another to be too gradual to warrant making a variety
based on vestiture alone. Hassler (23) considered M/. morongii to be a
variety of M. parvifolia H.B.K., and he described three forms in the

227-051—67——_9

318 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

variety. The leaves of Af. morongii are generally ovate to lanceolate,
those of Af. parvifolia are usually obovate to elliptic. The leaves are
usually larger and usually hairier than those of Af. parvifolia. The
peduncle is usually longer than that of Af. parvifolia. Melochia
parvifolia is a taller plant and its flowers are usually white rather than
usually colored.

The species is also related to Mf. anomala Griseb., but the inflo-
rescences and peduncle are shorter in the latter species, and the
serrations of the leaves are usually larger.

43. Melochia anomala Griseb. Abh. Wiss. Goett. 19:93. 1874. PLATE 7

A more or less erect shrub 0.5—1 meter high with several (4) stems
from the base, the root thickened, 0.5-1.0 em. wide, twice the di-
ameter of the stem, the young stems and petioles usually lightly
tomentose-hirsute, somewhat canescent, hairs simple, forked, and
stellate, 0.3-1.4 mm. long, internodes 1-5.5 em. long; stipules 3.5-
6.0 mm. long, 0.2-0.9 mm. wide, deltoid-acuminate, lanceolate, or
nearly linear, ciliate, also a few simple hairs, rarely also forked on the
abaxial surface, 0.3-1.2 mm. long; petiole 0.2—2.5 em. long.

Lamina 1-6 em. long, 0.8-4.0 cm. wide, ovate, ovate-lanceolate,
lanceolate, or oblong, the base rounded, the apex rounded to acute,
the upper surface green, covered with appressed simple hairs, rarely
also forked and stellate, 0.2-1.3 mm. long, the lower surface lightly
tomentose, slightly canescent, hairs usually simple, forked, and
stellate, 0.3-1.2 mm. long, the margin irregularly crenate-serrate
almost to the base, serrations 1-3.8 mm. wide, 0.4—2.5 mm. high, 6-7
(-10) pairs of primary veins, 2 are basal.

Inflorescences leaf-opposed at 3-6 upper nodes, in subcapitate
umbels 1-1.3 (-3.0) em. long, 3-7 (-12) flowers per umbel, the pe-
duncle 0-0.5 (-2.3) em. long, the pedicel 1-2 mm. long; bracts 3—5.5
mm. long, 0.2-0.5 mm. wide, linear to lanceolate, mostly ciliate, few
hairs on the abaxial surface, hairs simple, 0.2-1.3 mm. long.

Calyx 7-7.5 mm. long, 5.6-6.0 mm. wide, tomentose-canescent,
also ciliate, hairs simple, forked, and stellate, 0.1-1.3 mm. long, the
teeth 5-6 mm. long, 1.2-1.8 mm. wide, deltoid- to lanceolate-acumi-
nate, the sinus narrowly rounded; petals violet, purple, rose, blue,
rarely whitish, 9.8-10.5 mm. long, (1.7-) 3.5-3.9 mm. wide, usually
obovate, with scattered glandular hairs 0.05 mm. long, attached to
the staminal tube for 2-3 mm., the claw 1-2.5 mm. long.

Longistylous form. Stamens 5.4-5.7 mm. long, the filaments free
for 1.6-2.5 mm., the anther 1.3-1.6 mm. long, 0.6-1.0 mm. wide,
oblong, slightly emarginate at both ends; pistil 7.8 mm. long, the
styles filiform, free for 3.4-3.7 mm., united for 2 mm., papillose for
0.8-1.0 mm., the ovary ovoid, mostly upper part with forked and

GOLDBERG—GENUS MELOCHIA 319

stellate hairs 0.3-1.0 mm. long, and stellate hairs 0.2 mm. long nearly
to the tips of the styles.

Brevistylous form not seen.

Fruit (5-) 6.5-8.0 mm. long, of which the rostrum is 2.0-2.5 mm.,
(5.5—) 6.2-7.0 (-9.0) mm. wide, pentapterous, pyramidal, the angle of
wing acute, about the middle of the fruit, the sulci extending one-
third the way to the center of the fruit, tomentulose, hairs stellate,
0.1-0.5 mm. long, a few forked, up to 0.7 mm. long, stipe none; de-
hiscence loculicidal along the dorsal suture; seeds (not ripe) 2.5 mm.
long, 1.5 mm. wide, obovate, trigonal, brownish, one per cell.

Flowering specimens collected from October to April; fruiting ones,
from October to May.

Type ocauity: On hills exposed to the sua near Malagueno,
Cérdoba, Argentina. Type collected by Lorentz, no. 503, cited below.

Distrisution: Bolivia and Argentina on slopes of hills in red soil,
campo, uncultivated, dry, hard land, at 100-1,200 meters elevation,

BOLIVIA: Santa Cruz: Weddell 3588 (P). Cabezas (Cordillera), Peredo 39
(UC), 260 (UC). a. rBaANuz: La Guardia, Krapovickas 9038 (LIL). Tara:
Fries 1135 (8).

ARGENTINA: Saura: antittas: Cerro Negro, Venturi 10292 (A, 8, US).
orAN: Tartagal, Schreiter 3893 (S). Abra Grande, Venturi 5519 (GH, 8, US).
La Cantera, Pierott’ 205 (UC). La Calera, Pierotté 197 (UC). At foot of hill E.
of Salta, Bartlett 19722 (US). ‘Monte’ at Experiment Station, Coronel Moldes,
Barilett 19668 (US). Capital, San Lorenzo, Venturi 5085 (A, US). Joaquin V.
Gonzdles, Aguilar 257 (UC). TucumM4n: BuRROYACO: El Puestito, Venturi 7529
(S, US). El Timb6é, Venturi 2430 (US). Rio cHico: La Cocha, Bailetti 9 (8).
cAPITAL: Rio Sali, Venturi 2756 (A, US). ?Mataderos, Venturi 2190 in part
(US). vTRANcAS: Tapia, Venturi 2360 (A, US). Road to S. Pedro de Colalao,
Venturi 4219 (US). cruz atta: Campo Alegre, Venturi 2012 (US). Las Cajas,
Venturi 2108 (US). SantTIAGO DEL EsTERO: C. PELLEGRINI: Carro del Remate,
Venturi 5972 (GH, US). Caco: Border of Rio Las Garzas, E. R. Wagner s.n.
(P). NAPaLPf: Pampa del Infierno, Buratovich 525 (W). Cérvosa: Kuntze
Dec. 1891 (NY). F neg. 9596 ex. B of Lorentz 110 (US). Maleguefi, foothills
of Cérdoba, Lorentz 503 (GOET, type, also US neg. 5239). Pueblo Giiemes,
Ruiz de Huidobri 413 (UC). puntita: San Roque, Hunziker dupl. 5997 (S).

This species is closely related to M. hermannioides St.-Hil., but
M. anomala has larger leaves and usually shorter peduncles. The
entire plant is hairier, particularly the leaf and calyx. It is also
related to M. parvifolia H.B.K., but it has larger, hairier leaves with
larger serrations.

44. Melochia werdermannii A. Goldberg, sp. nov. PLATE 8

Suffrutex, utrinque valde tomentosus molle incanus, pilis stellatis,
0.3-1.0 mm. longis praeditus; petiolis 1-1.7 em. longis; laminis foli-
orum 4.5-7.0 cm. longis, 2.2-4.9 cm. latis, ovatis vel aliquatenis
rotundatis; inflorescentiis oppositifoliis, cymoso-umbellatis, umbellis
5-15 floris, pedunculis brevis et crassis, 0.2-0.8 cm. longis, 1.2-1.5

320 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

mm. latis, pedicellis brevis, 1.5-2.5 mm. longis; calycibus 7-9.2 mm.
longis; fructibus perangustatis, 6-7 mm. longis, 3.8-4.8 mm. latis,
pentapteris, pyramidatis, angustatis ad basin, alis perangustatis,
acuminibus lateralibus rotundatis vel subacutis, sessilibus; dorsaliter
plene loculicide vix septicide dehiscentibus.

An erect, usually branched suffrutex, the young stems and petioles
densely tomentose, canescent, hairs stellate, 0.5-1.0 mm. long, inter-
nodes 2-6 cm. long; stipules 4-6.5 mm. long, 0.5-0.8 mm. wide, lance-
olate, densely tomentose, hairs stellate, 0.2-0.5 mm. long; petiole
1-1.7 em. long.

Lamina 4.5-7.0 em. long, 2.24.9 em. wide, lanceolate-ovate to
ovate, sometimes also suboblong, the base rounded, the apex acute
to somewhat rounded, both surfaces densely tomentose, canescent,
hairs stellate, 0.3-0.5 mm. long, velvety to the touch, the margin
irregularly crenate-serrate, serrations 1-6 mm. wide, 0.31.6 mm. high,
10-12 pairs of primary veins, 2 are basal.

Inflorescences 1-2 em. long, leaf-opposed, at 5-9 nodes, umbellate,
5-15 flowers per umbel, the peduncle short and relatively thick,
0.2-0.8 cm. long, 1.2-1.5 mm. wide, the pedicel 1.5-2.5 mm. long;
bracts 2-5.3 mm. long, 0.2-0.6 mm. wide, linear to lanceolate,
tomentose, hairs stellate, few forked and simple, 0.2-1.8 mm. long.

Calyx 7-9.2 mm. long, 4.3-5.5 mm. wide, densely tomentose,
canescent, hairs stellate, 0.2-0.7 mm. long, the teeth 4.7-5.7 mm.
long, 1.7-2.3 mm. wide, deltoid- to ovate-acuminate, or sublanceolate,
the midrib often prominent, the sinus acute to narrowly rounded;
petals 7.5-10.8 mm. long, 1.5-3.4 mm. wide, obovate-oblanceolate,
attached to the staminal tube for 2.5-3.0 mm., the claw 1.5-3.0
mm. long, with scattered glandular hairs 0.05 mm. long, faint indica-
tion of purple or rose in the dried condition.

Longistylous form. Stamens 4.7-5.8 mm. long, the filaments free
for 1.7 mm., the anther 1.5 mm. long, 0.8 mm. wide, oblong, slightly
emarginate at both ends; pistil 7.2 mm. long, the styles free for 3.2
mm.

Brevistylous form. Stamens 7-10 mm. long, the filaments free for
4.0-6.5 mm., the anther 1.6—-1.8 mm. long, 0.6-0.9 mm. wide, shape
same as that above; pistil 5.2-5.7 mm. long, the styles filiform, free
for 1.5-1.8 mm., united for 0.5-1.5 mm., stigmatic for 0.3-0.7 mm.,
the ovary ovoid, considerably narrowed at the base, tomentulose,
hairs stellate, 0.2—-0.3 mm. long.

Fruit narrow, 6-7 mm. long, 3.8-4.8 mm. wide, narrowed toward
the base, the rostrum 1.2—-2.0 mm. long, pentapterous, pyramidal, the
wings very narrow, outer angle of the wing slightly below the middle
of the fruit, rounded to subacute, the sulci between the carpels

GOLDBERG—GENUS MELOCHIA 321

extending a fifth to a third of way to the center of the fruit, tomentulose,
hairs stellate, 0.2-0.3 mm. long, stipe none; dehiscence loculicidal all
along the dorsal suture, sometimes tardily so, very slightly septicidal ;
seeds 2.4-2.9 mm. long, 1.4-1.6 mm. wide, obovate, trigonal, brown,
1 per cell.

Flowering and fruiting specimens collected in February and
October, few fruits present in the latter month.

TyPE Locauity: Missiones Guarayos-Santa Cruz de la Sierra,
Santa Cruz, Bolivia. The type was collected by Werdermann, no.
2606, and is cited below.

Disrrisution: Santa Cruz, Bolivia, at 250-300 meters elevation.

BOLIVIA: Santa Cruz: Missiones Guarayos-Santa Cruz de la Sierra,
Werdermann 2606 (S, type, also US neg. 5607). cHIQurTos: Roboré, Krapovickas
9190 (LIL).

This species differs from the other densely stellate-tomentulose,
canescent species of this section in having the following combination
of characters: Very narrow fruits, short, thick peduncles, and a
relatively large lamina, velvety to the touch. It is most closely
related to M. anomala.

45. Melochia illicioides Schum. in Mart. Fl]. Bras. 12(3):31. 1886.
Melochia decandra “‘ Willd.”’ ex Schum. in Mart. Fl. Bras. 12(3):31. 1886, in
synonymy.

A simple or branched erect suffrutex up to 1 meter high, the young
stems tomentulose-canescent, hairs stellate, 0.1-0.2 mm. long,
internodes 1-8 cm. long; stipules caducous, 1.7 mm. long, 1 mm. wide,
deltoid-acute, tomentulose, hairs stellate, 0.1 mm. long; petiole
0.4-3.3 cm. long.

Lamina 2-5.2 cm. long, 1.2-3.6 cm. wide, lanceolate to lanceolate-
ovate, the base subcordate, the apex acute, the upper surface greenish-
canescent, tomentulose, hairs stellate, 0.1 mm. long, the lower surface
canescent, tomentulose, hairs stellate, 0.1-0.2 mm. long, both surfaces
velvety to the touch, the margin irregularly crenate-serrate, serrations
1.2-6.0 mm. wide, 0.5-1.6 mm. high, 8 pairs of primary veins, 2 are
basal.

Inflorescences at the upper 3-7 nodes, apparently axillary and leaf-
opposed at the same node, or one cluster at a node, 1.5-3.0 em. long,
apparently sometimes also terminal, extending 1.7 cm. beyond the
first leaf, umbellate, 3-6 flowers per umbel, the peduncle 0.4-1.5
(-2.5, Schum.) cm. long, sometimes with a primary branch 0.5-1.1
cm. long and a secondary branch 0.5 cm. long, the pedicel 1.7-4.0
mm. long; bracts 1-1.7 mm. long, 0.2-0.5 mm. wide, lanceolate to
nearly linear, tomentulose, hairs stellate, 0.1 mm. long.

322 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Calyx 5.2 mm. long, 4.2 mm. wide, tomentulose-canescent, hairs
stellate, 0.1-0.2 mm. long, also a few glandular granules and glandular
hairs 0.1-0.2 mm. long, the teeth 3-3.5 mm. high, 1.5 mm. wide,
delioid-acuminate, the sinus acute to slightly rounded; petals white,
9-9.7 mm. long, 2 mm. wide, oblanceolate, the claw 3-4 mm. long,
united to the staminal tube for 2.7 mm.

Longistylous form. Stamens 5.5 mm. long, the filaments free for
1.5 mm., the anther 1.1 mm. long, 0.7 mm. wide, oblong, emarginate
at both ends; pistil 9 mm. long, the styles filiform, free for 5.5 mm.,
united for 1 mm., papillose for 0.8 mm. at the apex, the ovary pentag-
onal-ovoid, tomentulose, hairs stellate, 0.1-0.2 mm. long, the styles
with scattered hairs up to the stigmas.

Brevistylous form (according to Schumann). Stamens 8 mm.
long, the filaments filiform, free for 6 mm., the anther 1.5 mm. long,
deeply divided, divergent above; pistil 4 mm. long, the ovary setose
at the apex, the styles flat and free (in one specimen 6), the apex
clavate, slightly thickened, glabrous.

Fruit relatively large, 1-1.2 cm. long, 1.9-2.2 em. wide, chartaceous,
pentapterous, pyramidal, truncate at the apex or with a short rostrum
1 mm. long, the outer angle of wings basal, rounded and mucronate
for 0.2 mm., or acute, the sulci between the carpels extending half
way to the center of the fruit, stipe none, but the base of the capsule
stipelike for 0.5 mm., sometimes 6-7 carpels; dehiscence loculicidal
along the dorsal and ventral sutures; seeds (Schumann) 2.5 mm.
long, compressed, black, glabrous, chalaza large, cinereous-fuscous.

Tyre Locatiry: Bahia State, Brazil, near Barra da Vareda. Type
collected by Prinz zu Wied-Neuwied, number 2.

Distrispution: Bahia, Brazil, and Honda, Colombia (Schumann),
in forest and along roads.

BRAZIL: F neg. 9604 ex B (US). Banta: Maritiba, Blanchet 3472 (NY).

This species can be distinguished from other species by its rela-
tively large capsules that are chartaceous rather than somewhat
indurate and truncate or very shortly rostrate rather than long
rostraie. It is perhaps most closely related to Melochia betonicifolia
St.-Hil.

The collection of the specimen photographed as F 9604 ex B is
attributed to Poeppig, no. 2519, but Schumann (47) cited Poeppig
2519 as M. littoralis (Poepp. & Endl.) Schum. The photograph fits
the description of M. illicioides Schum., and the specimen was identi-
fied as such by Schumann. Since there is a number 2 pasted on the
specimen sheet, the specimen may have been collected by Prinz zu
Wied-Neuwied, number 2, which is the only Brazilian collection of
this species mentioned by Schumann (47).

GOLDBERG—GENUS MELOCHIA 323

46. Melochia betonicifolia St.-Hil. Fl. Bras. Mer. 1:165. 1825.
Melochia nepetoides St.-Hil. Fl. Bras. Mer. 1:165. 1825.
Melochia cordiformis St.-Hil. Fl. Bras. Mer. 1:166. 1825.

A branched, erect suffrutex to frutex, 0.45-4.0 meters high, the
young stems and petioles tan-canescent, tomentulose, hairs stellate,
0.1-1.0 mm. long, internodes 0.3-5.0 cm. long; stipules 1.7-8.0 mm.
long, 0.4-3.7 mm. wide, deltoid-acuminate or deltoid-acute or lan-
ceolate, tomentulose on the abaxial surface, also ciliate, hairs simple,
forked, and stellate, 0.1-1.2 mm. long; petiole 0.5-4.4 em. long.

Lamina 2.5-8.0 cm. long, 1.2—5.8 cm. wide, ovate to lanceolate, the
base rounded to cordate, the apex acute, the upper surface green,
hirsute-tomentulose, hairs stellate, forked, and simple, 0.1-0.5 (—0.8)
mm. long, the lower surface greenish or slightly to considerably
canescent, tomentulose, hairs stellate, 0.1-0.6 mm. long, velvety or
rough to the touch, the margin irregularly crenate-serrate, serrations
1.1-7.0 mm. wide, 0.5-2.6 mm. high, (5-) 7-10 pairs of primary
veins, 2 (1-3) are basal.

Inflorescences 1.3-4.7 cm. long, usually axillary at the upper 2-5
nodes, rarely also terminal and leaf-opposed, sometimes on short
axillary branches, the clusters in subcapitate umbels of 5-20 flowers,
the peduncles (0.2—) 0.8-2.9 em. long, 1-1.2 mm. wide, rarely with a
branch 0.2 cm. long, the pedicel 1-3.7 mm. long; bracts 1-4.5 mm.
long, 0.2-1.2 mm. wide, deltoid-acuminate or deltoid-acute to lan-
ceolate and linear, hirsute-tomentulose and ciliate, hairs simple,
forked, and stellate, 0.1-0.8 mm. long.

Calyx 6-8.3 mm. long, 4.5-6.0 mm. wide, canescent, densely tomen-
tulose, hairs stellate and sometimes also forked, 0.1-0.3 (-1.0) mm.
long, the teeth 3.3-5.3 mm. long, 1.5-2.4 mm. wide, deltoid to ovate,
acuminate or acute, the sinus acute to rounded; petals white,
yellow or purple, 7.5-12.5 mm. long, 2-4.5 mm. wide, oblanceolate- or
obovate-cuneate, attached to the staminal tube for 14.5 mm., the
claw 2-3 mm. long.

Longistylous form. Stamens 5.5-7.1 mm. long, the filaments free
for 0.7-1.5 mm., the anther 1.1—-1.4 mm. long, 0.5-0.7 mm. wide, oblong,
slightly emarginate at both ends; pistil 8.4-10.4 mm. long, the styles
filiform, free for 4.3-6.7 mm., united for 1-2 mm., papillose for 0.7-1
mm. at the apex, the ovary ellipsoid-ovoid, the styles and ovary
hirsute-tomentulose, hairs stellate, forked and simple, 0.1-0.6 mm.
long, sometimes 7 carpels present, the stipe 0.1 mm. long.

Brevistylous form. Stamens 6.5-10.2 mm. long, the filaments free
for 2.5-6.0 mm., the anther 0.8-1.4 mm. long, 0.6-0.8 mm. wide;
pistil 5-5.2 mm. long, the styles free for 2-4 mm., united for 1-2.8
mm., slightly clavate and stigmatic for 0.5-0.7 mm., the styles and
ovary tomentulose, hairs stellate, 0.2-1.0 mm. long.

324 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Fruit sometimes with 8 carpels, 6.7-8.5 mm. long, 2.5 mm. being
rostrum, 11.5-17 mm. wide, pyramidal, pentapterous, the points of
wings acuminate, tomentulose, hairs stellate, 0.2-1.0 mm. long, the
capsule narrowed at the base for 1 mm., the stipe 0.5 mm. long;
dehiscence loculicidal along the dorsal and ventral sutures; seeds
2.5-2.7 mm. long, 1.6-1.8 mm. wide, brown.

Flowering specimens collected from December to February, April
to June, and in August and September; fruiting ones, in February,
April, August, and September.

Type Locatity: The type was collected by St.-Hilaire near the
praediolum Bom Jardin, not far from the praesidio called S. Miguel
de Jiquitinhonha in Minas Gerais, Brazil, commonly in caatinga
woods where the leaves are shed every year. The isotype is cited
below.

DistrisutTion: Northern and eastern Brazil in caatinga woods in
which the leaves are shed during the hot and dry season, at low ele-
vations (50 meters).

BRAZIL: Crard: Four km. 8. of Mecejana, Cutler 8393 (US). Fortaleza,
Aldeota, Ducke 2377 (US). PrErNamBuco: Paquaretinga, Pickel 1504 (US).
Banta: Maracds, Ule 6967 (K, L). 8S. Thome, Blanchet 3791 (P). ?Bauta:
Coragao, Blanchet 3306 (NY, P). Minas Gerais: 8S. Miguel da Jequitinhonha,
St.-Hilaire cat. B, 1561 (P, isotype), also F neg. 35377 ex P (US). Near 8. Do-
mingos, St.-Hilaire cat. B, 1579 (P, type of Melochia cordiformis St.-Hil., also F
neg. 35378 ex P [US]). Rivulet Sucurid, Si.-Hilaire 1405 (P, isotype of Melochia
nepetoides St.-Hil., also F neg. 35384 ex P (US). Arassuahy near Rio Manso,

Glaziou 13557 (P), in part (R). Rio pE JANEIRO: Environs of Rio de Janeiro,
Glaziou 13557 (K). Serra da Estrella, Glaztou 12459 (R).

This species is similar to MZ. tomentosa L., but generally more robust;
the inflorescences generally contain more flowers, often white or
yellow; the pedicels are usually shorter. It is also related to M.
caracasana Jacq.; the inflorescence is usually leaf-opposed in M.
caracasana, but usually axillary in M. betonicifolia. The capsules are
generally wider in M. betonicifolia than in M. caracasana.

M. betonicifolia, M. nepetoides, and M. cordiformis were published
at the same time. In combining them, in accordance with the
International Code of Botanical Nomenclature (33), I select M.
betonicifolia as the correct name.

47. Melochia caracasana Jacq. Coll. 2:369. 1788.
Melochia macrophylla H.B.K. Nov. Gen. & Sp. 5:324. 1823.
Mougeotia caracasana (Jaeq.) H.B.K. Nov. Gen. & Sp. 5:329. 1823.
Riedleia Caracasana (Jacq.) DC. Prodr. 1:492. 1824.
Visenia caracasana (Jacq.) Spreng. Syst. 3:30. 1826.

An erect, branched shrub, sometimes herb, 0.45-5.0 meters high,

the young stems sordid canescent, densely tomentose, hairs stellate,
0.3-1.0 mm. long, internodes 1-9 cm. long; stipules 2-7 mm. long,

GOLDBERG—GENUS MELOCHIA 325

1.2-2.0 mm. wide, deltoid-acute to deltoid-acuminate, tomentose
and ciliate, hairs simple and forked, 0.2-1.0 mm. long; petiole 0.5-
5.1 cm. long.

Lamina 1.4-12.7 cm. long, 0.8-7.7 cm. wide, lanceolate-ovate to
ovate, the base rounded to cordate, the apex usually acute, the
upper surface green, sparsely to densely puberulous, hairs stellate,
forked and simple, 0.4 (0.2-0.9) mm. long, the lower surface yellowish-
tan-canescent, tomentose, hairs stellate, 0.3-0.8 mm. long, both
surfaces woolly to the touch, the margin irregularly crenate to crenate-
serrate, crenations or serrations 2-5 mm. wide, 0.7-2.0 mm. high,
5-10 pairs of primary veins, 2 are basal.

Inflorescences 1-3 cm. long, leaf-opposed, axillary, or both, at the
upper 2-6 nodes, apparently sometimes also terminal, the clusters in
rather dense subcapitate umbels of 3-20 flowers, the flowers fragrant,
the peduncle thick, 0.4-2 cm. long, 0.8-1.4 mm. wide, sometimes
with a branch 0.3-0.5 em. long, the pedicel 1-4 mm. long; bracts
1.8-6.0 mm. long, 0.2-1.0 mm. wide, deltoid-acute to deltoid-
acuminate, or linear, tomentose.

Calyx 5-8 mm. long, 3.5-6.0 mm. wide, sordid canescent and
tomentulose, hairs stellate, 0.3 (-2.0) mm. long, the teeth 2.5-6.0
mm. long, 0.8-2.0 mm. wide, deltoid-acute to deltoid-acuminate,
sometimes with rounded sides, the sinus usually acute; petals white
to yellow and brownish, 6-10 mm. long, 1.5-2.6 mm. wide, oblanceo-
late to obovate, spatulate, attached to the staminal tube for 1.5-1.8
mm., the claw 1 mm. long.

Longistylous form. Stamens 5.0-6.5 mm. long, the filaments
free for 1-3 mm., the anther 1.4-1.7 mm. long, 0.5-1.0 mm. wide,
oblong, emarginate at both ends; pistil 7.2-10 mm. long, the styles
filiform, free for 4.5-6.0 mm., united for 1.6 mm., papillose and slightly
clavate for 0.5-1.0 mm. at the apex, the ovary ovoid to ellipsoid,
slightly pentagonal, hirsute at the apex, hairs stellate, forked and
simple, 0.2-0.7 mm. long, the stipe 0-0.8 mm. long.

Brevistylous form. Stamens 7-8.5 mm. long, the filaments free
for 4-5.5 mm., the anther 0.8-1.6 mm. long, 0.4-0.7 mm. wide, ellip-
sold, ovoid, or oblong and emarginate at both ends; pistil 4-6.5 mm.
long, the styles 1.5-2 mm. long, free to the ovary.

Fruit 7-10 mm. long, 2 mm. is rostrum, 6-11 mm. wide, pyramidal,
pentapterous, the points of wings usually basal, usually recurved and
acute to acuminate, rarely blunt, usually canescent, densely pubescent,
hairs stellate, up to 0.5 mm. long, the stipe 0-1 mm. long; dehiscence
loculicidal along the dorsal suture; seeds 2.2-2.7 mm. long, 1.1-1.5
mm. wide, trigonous, brown, sometimes striate.

Flowering and fruiting specimens collected in May, June, and
August to December.

326 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Type Locauity: The type was collected by Jacquin near Caracas,
Venezuela, and is cited below.

Distripution: Northern Venezuela, Colombia and Brazil, in
thickets, on open, dry, rocky hillside, in ravines, or in savannas, at
0-1,700 meters elevation.

VENEZUELA: Sucre: F neg. 35383 ex P of type of Melochia macrophylla
H.B.K. (US). Muiranpa: Petare and vicinity, near Caracas, Pittver 7146 (US).
FrepEeraL District: Caracas, Jacquin (W, type); van Landsberge 253 (8); Pittier
6149 (US). El Valle, Fernandez 154 (US). On road to La Guaira from Caracas,
between Catia and Las Trincheras, EF. Pittier 12 (US). Lower Cotiza, Pittier
7178 (US). Turmerito, Ingomar, Killip 37728 (US). El Paraiso, Caracas,
Lasser 744 (US). Between Caracas and La Guaira, Mr. & Mrs. J. N. Rose
21820 (US). On the old road from Caracas to La Guaira, Pittter 9545 (US).
La Guaira, André 109 (NY). ?FeEpERAL Disrricr: Valle, van Landsberge Nov.
1847 (S). El Valle and Laguna de Espino, Eggers 13130 (L, US). Aragua:
Maracay, Vogel 340 (8S). Colonia Tovar, Fendler 109 (P, US). Carasono:
Hacienda Taborda, near KE] Palito, on the road from Valencia to Puerto Cabello,
Pittier 7666 (US). Puerto Cabello, Valle Seco, Engstedt 3 (S). El Palito,
Valencia, Warming 264 (US). Lara: Near Barquisimeto, Saer 26 (US).
Méripa: Mocquerys 1234 (US).

COLOMBIA: MaapaLena: Santa Marta, Dawe 500 (US). SANTANDER:
Rio Suraté valley, near Bucaramanga, Killip & Smith 16226 (US).

BRAZIL: Creard: Alleméo & de Cysnetros 134 (R, US).

This species is similar to M. tomentosa L., but generally larger and
more robust; the inflorescences generally contain more flowers, white
or yellow rather than purple; the peduncles are thicker; the pedicels
are usually shorter; the leaves have longer hairs. It is also similar
to M. betonicifolia St.-Hil., but the inflorescence is most commonly
axillary in M. betonicifolia and usually leaf-opposed in M. caracasana.
The leaves of M. caracasana are usually hairier than those of M.
betonicifolia and the hairs are yellowish rather than white.

It is illustrated by Jacquin (32).

48. Melochia lanata St.-Hil. Fl. Bras. Mer. 1:167. 1825.
Melochia lanata St.-Hil. var. inclusa St.-Hil. Fl. Bras. Mer. 1:167. 1825.

An erect, branched suffrutex, at least 0.4 meter high, the young
stems and petioles densely lanate, canescent, hairs stellate and
apparently forked and simple, 0.2—-1.6 mm. long, internodes 2-7 cm.
long; stipules 4.1-7.0 mm. long, 0.2—0.5 mm. wide, lanceolate to sub-
linear, mostly ciliate, some hairs on the abaxial surface, most simple,
some forked, 0.4—-2.0 mm. long; petiole 0.5-1.5 em. long.

Lamina 2-5.5 cm. long, 1-3.4 cm. wide, ovate, ovate-lanceolate,
lanceolate, oval or elliptic-oblong, the base rounded, the apex rounded
or acute, the upper surface sericeous-tomentose, lanate, hairs yellowish-
tan-canescent, simple, appressed, 0.5-2.0 mm. long, the lower surface
densely lanate, yellowish- to tan-canescent, hairs stellate, forked, and
simple, 0.7—1.6 mm. long, the margin irregularly and mostly shallowly

GOLDBERG—GENUS MELOCHIA 327

crenate-serrate, serrations 0.8-3.9 mm. wide, 0.3-1.3 mm. high,
7-9 pairs of primary veins, 2 are basal.

Inflorescences 2-5 cm. long, leaf-opposed, 90° from the leaf, or
axillary, 3-15 flowers in dense subcapitate, umbellate clusters at
3-6 upper leaf nodes, 1 or 2 clusters per peduncle, the peduncle 0-3
cm. long, 1.2 mm. wide, the pedicel 2-4 mm. long; bracts 2.5-6.5
mm. long, 0.2-0.3 mm. wide, sublinear to linear, lanate-tomentose
and ciliate, hairs mostly simple, few forked, 0.3-1.8 mm. long.

Calyx 6.8-9.2 mm. long, 4.7-5.2 mm. wide, lanate-tomentose on the
outside, hairs stellate, forked and simple, 0.2-1.5 mm. long, the teeth
3.8-5.8 mm. long, 2-2.2 mm. wide, ovate- to deltoid-acuminate, the
sinus acute; petals 8.6-14 mm. long, 2.2-2.5 mm. wide, oblanceolate,
with scattered glandular hairs 0.05 mm. long, attached to the staminal
tube for 3.5-3.7 mm., the claw 1.5-2.5 mm. long.

Longistylous form not seen.

Brevistylous form. Stamens 7.7-10.8 mm. long, the filaments free
for 5-6.5 mm., the anther 1.2-1.7 mm. long, 0.6-0.8 mm. wide, oblong,
slightly emarginate at both ends; pistil 5.2-5.5 mm. long, the styles
filiform, stigmatic for 0.8 mm., free for 2-2.2 mm., united for 1-1.2
mm., the ovary ovoid to ellipsoid, the upper half of the ovary and
base of the styles tomentose-sericeous, hairs forked and stellate,
0.2-1.3 mm. long, stipe none.

Fruit not seen.

Flowering specimens collected from September to November.

TypE Locauity: Sertaéo near Olho d’Agua, Minas Gerais, Brazil,
isotype cited below.

DistriputTion: In herbaceous fields in the desert part of Minas
Gerais, Brazil.

BRAZIL: Minas Gerais: Near Olho d’Agua, St.-Hilaire (P, isotype, also
US neg. 5661); F neg. 35382 ex P of type or isotype. Rico Pares?, Vincent 818
(L).

The position of the inflorescence, often leaf-opposed, and the
relatively long attachment of the petals to the staminal tube indicate
that this species belongs to the section Pyramis. M. lanata var.
inclusa St.-Hil. appears to be the longistylous form of the species.
The species appears to be closest to M. caracasana Jacq., but differs
considerably from it in vestiture.

49. Melochia tomentosa L. Syst. ed. 10, 1140. 1759.

Usually a shrub or subshrub, rarely an herb, 0.3-4.0 meters high,
slender, erect or spreading, usually branched, sometimes several
branches from the base, perennial, sometimes flowering the first year,
the root sometimes thicker than the base of the stem, the bark of the
stem blackish or purplish, the young stems and petioles tomentulose,

328 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

usually canescent, hairs stellate, 0.1-0.2 mm. long, internodes 0.8-5.5
cm. long; stipules sometimes deciduous, 2-9 mm. long, 0.4-1.5 mm.
wide, deltoid-acuminate or deltoid-acute to nearly linear, densely
tomentulose, sometimes also ciliate, hairs stellate, sometimes also
simple and forked, 0.1-0.3 (—0.7) mm. long; petiole 0.3-2.0 (-3.0) cm.
long.

Lamina (1.2-) 2-6 (—10) em. long, 0.7-4.0 (—5.0) em. wide, ovate to
lanceolate, rarely suborbicular, the base rounded, subcordate, rarely
also subtruncate or subcuneate, the apex acute to rounded, both
surfaces usually tomentulose, sometimes hirtellous, usually canescent,
sometimes only the lower, or neither surface canescent, hairs stellate,
0.1-0.5 mm. long, the margin irregularly crenate or crenate-serrate,
crenations or serrations 0.7—5.0 mm. wide, 0.3-2.0 mm. high, (5-) 6-9
(-11) pairs of primary veins, usually 2 are basal, the veins impressed
above, prominent beneath.

Inflorescences usually axillary, 1.5-5.0 em. long, sometimes also
leaf-opposed, 2~3 cm. long, or terminal, extending 3-3.2 cm. beyond
the first leaf, umbellate, 2-10 (-20) flowers per umbel, at (1—-) 3-9
upper nodes, sometimes at 2-3 naked nodes, the peduncle 0.3-2.5
(-5.0) em. long, 0.5-0.6 mm. wide, rarely with a branch 0.5 em. long,
rarely two in an axil, the pedicel 2-5.7 mm. long; bracts 1-6.1 mm.
long, 0.1-0.7 mm. wide, shape and pubescence like those of the
stipules.

Calyx green, rarely dull red distally, 4.9-8.6 mm. long, 3.2-7.0 mm.
wide, densely tomentulose, canescent, hairs stellate, sometimes
also glandular, 0.1-0.3 mm. long, the teeth 2.8-5.8 mm. long, 0.7-
2.2 mm. wide, deltoid or ovate, acute or acuminate, the sinus
acute to rounded; petals purple, pink, violet, blue, sometimes paler
proximally, rarely red or white, 8-13.5 (-17) mm. long, 2.0-3.8
(-6.4) mm. wide, oblanceolate- to obovate-cuneate and spatulate,
attached to the stamens for 2.8-5.4 mm., the claw 1.7-3.0 mm.
long, sometimes united for 2 mm.

Longistylous form. Stamens 4-6.5 (-9.0) mm. long, the filaments
free for 1.8-2.7 mm., the anther 1.1-1.2 mm. long, 0.6—-0.8 mm. wide,
oblong, emarginate at both ends; pistil 7-10.9 (-12) mm. long, the
styles free for 3.2-6.0 mm., united for 3 mm., papillose and slightly
clavate for 0.7-1.5 mm. at the apex, the ovary and usually the base
of the styles tomentulose, hairs stellate, 0.1-0.4 mm. long, the ovary
ellipsoid to ovoid, the stipe 0-1 mm. long.

Brevistylous form. Stamens 7-12.2 mm. long, the filaments free
for 2.5-6.3 mm., the anther 0.9-1.5 mm. long, 0.5-0.9 mm. wide;
pistil 4.5-5.5 (-8.0) mm. long, the styles free for 1.5-2.6 mm., united
for 1.5-3.0 mm., papillose for 0.4-1.0 mm. at the apex, the ovary

GOLDBERG—GENUS MELOCHIA 329

and base of styles densely tomentulose, hairs stellate, 0.1-0.6 mm.
long, the ovary ellipsoid, the stipe 0-2 mm. long.

Fruit 6.5-10.5 mm. long, 6.6-10.5 mm. wide, long-rostrate, the
rostrum 1.3-5.0 mm. long, pyramidal, sometimes suboblong, pentap-
terous, the lateral points of wings at or below the middle of the fruit,
rather short, acute or rounded, the sulci extending a third to half
way to the center of the fruit, the base of the fruit narrowed into a
stipe 1-1.5 mm. long, tomentulose, hairs stellate, forked, 0.1-0.6
mm. long; dehiscence loculicidal all along the dorsal and ventral
sutures, eventually also septicidal; seeds 2-3 mm. long, 1.1-2.0 mm.
wide, obovate, red-, purple-, chocolate-, or light-brown, sometimes
longitudinally striate, sometimes canescent at the ends and along the
raphe, 1 or 2 per cell.

Flowering and fruiting specimens collected throughout the year.

Key to Varieties of M. tomentosa

1. Lamina green on both surfaces, hispidulous; petals often large, sometimes
15-17 mm. long; plant very slender; inflorescences at few nodes.
49d. M. tomentosa var. speciosa
1. Lamina canescent and tomentulose at least on the lower surface; petals up
to 13.5 mm. long; plant usually robust; inflorescences often at many nodes.
2. Upper surface of lamina green, lower canescent, serrations relatively few
and broad. . . . . . . « . . .49c. M. tomentosa var. turpiniana
2. Upper and lower surfaces of lamina canescent, serrations relatively numerous
and narrow.
3. Leaves medium-sized, lamina typically 2-6 cm. long, 0.7-4.0 cm. wide,
petiole often more than 1 cm. long.
49a. M. tomentosa var. tomentosa
3. Leaves small, lamina typically 1-3 em. long, up to 2 cm. wide,
petiole less than 1 em. long; stems much more branched and canescent
for a shorter distance from the apex than in the typical variety.
49b. M. tomentosa var. frutescens
49a. Melochia tomentosa L. var. tomentosa
Melochia crenata Bert. ex. Spreng. Syst. 3:29. 1826; not Vahl 1794.
Melochia portoricensis Spreng. Syst. 3:29. 1826.
Melochia plicata Presl, Rel. Haenk. 2:145. 1835.
Melochia hypoleuca Mig. Linnaea 22:468. 1849.
Ridleta hypoleuca (Miq.) Walp. Ann. 2:166. 1852.
Melochia tomentosa L. var. typica Schum. in Mart. Fl. Bras. 12(8):34. 1886.
?Melochia tomentosa L. var. Bahiensis Schum. in Mart. Fl. Bras. 12(8) :34.
1886.
Melochia arida Rose, Contr. U.S. Nat. Herb. 8:321. 1905.
Moluchia tomentosa (L.) Britt. Brooklyn Bot. Gard. Mem. 1:69. 1918.
Sida rajoides M. E. Jones, Extracts from Contr. West. Bot. 18:58. 1933.

The leaves of the typical variety are tomentose and canescent
on both surfaces. The lamina is typically 2-6 cm. long and 0.7-4.0
cm. wide.

330 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

TYPE LOCALITY: Jamaica. There are two specimens of MM. tomen-
tosa in the Linnaean herbarium. One appears to be M. tomentosa
var. tomentosa and the other Af. tomentosa var. frutescens. Localities
of collections are not given.

Distripution: Southern Florida, southern Texas to Nicaragua,
West Indies, Venezuela, Colombia, Surinam, and _ northeastern
Brazil to Bahia, usually in dry, open places, on rocky limestone hills,
in coastal thorn thickets, savanna, pineland, along the roadside, in
granite sand, in loam, on volcanic slopes, at 0-1,250 meters elevation,
usually below 500 meters.

UNITED STATES: Frioripa: pape: Buena Vista, Small 7268 (8, US). Texas:
HIDALGO: Mission, Shiller 644 (US).

MEXICO:58. Augustin, Liebmann 11880 (US). Lower Catirornia: Cerralvo,
Rose 16889 (US). Loreto, above Primiera Agua, Jones 27167 (US), 27170 (US).
Aguaje de San Esteban 25 mi. N. of San Ignacio, Nelson & Goldman 7205 (US).
About 5-10 mi. W. of San Ignacio, Wiggins 5443 (MICH, US); Thomas 8330
(US). About 26-27 km. NW. of San Ignacio, Carter et al. 1967 (US), 2519 (US).
Forty mi. E. of San Ignacio, Shreve 7056 (MICH). From Agua Colorado to Cerro
Colorado, Nelson & Goldman 7321 (US). Carmen Island, EH. Palmer 823 in
1890 (US). Isla Partida, Collins et al. 150 (US). North of Comondu, Wiggins
5476 (MICH, US). Forty mi. NE. of Pozo Aleman, Wiggins 7817 (US). North-
east of Pozo Aleman, Shreve 6984 (MICH). Two mi. E. of Pozo Aleman, Epling
& Robinson s.n. (MICH). About 28 mi. 8. of Pozo Aleman, Wiggins 7876 (US).
San Jose de Magdalena, Wiggins 11381 (US). Magdalena Island, Brandegee s.n.
(US); Orcutt 46 (US). San Nicholas Bay, Johnston 3734 (US). Puerto Escon-
dido, Johnston 3853 (US). Ceralbo Island, Gordas Point, Johnston 4033 (US).
La Paz and vicinity, Collins et al. 56 (US), 60 (US); HE. Palmer 121 in 1890 (US);
Hammerly 208 (US); Shreve 7201 (MICH). Between La Paz and Santo Domingo,
Thomas 8452 (US). Between Santonio and Puerto de Bahia de los Muertos,
Wiggins 5637 (MICH, US). Todos Santos, Jones 24108 (MICH). San José del
Cabo and vicinity, Brandegee s.n. (US); Thomas 7761 (US); Purpus 318 in part
(US); Carter et al. 2194 (US); Dawson 1191 (MICH). Cape San Lucas, Pose
16360 (US); Xantus 13 (US). Sonora: Guaymas, Rose et al. 15015 (GH, US);
Rose 1205 (US, type of Melochia arida Rose); Drouet & Richards 4029 (MICH);
Gentry 4714 (MICH). Bahia San Carlos, Dawson 1062 (MICH). TamMautipas:
Viereck 641 (US). Matamoros, Runyon 423 (US). Buena Vista Hda., Wooton
s.n. (US). San Miguel La Tamaulipeca, Bartlett 10585 (MICH), 10679 (MICH,
US). El Limon, Kenoyer & Crum 3637 (MICH). Victoria, Leavenworth 240
(GH). Sinavoa: Vicinity of Culiacan, Ortega 6691 (US). Altata, Gentry, 5437
(MICH). Durango: Rose 2272 (US). Zacatecas: San Juan Capistrano, Rose
2413 (US). San Luis Porosf: Las Palmas, Rose & Hough 4880 (US). Valles,
Kenoyer s.n. (MICH). NAyARIT: TRES MARIAS ISLANDS: Maria Madre Island,
Nelson 4205 (US); Tepic, Cafion de Jesus Maria, Goldsmith 145 (GH). VERA
Cruz: Zacuapan, Barranea de Santa Maria, Purpus 2357 (US); Rozynski 608 (R).
Banos del Carrizal, Purpus 6134 (US). Pursua: San Luis Tultitlanapa near
Caxaca, Rio de Santa Lucia, Purpus 2612 (US). Micnoacdn: APATZINGAN!
Tiquiluea, Hinton 12121 (US). Cotima: EH. Palmer 956 in 1891 (MICH, US).
GvuERRERO: Acapulco and vicinity, Bailey 514 (MICH); Haenke (PR 174399a, b,
c, type of Melochia plicata Presl, also US neg. 5550, W, isotype); H. Palmer 258
in 1894-95 (MICH, US). Oaxaca: Near Tomellin, Rose et al. 10083 (US).

GOLDBERG—GENUS MELOCHIA 331

Tomellin Canyon, J. N. & J. 8S. Rose 11330 (US). Estacion Almoloyas, Cafion
de Tomellin, Conzatti 1685 (US), 2034 (MICH). Playa de Coyula, Reko 3664
(US). ‘“Cancatlan’” (?Cuicatlin), Rusby 64 (US). Vicinity of San Juan Guichi-
covi, Nelson 2716 (US). Ixtepec, Fisher 35467 (US). San Geronimo, Mell 2111
(US), 2147 (MICH). Cuicatlin, Gonzalez 978 (US). Estacion de Quiotepec,
Conzattt 4588 (US). Salina Cruz and vicinity, King 209 (MICH); Deam s.n.
(MICH). Tehuantepec and vicinity, Matuda 923 (MICH); King 862 (MICH),
441 (MICH), 346 (MICH). About 5.5 km. NE. of Juchitan, King 364 (MICH).
CamprecHE: Konchen, Lundell 1388 (MICH). Yucardn: Balantun, Gaumer
23267 (US). Chichankanab, Gaumer 1799 (US). Mérida, Collins 13 (US);
Souza Novelo 259 (US). Progreso, Steere 3029 (MICH). Muna, Steere 2136
(MICH). QuintTana Roo: Coba on bank of aguada W. of Lake Coba, C. L. &
A. A, Lundell 7670 (MICH, US).

HONDURAS: Cortss: Rio Lindo, Edwards P-698 (US).

NICARAGUA: Managua: Managua and vicinity, Garnier 1168 (MICH, US);
Chaves 72 (US). Granapa: Granada, Baker 213 (MICH).

BAHAMAS: Eleuthera, Harbor Island, EH. G. Britton 6387 (US). New
PROVIDENCE: Near Nassau, Curtiss 12 (L, US). ANpros: Coppice, Smith Hill,
Long Bay Cays, Small & Carter 8666 (US). Exuma Cuain: Great Guana Cay,
Britton & Millspaugh 2877 (US).

CUBA: Camactey: Vicinity of Pueblo Romano Cayo Romano, Shafer 2437
(US). Six mi. from NE. end of Cayo Coco, Shafer 2728 (US). Paloma, Cayo
Sabinal, Shafer 816 (US). Cayo Guajaba, Shafer 640 (US). OnzentE: North-
ward of Raimon, Nipe Bay, Shafer 1784 (US). Santiago and vicinity, Taylor 17
(US). Lopez Figueiras 189 (US). Siboney, Morton 3970 (US). Yateritas, Via
Azul, Bro. Alain & Morton 4949 (US). Manigua costera, en el Socucho, Playa
de Puerto Padre, Lopéz F 2906 (US).

HAITI: Port-au-Prince and vicinity, Leonard 2816 (US); Bailey Haiti 132 (US),
Haiti 283 (US). Torrur Isuanp: Vicinity of La Vallée, FE. C. & G. M. Leonard
11647 (MICH, US), 15406 (US). Gonave IsLANp: Pte. & Raquette, Eyerdam
223 (US). Vicinity of Etroite and Anse Galette, Leonard 3112 (US), 3351 (US).
Gonaives and vicinity, Nash & Taylor 1546 (US); Buch 283 (L, 8S); Leonard 10061
(US). Vicinity of St. Raphael, Leonard 7692 (US). Vicinity of Jean Rabel,
E. C. & G. M. Leonard 13829 (US). Vicinity of Mole St. Nicholas, H.C. & G. M.
Leonard 13210 (US). Vicinity of Cabaret, EH. C. & G. M. Leonard 12016 (US).
Vicinity of Fond Parisien, Etang Saumatre, Leonard 4117 (US), 4132 (US).
Vicinity of St. Mare, Leonard 2876 (US).

DOMINICAN REPUBLIC: C. Wright et al. 45 (US), 108 (US). Azua, Rose
et al. 4024 (US). Sanriaco: Guayacanal, Jiménez 2263 (US). Monre CristT!1:
Guayubin, Abbott 941 (US). Baranona: Fuertes 382 (US).

JAMAICA: Kingston and vicinity, Killip 11 (US); Mazon & Killip 1385 (US),
308 (US). Near Falmouth, W. Harris 7222 (US). Near Alligator Pond, W.
Harris 7230 (US). Watsons Hill, W. Harris 6315 (US). Long Mountain Road,
Fawcett 7313 (US). East slope of Hellshire Hills, Proctor 7502 (US). Port
Henderson, J. A. Harris & Lawrence C1537 (US). Granp CayMAN ISLAND:
Proctor 15054 (IJ).

PUERTO RICO: Cayo Muertos, Britton et al. 4971 (US). San Ildefonzo,
Goll et al. 701 (US). Coamo, Sintenis 3150 (US).

LESSER ANTILLES: U.S. Viren Isuanps: st. croix: Bassin, Ricksecker
127 (US). sr. Jonn: Near Lamosure, Britton & Shafer 644 (US). MarrTiniquBE:
Duss 1360 (US). H. & M. Stehlé 6861 in part (US). Sieber 321 (L). Case
Pilote, Hahn 248 (GH, L, US), 240 (R). Grenapa: St. George, Broadway in
1896 (US).

332 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

DUTCH WEST INDIES: Curagao: Various localities, Suringar, Jan. 1885
(L). Near Willemstad, Britton & Shafer 2913 (US). Bonaire: Suringar Feb.
20-23, 1885 (L). Rincon, Boldingh 7010 (L). Arusa: Koolwijk, Suringar Jan.-
Feb, 1885 (L).

TRINIDAD: Patos Island, Broadway 2716 (S).

VENEZUELA: Valley of Puerto La Cruz, Hacienda Panarigua, Pittier 9199
(US). Nueva EsparTa: MARGARITA ISLAND: El Valle, Miller & J. O. Johnson
44 (US). Sucre: Vicinity of Cristébal Colén, Broadway 133 (US). Frprrau
District: Around Caracas, Pittier 6170 (US). Lower Cotiza, near Caracas,
Pittier 7257 (US). Vicinity of Cabo Blanco, Pittier 10274 (US). La Guaira,
Curran & Harman 846 (US), 876 (US). Catia de la Mar, Tamayo 399 (US);
Vogl 617 (8S). Araaua: Colonia Tovar, Fendler 107 (US). Carasoso: El Palito,
on road from Valencia to Puerto Cabello, Pitter 7668 (US). Fatc6n: Paraguans,
Pueblo Nuevo, Tamayo 1033 (US). Mé&rripa: Brecenio Hacienda 23 mi. SE. of
Mérida, Reed 547 (US). Vallée du Rio Chama, en aval d’Egido, Humbert 26537
(P, US). Boxfivar: Ciudad Bolivar, Holt & Gehriger 85 (US), 100 (US); L. H. &
E. Z. Bailey 1508 (US).

COLOMBIA: Magpa.ena: Santa Marta, Curran s.n. (US). Near Fonseea,
Haught 4311 (US). Cerrején, Haught 6639 (US). ArLAntico: Barranquilla,
Bro. Elias 134 (US). Botivar: Vicinity of Cartegena, Bro. Heriberto 9 (US).
Norte DE SANTANDER: Vicinity of Cicuta, Killip & Smith 20981 (US); Molina &
Barkley 18 N. 8. 011 (US).

BRAZIL: Pravf: Sete Lagoas, Liitzelburg 1476 (RB). Crear: Motuquinha,
Léfgren 460 (R, S$). Paraisa: de Moraes 804 (US). PeRNAmBuco: Petrolandia,
Kuhlmanns.n. (RB). Alagéa de Baixo, Monteiro 238 b (R). Caruart (Carapotoz),
Pickel 3511 (US). Auagoas: Gardner 1258 (5S, US). Banta: Jacobina, Schreiner
s.n. (R). Joazeiro, Léfgren 961 (RB). Joazeiro, Horto Florestal, Zehntner 148
(R). Frageiro, Rio 8. Francisco, Zehniner 768 (M). Queimadas, Rose &
Russell 19836 (US).

This species is related to Melochia pyramidata L. The fruit of the
latter is not as hairy, and is frequently purple blotched. Also, the
fruit of MZ. tomentosa has a longer rostrum and the lateral angle of the
wings is acute or rounded, but is acute or acuminate in M. pyramidata.
Melochia tomentosa is usually a taller and hairier plant than M.
pyramidata. The leaves of M. pyramidata are usually subglabrous
and usually have some simple hairs on the upper surface, those of AZ.
tomentosa are usually tomentose and canescent and do not have
simple hairs. The inflorescences of M. pyramidata are usually leaf-
opposed, those of AM. tomentosa are usually axillary.

The flowers open in the morning and close in the afternoon (Run-
yon), and are very fragrant (MZ. arida Rose) (Rose).

The varieties of M. tomentosa grade into one another.
49b. Melochia tomentosa L. var. frutescens (Jacq.) DC. Prodr. 1:490. 1824.

Melochia frutescens Jacq. Obs. Bot. 2:22, pl. 44. 1767.
Moluchia frutescens (Jacq.) Medic. Malv, 10, 1787.

The leaves of this variety are smaller than those of the typical
variety. The petiole is less than 1 cm. long, and the lamina is typi-

GOLDBERG—GENUS MELOCHIA 333

cally 1-3 cm. long and less than 1 cm. to 2 cm. wide. The stem
branches more than that of the typical variety.

TYPE LOCALITY: Saint Martin, West Indies.

Distribution: Mexico, West Indies, Venezuela, and Colombia.

MEXICO: Lower Catrrornia: San Francisquito Bay, Rose 16733 (US);
Johnston 3568 (US). Espiritu Santo Island, the Isthmus, Johnston 3962 (US).
NAYARIT: TRES MARIAS ISLANDS: Maria Madre Island, Maltby 57 (US); Solis 12
(US). Oaxaca: Puerto Angel, Morton & Makrinius 2641 in part (US). Teotit-
lin del Camino, Seler 809 (US).

BAHAMAS: ELeutTHera: Rock Sound, Krebs s.n. (8). ?Siklins Island, Eggers
s.n. (S). ANpRos: Mangrove Bay, Brace 4907 (US).

TURKS and CAICOS: Proctor 8780 (IJ), 9177 (IJ).

CUBA: C. Wright 40 (S, US). Camaairy: Pastelillo, Cayo Sabinal, Ekman
15413 (S), 15536 (8S). Orrenre: Santiago de Cuba W. of Playa Siboney, Ekman
7748 (S), 8730 (S). Guanténamo at Caimanera, Ekman, 2866 (8S). Guantdnamo
Bay, Britton 1948 (US). Vicinity of El Cobre, Pollard & W. Palmer 384 (US).
Berraco, vicinity of Daiquiri, Britton & Cowell 12672 (US); Mazon 4028 (US).

HAITI: Port de Paix, Ekman 3837 (8S). Vicinity of St. Michel de l’Atalaye,
Leonard 7327 (US), 7335 (S, US). Vicinity of Ennery, Leonard 10025 (US).
Gonaives and vicinity, Leonard 10067 (US). Vicinity of Cabaret, HE. C. & G. M.
Leonard 11874 (US), 11936 (US).

DOMINICAN REPUBLIC: Near Manuel de Ocoa, Tiirckheim 3690 (L, 8,
US). Moncién, Valeur 158 (S, US). Gurabito, Eggers 1991 (US). Monrs
Cristi: Near Rio Yaque del Norte, R. A. & E. S. Howard 9608 (US), 9619
(US). Baranona: Trujin, Abbott 1726 (US). La Salina, Rk. A. & EF. 8S. Howard
8364 (US). Beata Island, Howard 12498 (S, US); Jiménez 3772 (US).

JAMAICA: Kingston, Orcutt 4956 (S).

PUERTO RICO: Stahl 1027 (L,8). Wydler 26 (8S). Cabo Rojo, Las Salinas,
Otero M-33 (MICH); Sintenis 628 (L, 8S, US). Morrillos de Cabo Rojo, Britton
et al. 4715 (US). Ponce and vicinity, Underwood & Griggs 698 (US); Heller 482
(US), 6064 (L, US). Coamo Springs, Underwood & Griggs 562 (US). Guanica,
Sintenis 3499 (US). Brigadier Point to Puerto Negro, Shafer 2921 (US). ?La
Paguera, Sargent 155 (US). ?Tajarto, Sintenis 1602 (US).

LESSER ANTILLES: U.S. Virgin Isuanps: st. crorx: Benker s.n. (8).
Salt pond, Raunkiaer s.n. (8S). Hansen 109 (L). Frederiksted, Rose et al. 3525
(US), 3226 (US). str. THOMAS: O6ersted collected 1845-48 (MICH, 58, US).
Schomburgk 42 (W). Eggers 49 (US), 96 (MICH), 194 (S), 262 (L). Wydler
s.n. (L). Water Island, Britton et al. 108 (US). st. soun: Breutel s.n. (L).
BriTIsH VIRGIN IsLANDs: Tortola, Fishlock 43 (US). St. Martin: Suringar
May 1, 1885 (L). Sr. Bartruétemy: Forsstrém s.n. (8), Euphrasén s.n. (8),
von Goes s.n. (S). St. Jean, Stehlé 911 (US). Sv. Eusratius: Suringar Apr. 8,
1885 (L). Sr. Kirrs: Canada Estate, Britton & Cowell 740 (US). ANTIGUA:
Slopes and summit of Boggy Peak, Shekerley Mts., A.C. Smith 10421 (S, US).
Near English Harbor, Rose et al. 3357 (US). Lyons, Bor 997 dupl. (US). Monr-
SERRAT: Shafer 418 (US). GuapELouPE: Duss 480 (US), 2901 (US). GreENna-
DINES: West coast of Mustique, A. C. Smith 10163 (S, US).

CURACAO: Killip & Smith 21056 (US). Curran & Haman 26 (US).

TRINIDAD: Patos Island, Britton et al. 526 (US).

VENEZUELA: Northern Venezuela, Curran & Haman 876 (US). Nurva
Esparta: Robledar, Gines 3191 (US), 3871 (US). El Valle, Gines 4126 (US).
R. Matasiete, Gines 2759 (US), 3195 (US). Salamanca, Gines 3647 (US). Cer-

227-051—67——10

334 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

rado, Gines 3768 (US). Sucre: Valley of Bordones SW. of Cumand, Steyermark
62891 (US). Lara: Barquisimeto, Tamayo 2616 (US). Carora, Jahn 184
(US). Trusttto: La Conception, Reed 1011 (US). Meriva: El Molino,
just above Lagunillas, Steyermark 56222 (US). Bottvar: Along Rfo Caronf,
near Puerto Ordaz, Steyermark 94224 (US).

COLOMBIA: Magpatena: Near Codazzi, about 90 km. NE. of Chiriguan4,
Haught 2306 (MICH, US), 3734 (US).

49c. Melochia tomentosa L. var. turpiniana (H.B.K.) Schum. in Mart. Fl. Bras.
12(3) :34. 1886.
Melochia Turpiniana H.B.K. Nov. Gen. & Sp. 5:323, pl. 482. 1823.

The leaves of this variety are about the size of those of the typical
variety, but differ in that the upper surface is green rather than ca-
nescent and the serrations are broader and fewer.

Tyre tocauiry: ?Colombia. The type was collected by Hum-
boldt and Bonpland who were not certain of the locality.

DistriputTion: Mexico, West Indies, Venezuela, and Colombia.

MEXICO: Mexico und Umgebung, Wawra 394 (W). Jauisco: Tequila,
Herrerén Nov. 1902 (8). Corcovado Canyon, NE. of Autlin, toward Guadalajara,
Rk. LL. & C. k. Wilbur 2378 (MICH, US). Micuoacdn: Zitacuaro-Tuzantla,
Hinton 13064 (US). Between San Juan de los Plitanos and Amatlin, McVaugh,
17956 (MICH, US). Cotrma: Manzanillo and vicinity, Ferris 6150 (US).
Oaxaca: Tomellin Canyon, Pringle 4631 (MICH, S, US). About 4-5 km. E. of
Juchitain, King 1631 (US). YucarA4n: Gaumer 487 (MICH, S, US). Quinrana
Roo: Cozumel Island, Gawmer 88 (K).

HAITI: Port-au-Prince and vicinity, Ekman H. 9342 (8, US); Rose 22096
(GH, US); Holdridge 1005 (MICH, U8).

DOMINICAN REPUBLIC: Santo Domingo, Valle del Cibao, Ekman H.
15983 (S).

JAMAICA: Swartz s.n. (8). D. D. Wright s.n. (8S).

LESSER ANTILLES: Martinique: Near Casa Pilote, Hahn 240 (S). Grena-
DINES: Bequia, Eggers 7058 (US). Cannovan I., Beard 1409 (S, US).

VENEZUELA: Anzofrecui: Puerto La Cruz, Guaraguao, Pittier 14909 (US).
Carasoso: Puerto Cabello, Linden 1549 (W). Lara: Near Barquisimeto,
Saer 233 (US).

COLOMBIA: Wagener 32 (US). ?CoLtomBra: Humboldt & Bonpland, F neg.
35386 ex P of type of Melochia Turpiniana H.B.K. (US). MagpaLtena: Around
Rio Frio, between the Cienaga de Santa Maria and the foothills, Pittéer 1601 (US).
Santa Marta, HW. H. Smith 474 (L,8, US). AruAntico: Barranquilla and vicinity,
Bro. Elias 186a (US); Torregroza et al. 18 At 505 (US); Dugand 3254 (US); Bro.
Paul C-2 (US). Near Puerto Colombia, Kattéh et al. 19 At 033 (US); Bro.
Elias 24 (US), 824 (US). Botfvar: Cartagena and vicinity, Billberg 33 (8);
Engstedt 45 (8); Dahlin s.n, (8); Killip & Smith 14017 (US), 10462 (US).

49d. Melochia tomentosa L. var. speciosa (S. Wats.) A. Goldberg, comb. et
stat. nov.

Melochia speciosa 8. Wats. Proc. Amer. Acad. 24:42. 1889.
Melochia pterocarpa J. Ar. in Mém, Inst. Sci. Madag., ser. B, 7:68. 1956.
The leaves of this variety are small and differ from those of the
other varieties in being green on both surfaces and hispidulous rather

GOLDBERG—GENUS MELOCHIA 335

than tomentulose. The plant is very slender, has inflorescences at
few nodes, and the flowers are larger than average for the species.

The zygomorphy of the corolla and androecium reported by Arénes
for his M. pterocarpa may have been due to differential shrinkage in
drying or to an anomalous condition. I dissected flowers from the
type specimen and found petal sizes intermediate between the ex-
tremes reported by Arénes. The range in length of petals from one
corolla was 13.3-14.4 mm. The calyx teeth of M. tomentosa may vary
slightly in size in a single calyx.

Type Locauiry: The type was collected by EH. Palmer, no. 650, at
Guaymas, Sonora, Mexico, and is cited below.

DistripuTion: Endemic to Western Mexico, adventive in
Madagascar.

MEXICO: Sonora: Guaymas, FE. Palmer 148 in 1887 (MICH, US), 266 in
1897 (S, US), 650 in 1887 (US, type of Melochia speciosa 8. Wats.). Three mi, W.
of Jori, Pennell 19458 (US). Srnatoa: Vicinity of Culiacan, Brandegee s.n. (US).
Vicinity of Topolobampo, E. Palmer 205 in 1897 (MICH, US); Rose et al.
13275 (US). Cerros del Fuerte, 18-24 mi. N. of Loe Mochis, Gentry 14288 (US).
ZACATECAS: San Juan Capistrano, Rose 3551 (US). Jaxisco: Near Huejuquilla,
Rose 3558 (US). MicnoacAn: Zitacuaro-Tuzantla, Hinton 13064 (MICH, US).

MADAGASCAR: Scott Elliot 650 (P), type of M. pterocarpa J. Ar.

50. Melochia crenata Vahl, Symb. 3:86, pl. 68. 1794.
Riedleia crenata (Vahl) DC. Prodr. 1:491. 1824.
Visenia crenata (Vahl) Spreng. Syst. 3:30. 1826.
Melochia diffusa Bert. ex Spreng. Syst. 3:30. 1826.

A sparingly branched prostrate subshrub or herb with shoots to 1
meter long, the young stems and petioles densely tomentose, somewhat
canescent, hairs stellate, 0.3-'.3 mm. long, internodes 2.3-8.3 cm.
long; stipules 2.5-3.7 mm. long, 0.8-2.0 mm. wide, ovate to lanceolate,
acute, mostly ciliate, hairs simple, 0.3-1.0 mm. long, shorter stellate
hairs 0.3-0.5 mm. long on the abaxial surface; petiole 0.7—2.0 cm. long.

Lamina 1.4—-3.5 em. long, 1.1-2.6 em. wide, oval to broadly ovate,
the base rounded, rarely slightly cuneate, the apex rounded, one side
sometimes slightly larger than the other, both surfaces densely tomen-
tose, somewhat canescent, velvety to the touch, hairs stellate, 0.1-0.4
mm. long, the margin irregularly crenate-serrate, serrations 1.3-3.5
mm. wide, 0.5-1.1 mm. high, 6-7 pairs of primary veins, 2 basal.

Inflorescences 2.0-3.7 em. long, 90° from the leaf to leaf-opposed,
at 2-9 nodes, the peduncle 0.7—2.1 cm. long, 1 or 2 at a node, terminated
by a 2—5-flowered umbel, the pedicel 4.0—-7.0 mm. long; bracts 1.0—-4.5
mm. long, 0.4-0.8 mm. wide, lanceolate or deltoid- to ovate-acute,
mostly ciliate, hairs simple, forked, and stellate, 0.3-1.0 mm. long.

Calyx 5.2-6.2 mm. long, 4.2-5.2 mm. wide, densely tomentose,
hairs stellate, 0.3-1.0 mm. long, the teeth 3.0-3.6 mm. long, 1.5-1.9

336 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

mm. wide, deltoid-acute to deltoid-acuminate, the sinus acute; petals
purple, magenta to rather pale red, 11.2-15.5 mm. long, 3.6-5.2 mm.
wide, obovate-cuneate, sometimes with scattered, mostly ciliate, gland-
ular hairs 0.05 mm. long, attached to the stamens for 2.5-3.3 mm., the
claw 1 mm. long.

Longistylous form. According to Fawcett & Rendle (13), stamens
7-8 mm. long, the filaments free for 2.5-3.0 mm.; pistil about 11 mm.
long, the styles united shortly (1.5-2 mm.) at the base, the ovary and
styles somewhat hirsute with branched hairs.

Brevistylous form. Stamens 10.4-11.3 mm. long, the filaments
flattened, 0.3 mm. broad, free for 5.5-6.5 mm., with scattered glandu-
lar hairs 0.05 mm. long, the anther 1.3-2.0 mm. long, 0.6—0.8 mm.
wide, oblong, slightly emarginate at both ends; pistil 4.5-6.9 mm. long,
the styles free for 2.1-3.0 mm., united for 0.5-1.0 mm., filiform, papil-
lose for 0.7-0.8 mm., the ovary narrowly ovoid, the united part of
styles and ovary hirsute-tomentose, hairs 0.5 mm. long.

Fruit 10.0-10.5 mm. long, of which the rostrum is 1.5-2.0 mm.
long, 6.7-8.7 mm. wide, pentapterous-ovate, narrowed at the base to
a stripe 0.5-1.0 mm. long, the outer angles of wings rounded, hardly
evident, below the middle of the capsule, the sulci between the carpels
extending a third to half way to the center of the fruit, densely tomen-
tose, hairs stellate, 0.2-0.5 mm. long; dehiscence loculicidal along the
dorsal suture; seeds not quite ripe, 2.9-3.0 mm. long, 1.7-2.0 mm.
wide, brown, usually 2 per cell.

Flowering specimens collected in July, September, and October;
fruiting ones, in July and October.

TypE Locauity: Near Santa Marta, Colombia. Type collected by
von Rohr.

DistripuTion: Colombia and Jamaica in sandy soil near sea level.

?JAMAICA: ?W. Wright (W).

COLOMBIA: Goasira: Dawe 532 (US). Magpatena: Santa Marta, H. H.
Smith 473 (K, P, 8, US), 475 (A). F neg. 21602 of type ex C (US). ?Santa
Marta, Balbis (8S). Riohacha, Haught 4409 (US).

The species can be distinguished from other members of the section
by its fruits, which are not pyramidal and have rounded rather than
acute or acuminate wings. Its trailing habit, leaves rounded at the
apex, and long pedicels are also aids in identification. It is perhaps
most closely related to Mf. tomentosa L.

The collection of the specimen photographed as F 21602 ex C is
attributed to Ryan, but the type was collected by von Rohr.

This species is cited as occurring in Peru by Macbride (39). To my
knowledge, it does not occur there.

GOLDBERG—GENUS MELOCHIA 337

51. Melochia pyramidata L. Sp. Pl. 674. 1753.

A slender erect or spreading annual or perennial herb to shrub,
0.08—2.0 meters high, usually branched, sometimes several branches
from the base of the stem, the root sometimes thickened and 0.51.3
cm. wide, the young stems hirsute-tomentulose, hairs in a longitudinal
line or surrounding the stem, simple, sometimes also stellate and
forked, 0.2-0.5 mm. long, internodes (0.2—) 2-10 cm. long; stipules
1.5-5.3 mm. long, 0.2-1.0 mm. wide, deltoid-acuminate or lanceolate,
rarely linear, hirsute-tomentulose and ciliate, hairs simple, forked,
and stellate, 0.2-0.8 mm. long; petiole (0.1—-) 0.4-2.0 (-4.7) cm. long,
hirsute-tomentulose on the upper surface or all the way around.

Lamina (0.7—) 2-6 (-10.5) cm. long, (0.3-) 0.6-3.0 (-7.7) cm. wide,
the upper leaves tending to be lanceolate, the lower ovate, rarely
oblong, the base rounded, sometimes cordate or attenuate, the apex
acute, acuminate or rounded, the basal leaves usually smaller, some-
times nearly orbicular, the upper surface green, subglabrous or hirsute,
hairs simple, appressed, sometimes also forked and stellate, 0.1-1.0
mm. long, the lower surface green, with few hairs on the veins, some-
times slightly yellowish-canescent, or tomentulose, hairs stellate,
sometimes also forked, simple, and glandular, 0.1-0.4 (-0.7) mm. long,
the margin irregularly crenate-serrate, serrations 0.8-4.4 mm. wide,
0.2-2.0 mm. high, 5-11 pairs of primary veins, 1 or 2 (-3) are basal.

Inflorescences generally leaf-opposed, sometimes 90° from the leaf,
rarely also axillary or terminal, 1.3-5.2 cm. long, at (1-) 3-8 upper
nodes, umbellate, (1—-) 3-8 (-15) flowers per umbel, the peduncle
slender, (0—) 0.2—1.6 (—3.3) cm. long, 0.4 mm. wide, rarely branched,
the pedicel 1-5 (-6) mm. long; bracts 0.5-3.0 mm. long, 0.2-0.7 mm.
wide, deltoid-acuminate, lanceolate or linear, ciliate and_ hirsute-
tomentulose, hairs simple, forked, and stellate, 0.1-0.5 mm. long.

Calyx 3.7-6.2 mm. long, 2.1-4.0 mm. wide, ciliate and _hirsute-
tomentulose, hairs stellate, and forked, 0.1-0.5 mm. long, the teeth
2-4.9 mm. long, 0.6-1.5 mm. wide, usually deltoid-acuminate, some-
times deltoid-acute, the sinus acute to narrowly rounded; petals
usually purple, pink, violet, blue, sometimes yellow proximally,
rarely red, white or yellow, (8—-) 7-11.5 mm. long, (1.5-) 2.4-3.0 mm.
wide, obovate-oblanceolate-spatulate, attached to the staminal tube
for 1-3 mm., the claw 0.6-2.5 mm. long, sometimes glandular hairs
0.05 mm. long on the petals.

Longistylous form. Stamens (2.6—) 3.5-6.3 mm. long, the filaments
free for 0.7-2.0 mm., the anther 0.7-1.2 mm. long, 0.4—0.6 mm. wide,
oblong, emarginate at both ends; pistil 4.3-9.0 mm. long, the styles
free for 2-4.8 mm., united for 0.5-2.1 mm., filiform, papillose for
0.6-1.0 mm. at the apex, the ovary ellipsoid, the styles and ovary

338 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

hirsute-tomentulose, hairs stellate, 0.1-0.4 mm. long, the stipe 0—0.2
mm. long.

Brevistylous form. Stamens 5.7-7.5 mm. long, the filaments free
for 3-4 mm., the anther 1-1.2 mm. long, 0.5-0.6 mm. wide, shape
same as that above; pistil 3.7-5.0 mm. long, the styles free for 1.4—-
2.0 mm., the ovary ellipsoid to oblong-ellipsoid, subglabrous to
hirsute-tomentulose, hairs 0.1-0.2 mm. long.

Fruit often with purple blotches, 5-9.5 mm. long, (5—) 6.5-14 mm.
wide, the rostrum short, 0-2.1 mm. long, the base narrowed into a
stipe 0.2—-1.0 mm. long, occasionally 6 or 7 carpels present, pyramidal,
pentapterous, the points of wings acute or acuminate, at the middle
or toward the base of the fruit, the sulci between the carpels extending
half way to the center of the fruit, the base stipitate for 0.2-1.0 mm.,
subglabrous, hirsute or tomentulose, hairs only stellate or also simple
and forked, 0.1-0.5 mm. long; dehiscence loculicidal all along the
dorsal and ventral sutures; seeds 1.7-2.6 mm. long, 1.1-1.7 mm.
wide, trigonal, brown, 1 or 2 per cell.

Flowering and fruiting specimens collected throughout the year in
the West Indies and for 8 months in the colder regions near the ex-
tremes of the range.

Key to Varieties of M. pyramidata

1. Lamina subglabrous, hairs on upper surface often simple.

5la. M. pyramidata var. pyramidata
1, Lamina usually tomentulose beneath, hairs on upper surface more commonly

stellate. . 2. ...... =... . .51b. M. pyramidata var. hieronymi

51a. Melochia pyramidata L. var. pyramidata

Melochia domingensis Jacq. Select. Stirp. Amer. 194. 1763.

Moluchia fruticosa Medic. Malv. 10. 1787.

Moluchia herbacea Medic. Malv. 10. 1787.

Moluchia domingensis (Medie.) Steud. Nom. IT, 2:155, 1841.

Sida Sabeana Buckley, Proc. Acad. Phila. for 1861, 449. 1862.

?Melochia pyramidata L. var. normalis O. Ktze. Rev. Gen. Pl. 3(2):25. 1898.
Moluchia pyramidata (L.) Britt. Brooklyn Bot. Gard. Mem. 1:69. 1918.

The leaves of the typical variety are subglabrous.

Type Locauity: Brazil. The type is in the Linnaean herbarium.

DistrIBUTION: Southern and western Texas to Costa Rica, West
Indies, northern South America to northern Argentina, adventive in
tropical and subtropical areas in various parts of the world, usually
in open disturbed places, often along the roadside, margin of culti-
vated fields, and in waste places, in various types of wet or dry soil,
at 0-1,560 meters elevation, usually below 1,000.

UNITED STATES: Texas: Guadalupe River, Lindhetmer 356 (US, W).

Western Texas to El Paso, C. Wright 1334 (US). Mexican Boundary Survey,
Parry et al. s.n. (US). comaL: Comanche Spring, New Braunfels, Lindheimer 687

GOLDBERG—GENUS MELOCHIA 339

(US, W). Bracken, Groth 110 (US). Bexar: 25 mi. N. of San Antonio, Schulz
554 (US). cotorapo: Eagle Lake, E. J. Palmer 6601 (US); Barkley et al. 46449
(S, W). Gatvesron: San Leon, Fisher 5083 (US). uvaLpE: Uvalde, E. J. Palmer
12295 (US). Kinney: Near Rio Grande River, Correll 14901 (US). BRazorRIA:
Brazos River, Bush 271 (US). Columbia, Bush 1346 (US). maracorpa: Bay City,
Fisher 180 (US). E. J. Palmer 9625 (S, US). Near Colorado River W. of Bay City
Rogers 6599 (W). Palacios, gulf coast, Tharp 178 (MICH). Maverick: Eagle
Pass, Havard s.n. (US). caLHoun: Guadalupe River bottoms, 15 mi. SW. of
Port Lavaca, Cory 51145 (US). Kiepera: Kingsville, Bogusch S-130 (US).
WEBB: Laredo, J. N.& J. S. Rose 11019 (US). Laredo, Rio Grande River,
E. Palmer 117 in 1879-80 (MICH, US). urpatao: Mission, Shiller 843 (US).
Tabasco, Clover 218 (MICH). Six mi. 8. of Alamo, Clover 879 (MICH). Rio
Grande Valley, M. L. Walker 41 (8, W). witiacy: E. of San Perlita, C. L. & A. A.
Lundell 8782 (MICH). cameron: Brownsville, Hanson 300 (US); Rose & Russell
24245 (US); Runyon 554 (US); Tharp 1179 (US). Las Palmas Plantation, about
4 mi. SW. of Brownsville, Correll 14862 (US). Arroyo Colorado, Olmito road,
Clover 1261 (MICH).

MEXICO: Sartorius sn. (US). Mirador, Sartorius sn. (US). Sonora:
Vicinity of Alamos, Rose et al. 12912 (US); Gentry 4770 (MICH, US). On road
to Guaymas 16 mi. 8. of Hermosillo, Wiggins & Rollins 205 (MICH). Coanutta:
Sabinas, Nelson 6816 (US). About 23 mi. S. of Sabinas, Shreve 8418 (MICH,
US). Palm Canyon, Marsh Jr. 972 (GH). Muzquiz, Santa Anna Canyon,
Marsh Jr. 444 (GH). Monclova, White 1726 (MICH). Nusrvo Leon: Rio
Santa Catarina, Monterrey, Bro. Arséne 6304 (US). About 28 mi. 8. of Nuevo
Laredo on road to Monterrey, T. C. & E. M. Frye 2340 (US). Valley, Monterrey,
Pringle 1939 (MICH, 8, US, W). Forty mi. 8. of Laredo, Schery 7 (MICH).
Mts. near Monterrey, Diente Canyon, C. H. & M. T. Mueller 303 (W). Tam-
AuLIPAS: San Lucas, Viereck 737 (US), 876 (US). Victoria and vicinity, E.
Palmer 68 in 1907 (US); Fisher 3355 (US). Vicinity of El Mulato near Tanque,
Bartlett 10996 (MICH, US), 10994 (MICH). Vicinity of Marmolejo, Bartlett
10945 (MICH). Buena Vista Hda., Wooton s.n. (US). Ten mi. E. of Ciudad
Mante, Kenoyer & Crum 3665 (MICH). Huisachal, Stanford et al. June 29,
1949 (US). Srnatoa: Mazatlan and vicinity, Rose et al. 183676 (US); Lamb 354
(US); Brandegee, Oct. 8, 1893 (UC). Culiacan, Rose et al. 14856 (US); Brandegee,
Sept. 19, 1904 (UC). Vicinity of Villa Union, Rose et al. 13929 (US). Vicinity
of Topolobampo, Rose et al. 138274 (US). Imala, Gentry 4944 (MICH). San
Luis Potosf: Tamasopo, Fisher 3735 (US). Tancanhuitz, Nelson 4394 (US).
Nayarit: Vicinity of Acaponeta, Tepic, Rose 3114 (US); Rose et al. 14221 (US).
Jauisco: Near Chapala, Rose & Painter 7671 (US). Tequila, E. Palmer 371
in 1886 (MICH, US). Corcovado Canyon, 11 mi. NE. of Autlin, R. L. & C. R.
Wilbur 2376 (MICH, US). Five mi. E. of Autlin, R. L. & C. R. Wilbur 2234
(MICH, US). Querriraro: Bro. Aguiel 10446 (US). Hipauco: Huejutla,
C. & E. Seler 620 (US). Vera Cruz: Mirador, Wawra 968 (W). Soledad,
Wawra 544 (W). Barranca de Zacuapan, Purpus 2057 (US). SAN ANDRis
TUXTLA: Between Los Mangos and San Juan de Aziticar, Dressler & Jones 206
(US). Moretos: Yautepec, Rose et al. 5345 (US), 6560 (US). Cuantla and
vicinity, Pringle 9405 (US); Lyonnet 673 in part (US). Xochiltepec, Lyonnet
2159 (US). Mexico: Wawra 393 (W). TeMascaLTEPEC: Rincdén del Carmen,
Hinton 1742 (US). Tenayac, Hinton 4018 (US). Micuoacdn: One mi. W. of
Coahuayana, 20 mi. SE. of Tecoman, Turner 1973 (MICH). South shore of
Lake Chapala, Schery 173 (MICH). ziTacuaro: Zitacuaro-Tuzantla, Hinton
13073 (MICH, US). wurramo: Tacupa, Hinton 5622 (US). Coxima: Colima,
E. Palmer 165 in 1897 (MICH, S, US). Manzanillo, BE. Palmer 912 in 1890

340 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

(MICH, US). REVILLAGIGEDO ISLANDS: Clarion Island, Howell 8872 (US);
Mason 1560 (MICH, US); Anthony 408 (US). Socorro Island, Barkelew 247
(US). Gurrrero: Agua de Correa, Langlassé 403 (US). Acapulco and vicinity,
E. Palmer 193 in 1894-95 (US). Cafion de la Mano Negra, near Iguala, Rose
et al. 9329 (US). Oaxaca: Oaxaca and vicinity, Conzatti 1601 (US); Rose &
Hough 4646 (US). Between Mitla and Oaxaca, J. N. & N.S. Rose 11285 (US).
Half a km. E. of and 4 km. NNE. of Tehuantepec, King 320 (MICH). Puerto
Angel, Morton & Makrinius 2641 in part (US). TuxTEepPEec: Chiltepec and
vicinity, Martinez-Calderon 265 (US). Tarasco: Reforma, Belancan, Matuda
3189 (MICH). CamprcHe: Tuxpejia, Lundell 966 (MICH, US). Yucatan:
San Anselmo, Gauwmer 2208 (MICH, 8, US, W). Uxmal, Steere 2016 (MICH).
Chichen Itzd, Sleere 1894 (MICH). Yot Tzonot, Gawmer 13832 (MICH).

GUATEMALA: Agua Blanca, Tejada 104 (US). HupHvuETEeNANGO: Chorulé,
C. & E. Seler 2867 (US). Attra Verapaz: Chamiquin, von Tiirckheim 560 (US,
W). IzapaL: Quirigud, Standley 23796 (US). Zacara: Gualan, Kellerman
5617 (US). Santa Rosa: La Vega, Heyde & Lux 6079 (US).

BRITISH HONDURAS: Lower Belize River, Record s.n. (US). Corozal
District, Gentle 313 (MICH), 282 (MICH), 344 (MICH). Corozal, Lundell
4797 (MICH). Santa Rita, Gentle 579 (MICH).

HONDURAS: Mosquito coast, Wullschldgel 56 (W). Guaymas, Standley
55454 (US). Gracias: Hjalmasson s.n. (S). Santa BARBARA: San Pedro
Sula, Thieme 5154 (US). MorazAn: Zamorano, Molina R 371 (US).

SALVADOR: Renson 12 (US). Finca San Nicolas, Choussy 88 (US). Anuva-
cHAPAN: Padilla 410 (US). Vicinity of Ahuachapdn, Standley 19946 (US).
Santa ANA: Calderén 395 (US). Sonsonatre: Santa Emilia, Standley 22065
(US). Armenia, Standley 23436 (US). La Lrpertap: Vicinity of Ateos,
Standley 23380 (US). San Satvapor: Calderén 19 (US). Velasco 8923 (US).
Vicinity of San Salvador, Standley 19417 (US), 20571 (S, US). San Martin and
vicinity, Calderén 436 (US); Standley 22537 (US). San Miauex: Laguna de
Olomega, Standley 20999 (US). Hacienda Potrero Santo, 8. side of Lake Olomega,
Tucker 849 (US).

NICARAGUA: CuHrINANDEGA: Baker 581 (US). CuHontauss: Friedrichsthal
711 (W).

COSTA RICA: Nicoya, Cook & Doyle 664 (US).

BAHAMAS: New Provipence: Britton & Brace 594 (US). Near Nassau,
Curtiss 21 (L, US). Lake Killarney, FE. G. Britton 3285 (US).

CUBA: Jamain s.n. collected 1863 (W). de la Sagra 111 (W). C. Wright
3509 (US). Pinar pet Rfo: Near Guanajay, W. Palmer & Riley 668 (US).
Hapana: Baker & Wilson 363 (US). Vedado, Ekman 21 (8S). Maranzas:
Yumury Valley, Rugel 41 (L). Las Vituas: Vicinity of Soledad, Howard 5013
(MICH, US); Morton 4261 (US). District Cienfuegos, Combs 273 (US). Li-
mones, Soledad, Cienfuegos, Jack 6250 (8S). Banao Mts., Bro. Leén 21940 (US).
ORIENTE: Santiago de Cuba and vicinity, EH. Palmer 375 in 1902 (US), Ekman
8037 (8S). Vicinity of San Luis, Pollard & W. Palmer 331 (US). Bayate,
Central Miranda, Ekman 5681 (S), 9662 (S). Central highway, Santa Rita,
Jiguani, Bro. Clemente 6073 (US).

HAITI: Vicinity of Port-au-Prince, Leonard 2854 (US). Vicinity of St.
Marc, Leonard 2962 (US), 2966 (US). Vicinity of Fond Parisien, Etang Saumatre,
Leonard 4188 (US). Vicinity of Jean Rabel, EH. C. & G. M. Leonard 13041 (US).
Miragoane, Eyerdam 22 (US), 170a (US), 489 (US). Massif des Matheux,
Mirebalais, Ekman H2291 (8, US). Massif de la Selle, Petionville, Ekman
H8113 (8, US). Du Norp: Vicinity of St. Michel de l’Atalaye, Leonard 7310
(US), 7352 (US). Ds w’ArtTrBoniTB: Vicinity of Ennery, Leonard 8910 (US).

GOLDBERG—GENUS MELOCHIA 341

TortvuE Istanp: Basse Terre, E. C. & G. M. Leonard 12510 (US). La Vallée,
E. C. & G. M. Leonard 15358 (US), 15520 (US). Palmiste, Ekman H5123 (S).
GonaVvE IsuaNp: Vicinity of Pikmi, Leonard 5130 (US).

DOMINICAN REPUBLIC: Santo Domingo, Ritter Jan. 1823 (W); C.
Wright et al. 132 (US); von Tiirckheim 2502 (L, 8, US); Ekman H12505 (S, US);
Allard 13109 (US), 13375 (S, US), 13682 (US), 14028 (US). Santo Domingo,
Azua, Rose et al. 4418 (US), 4003 (US). Near Constanza, von Ttirckheim 3251
(L, 8, US, W). Puerto Plata, Raunkiaer 1103 (US), 1191 (US). Haina, Faris
21 (US), 30 (US), 80 (US), 156 (US). Santiago: Santiago, Raunkiaer 903
(US). Novarrete, Abbott 1106 (US), 1109 (US). La Delgada, Jiménez 1002
(US). Baranona: Near Barahona, Fuertes 66 (L, S, US, W). San Juan:
Juan Santiago, R. A. & E. 8S. Howard 9222 (8S, US). Monts Criste: Moncion
Valeur 183 (MICH, 8, US), 849 (US).

JAMAICA: Swartz s.n. (8). Cuming 99 (W). Sr. Exizasetu: Pepper, Harris
9861 (US). Lowrr CLiarenpon: Inverness, Harris 12751 (US).

PUERTO RICO: Wydler 305 (L). Toa Alta, Stahl 603 b (L). Road from
Guayama to Aguirre, Underwood & Griggs 388 (US), 412 in part (US). Aguirre,
Goll 549 (US). Road from Coamo to Ponce, Underwood & Griggs 579 (MICH,
US). Between Coamo and Aybonito, Sintenis 1971 (US). Coamo Springs, Goll
729 (US); Heller 6108 (L, US). Guaraguao, Ponce, Goll 818 (US). Magueyes,
Ponce, Goll 777 (US). Along Rio Portuguese, Ponce, Underwood & Griggs 726
(US). Guanica, Sargent 30 (US); Smyth 5013 (US). Cabo-Rojo, Sintenis
801 (L, $8, US). Between Pefiuelas and Tallaboa, Sintenis 4817 (L). ViequrEs
IsLaNnD: Santa Maria to Caballo Colorado, Shafer 2692 (US).

LESSER ANTILLES: U.S. Viren Istanps: st. croix: West s.n. (L).
Bassin, Ricksecker 98 (US). Frederiksted, Rose et al. 3201 (US). Grange, Oersted
s.n. (8S); Hansen s.n. (8S). Anna’s Hope, Thompson 878 (S). British VirGIN
IsLANDS: TORTOLA: Bank, Sea Cow Bay, Britton & Shafer 925 (US). Experi-
ment Station, Fishlock 12 (US), 163 (US). Sv. BarTu&Lemy: von Goés s.n. (8).
Forsstrém s.n. (8). St. Eustatius: Suringar May 1885 (L). Sr. Kirrs: Near
Sandy Point, Britton & Cowell 149 (US). AnTIGuA: Near Bellevue, Bor 932
dupl. (US). St. John’s, Ramsammy 110 (US). Montserrat: Plymouth, Shafer
1 (US); Potter 5553 (GH). GuaprLourE: Duss 2671 (US). Pointe-a-Pitre,
Hammarlund 32 (8). Jarry, Questel 598 (US). Gosier, Questel 2472 (US).
Marie GALANTE: Questel 1456 (US). Dominica: Wallhoure, Eggers 652 (L).
?Sr. Lucia: ?Grande Anse, Eggers 6460 (US). Grenapa: Eggers 6460 (US).
St. George, Broadway s.n. (US). Victoria, Suringar s.n. (L).

VENEZUELA: Las Adjuntas and Rio Macarao, Eggers 13277 (L, US; these
specimens lead toward M. pyramidata var. hieronymi). Muiranpa: Around Dos
Caminos and Los Chorros, Pittzer 5923 (US). FrprraL District: Caracas, San
Bernardino, Fernandez 194 (US). Lower Catuche wood above Caracas, Pittier
7026 (US). Aragua: Maracay, Vogl 336 (S).

COLOMBIA: Plages de la Magdalena, Goudot, collected 1844 (P). Maapa-
LENA: Santa Marta, H. H. Smith 472 (L, 8, US). AruAntico: Barranquilla and
vicinity, Bro. Elias 1343 (US, W). Usiacurf, Arroyo Cagén, Dugand & Garcia
Barriga 2346 (US). Between Palmar de Varela and Pondera, Dugand 4509 (US).
Puerto Colombia, Pennell 12025 (US); Dugand 5473 (US). Boxfvar: Cartagena
and vicinity, Billberg 34 (S); Killip & Smith 14159 (US). Sincelejo, Pennell 4073
(US). Turbaco, Bro. Heriberto 494 (US). ?Totima: Tocaima, Arbelaez 2092
(US). Hua: Rio Cabrera ca. 18 km. NE. of Villavieja, S. G. Smith 1197 (US).
Antioquia: Occidente, Peldez et al. 581 (US). VALLE peL Cauca: Cuchilla, E.
of Zarzal, Pennell et al. 8529 (US). Zaragoza, Cuatrecasas 22782 (US). Cauca:
Cali and vicinity, Pittier 619 (US). Cerrito in Cauca Valley, Lehmann 3866 (US).

342 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

ECUADOR: Guayas: Guayaquil and vicinity, Asplund 5113 (R, US); Fager-
lind & Wibom 105 (S); Mille s.n. .MICH); Haenke s.n. (W). Oil camp between
Guayaquil and Salinas, Hitcheock 19983 (US). Vicinity of Durdn, J. N. & G,
Rose 23615 (US).

PERU: San Martin: Pueayacu 11 km. from Tarapoto, Ferreyra 7752 (US).
Loreto: Maucallacta, Rfo Paranapura, Klug 3927 (8, US).

BOLIVIA: Santa Cruz: Cabezas, Peredo 443 (UC). Cabezas Laguna del
Turco, Peredo 146 (8).

BRAZIL: Sellow s.n. (US). MaranuAo; Island of Sio Luiz, Frées 11887 (US),
24080 (US). Crard: Alleméo & Cysneiros 133 (R, US). Fortaleza to Crato,
Duarte 1383 (US). YrorTALEzA: Floresta, Drouet 2324 (MICH, 8, US). Mara-
canat, Fernandez 30 (R); Ducke 2606 (US). Ipt, Léfgren 213 (8). Cedro,
Léfgren 74 (S). Paraipa: Xavier s.n. (R). Areia, Vasconcellos 271 (US). Prr-
NAMBUCO: Olinda, Bento 40 (RB). Sio Joao dos Pombos, Falcdo et al. 1015
(US). Tome, Recife, Monteiro 210 (R). Banta: Serra Jacobina, Blanchet 2677
(W). Sento 8é, Campos Porto 2470 (RB). Minas Gerats: Glaziou 13553 (P).
Arassuahy near Rio Manso, Glaziow 13557 in part (R). irururaBa: Funda,
Macedo 2449 (US). Go1ds: Gardner 3605 (W). Between Corallinho and Cazada
Telha, Pohl 2801 (W). About 13-15 km. E. of Formosa, Dawson 15098 (US).
Distrito FEDERAL: Occhioni 425 (US). Maro Grosso: Corumb4, Hoehne 2983
(R), 3385 (R). Aquidauana, Reiss s.n. (GH). GuanaBara: Mundo Novo,
Botafogo, Kuhlmann ?4151 (RB). Corcovado, Diomsio (RB). Margin of Lake
Rodrigo de Freitas, Guerra s.n. (US). Rio pe JANEIRO: Araruama near Cabo
Frio, Glaziouw 10326 (R). Sao Pauto: Sellow 5288 in part (R). Parand: S.
Jeronimo da Serra, Hatschbach 3726 (US). Procedéncia Cruzeiro do Oeste,
Serra Dourada, Rio Irai, Braga & Lange 60 (US). Rio GRANDE Do Sut: SB.
Borja, Rambo (PACA 3048). Fazenda do Jarau near Quaraf, Rambo (PACA
27197).

URUGUAY: Fray Bentos, Fruchard Feb. 11, 1877 (P, US). Crrro Largo:
Rio Negro y a: Palleros, Rosengurtt B2044 (US).

PARAGUAY: Upper part of River Apa, Hassler 8037 (8S). BETWEEN Rio APA
AND Rio AQUIDABAN: Centurién, Fiebrig 4009 (L) ; Caballero-cué, Fiebrig 4780 (L).

ARGENTINA: Corrientes: San Martin: La Cruz boca rios Aguapey and
Uruguay, Jbarrola 1792 (UC). Estacion Yapeyu-costa Uruguay, Jbarrola 1903
(8S). MONTE CASEROS: Libertad, Costa, Arroyo Curuzt, Jbarrola 2491 (5).
PASO DE LOS LIBRES: Arroyo Yatay, Cristébal et al. 17 (UC). Musrones: Can-
delaria, G. J. Schwarz 503 (US). Loreto, Montes 73 B (US). Posadas, Vattuone &
Bianchi L. 92 (US). Entre Rios: Conception del Uruguay, Lorentz Apr. 1876
(W).

MASCARENES: R&untion: St. Denis, D’ Alleizette sn. (L). Mauritius:
Commerson 456 (L). Bojer s.n. (W). Andersson s.n. (5).

SUNDA ISLANDS: Java: ParRoEROKA: Bantoer, Backer 36519 (L). Road
from Bantoer (to) south coast, Backer 3876 (L). CELEBES: Boeton, Kjellberg 28
(L, 8); C. B. Robinson 2494 (US). Timor: Colleetor not given (L).

MOLUCCAS: Kisar: E. of Wonreli, inland, Bloembergen 3867A (L).
TANIMBAR: JAMDENA: Sifnana near Saumlaki, Pleyte 35 (L).

NEW GUINEA: Papua: Cenrrau Div.: NAKEOo: Baroka, Brass 3753 (US).

AUSTRALIA: NorrHern Terrirory: 25 mi. NNE of Mt. Sanford Outstation,
Perry 2174 (US). QUEENSLAND: Dietrich sn. (US). Near Morestone Station,
Perry 1010 (US).

This species is related to Melochia tomentosa L. The latter is usually
a taller plant, with hairier leaves, often canescent on both surfaces.

GOLDBERG—GENUS MELOCHIA 343

The inflorescences of M/. tomentosa are usually axillary, those of M.
pyramidata are usually leaf-opposed. The fruit of M/. tomentosa has a
longer rostrum and is often supported by a longer stipe. The calyx
of M. tomentosa is often longer (5-8.6 mm. vs. 3.7—6.2 mm. long).

The stem contains fine silky fibers (Frées). The flowers open 8
a.m. and close 11 a.m. (Runyon).

51b. Melochia pyramidata L. var. hieronymi Schum. in Mart. Fl. Bras. 12(3) :35.
1886.

Melochia pyramidata L. Grisebachit O. Ktze. Rev. Gen. Pl. 3(2):25. 1898.

Melochia pyramidata L. flava O. Ktze. Rev. Gen. Pl. 3(2):25. 1898.

Melochia pyramidata L. var. Hieronymi Schum. f. follis luteo variegatis and
f. transitoria, both Schum. & Hassl. Bull. Herb. Boiss. II, 4:70. 1903.

Melochia tomentosa L. var. mattogrossensis R. E. Fries, Svensk. Vet. Akad.
Handl. 42(12):10. 1908.

?Melochia pyramidata L. var. paraguariensis Hassl. f. intermedia Hassl.
Buenos Aires Univ. Nat. Inst. Bot. y. Farm. Trab. 21:85. 1909.

?Melochia pyramidata L. var. pseudotomentosa Hassl. Repert. Nov. Sp. Fedde
8:122. 1910.

This variety is hairier than the typical variety. The upper surface
of the leaf is hirsute and the lower surface is tomentulose. Often
there are more flowers per umbel.

Type Locauity: Southern Brazil, Paraguay, and northern Argen-
tina. Sellow 1751 (W) collected in Brazil, and cited below, is selected
as lectotype.

DistrrisuTion: Northern South America to northern Argentina.

VENEZUELA: Frprrau District: Caracas, van Landsberge 340 (S). Above
Caracas, Middle Catuche wood, Pittier 9812 (US). Middle Cotiza, near Caracas,
Pittier 7059 (GH, US). Road from country club to Rio Chacito, Lasser 656 (US).

COLOMBIA: Norte pE SAnTanpeEr: Valle del rfo Peralonso near Santiago,
Molina & Barkley 18 N.S. 1387 (US). Sanranper: Near Quebrada Monte Grande
2km. W. of Jordan, Langenheim 3095 (US). CuNpbINAMARCA: Jerusalén, collector
not given (US). To tira: Valle del rio Magdalena, Armero, Cuatrecasas 10486
(US), 10486-B (US). VaLuE pet Cauca: Cali Loma, Dryander 2200 (US).
Cuchilla, E. of Zarzal, Pennell et al. 8529 (GH). Cauca: Cali and vicinity,
Figueroa P. 906 (US). La Manuelita, near Palmira, E. side of Cauca Valley,
Pittier 791 (US).

PERU: Loreto: Tarapoto, Ule 6529 (L). Junin: La Merced, Macbride 5312
(GH, US). TARMA: Between La Merced and Quimiri, Chanchamayo, Ferreyra
3693 (US).

BOLIVIA: Santa Cruz: sara: Rio Surutd, Steznbach 7241 (K, 8S, US neg. 5420
ex K).

BRAZIL: Sellow 1751 (W, lectotype of the variety). Parafsa: Experimental
Station de Alagoinha, Deslandes 4 (RB). Gor1ds: Fazendina in Via ad Caretao,
Pohl 1738 (W). Maro Grosso: Corumb4, Malme 3072 (S, type of M. tomentosa
L. var. mattogrossensis R. E. Fries, also US neg. 5609); Hoehne 3010 (R, US),
3011 (R), 3012 (R); Pereira et al. 113 (RB). Cd&ceres, Hoehne 4335 (R). SAo
Pau.Lo: Rio Piracicaba, de Lima s.n. (RB). Iporanga, margin of Rio Ribeira,
Hatschbach 5190 (US). Rio GraNpE po Sut: Fazenda Santa Cecilia near 8.
Gabriel, Rambo (PACA 25662), (PACA 25765).

344 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

PARAGUAY: Villa Rica, Jorgensen 3917 (S, US). Est. Armonia, Anisits 2000
(S). Cordillera de Altos, Fiebrig 370 (K, L). Asuncién, ?Dicters 20 (K).
Central Paraguay, Morong 588 (US). Atro Paraqguay: Chaco, 21° 8S. Lat.,
Fiebrig 1275 (S).

ARGENTINA: Satta: Chicoana, Zabala 121 (GH, US). Capital, Krapovickas
1445 (S). Salta City, N. Caseros, Wall & Sparre 1187 (8S). ANTILLAS: Cerro
Negro, Venturi 10404 (S). @uacutpas: Alemania, Venturi 9914 (GH, 8, US),
9971 (S, US). Tucumdn: Estacion agricola esperimental, Skotisberg s.n. (S).
Camino a San Andrés, Cuezzo 28 (8). Tusca and Taruca Pampa at La Cruz,
Lorentz & Hieronymus 61 (US). capiraLt: Mataderos ?, Venturz 2190 (A, S§),
2190 in part (US). tTRaNcaAs: Vipos, Venturt 5017 (GH, US). Tari: Tipa,
Rocha 3656 (US). Cuesta Pariquillo, Venturi 2621 (US). Santiaco pEL EstTErRo:
C. PELLEGRINI: Cerro del Remate, Venturi 5654 (US). Caco: Jorgensen 2541
(US). Fontana, Meyer 2309 (GIL). CorrrENTEs: concErcion: Loma Alta,
S. Ybarrola 1116 (US). Musrtones: Posadas, Bonpland, Ekman 1975 (8). CAN-
DELARIA: Puerto Santa Ana, G. J. Schwarz 612 (UC). Arroyo “Martin chico,”
G. J. Schwarz 568 (UC).

Section V. Physodium (Presl) A. Goldberg, stat. nov.

Physodium Presl, Rel. Haenk, 2:150. 1835,

Shrubs or small trees 2-10 meters high, the young stems and
petioles tomentulose, hairs yellow brown, stellate, sometimes also
glandular, 0.1-0.7 mm. long, internodes 1.0-5.5 cm. long; stipules
2.5-6.0 mm. long, 0.2-1.2 mm. wide, deltoid-acuminate, lanceolate
or nearly linear, densely pubescent, hairs stellate, 0.1-0.6 mm. long;
petiole 1-3.5 em. long.

Lamina 3.5-14.8 em. long, 2.6-11.5 em. wide, ovate, one side
usually larger than the other, the base rounded, truncate or cordate,
sometimes oblique, the apex acute to acuminate, woolly or scabrous
to the touch, the upper surface covered with stellate hairs 0.1-0.5
mm. long, sometimes also with resin-colored granules or pulverulent,
the lower surface tomentulose, hairs stellate, 0.2-0.6 mm. long, young
leaves densely covered with hairs, older leaves sometimes subglabrate,
the margin irregularly crenate-serrate, serrations 1.5-7.5 mm. wide,
0.5-3.2 mm. high, 5-7 pairs of primary veins, 2 or 3 are basal.

Inflorescences terminal, extending 2-9 em. beyond the first leaf,
corymbiform, the ultimate divisions umbellate, 3-9 flowers per umbel,
also axillary, 1.5-12.5 em. long, in the axils of the upper 1-11 or
more leaves, the peduncle 0.3-2.7 em. long, with a primary branch
0.2—2.4 cm. long, a secondary one 0-1.3 cm. long, and a tertiary one
0-0.7 cm. long, the pedicel 2.5-13 mm. long at anthesis, becoming
5-19 mm. long in the fruiting stage, covered with stellate, and some-
times also glandular hairs 0.1-0.8 mm. long; bracts 1.5-6.2 (-11)
mm. long, 0.2-0.7 (-1.6) mm. wide, lanceolate to nearly linear,
densely covered with stellate, and sometimes also glandular hairs
0.1-0.6 mm. long.

GOLDBERG—GENUS MELOCHIA 345

Calyx rose, rose purple or yellow, densely pubescent, hairs stellate
and sometimes also glandular, 0.1-0.8 mm. long, at anthesis 7-15
mm. long, 3.6-8.5 mm. wide, with teeth 1.7-4.5 mm. long, 1.5-3.5
mm. wide, accrescent, becoming 15-36 mm. long, 8-19 mm. wide,
with teeth 3-12 mm. long, 3-10 mm. wide, the teeth deltoid-acute or
deltoid-acuminate, the sinus acute, inflated calyx acutely pentagonal,
reticulate-veined and with a midrib leading to each tooth and a vein
to each sinus; sometimes with glandular tissue between the calyx and
corolla for 0.5-1.2 mm. and 0.8-1.0 mm. high between the bases of
the petals; petals red, pink, yellowish red, light blue, violet blue,
violet or mauve, 8-17.4 mm. long, 2.0-9.5 mm. wide, oblanceolate,
obdeltoid or obovate-oblong, spatulate, sometimes slightly emargi-
nate at the apex, attached to the staminal tube for 2.5 mm., the claw
2.5-6.5 mm. long, somewhat thickened, with a few glandular hairs
and sometimes also forked and stellate hairs, 0.1-0.7 mm. long on
both surfaces; anther 1.4-2.6 mm. long, 0.6-1.2 mm. wide, oblong,
slightly emarginate at both ends.

Longistylous form. Stamens 5.3-6.8 cm. long, with a few forked
or stellate hairs 0.1-0.2 mm. long at the base of the staminal tube, the
filaments free for 0.4-1.4 mm.; staminodes 0-0.2 mm. long; pistil 6.2—
7.8 mm. long, the styles 3.5-6.0 mm. long, free to the ovary, papillose
and slightly clavate at the apex for 0.7-1.0 mm., with forked, stellate,
and simple hairs 0.2 mm. long, the ovary globose, pentagonal, covered
with stellate or glandular hairs 0.1-0.2 mm. long, the stipe 1-4.3 mm.
long.

Brevistylous form. Stamens 6.2-10.3 mm. long, sometimes with
very few stellate hairs 0.1 mm. long at the base of the staminal tube,
the filaments free for 1.5—5.0 mm.; staminodes 0-1.2 mm. long; pistil
3.2-6.5 mm. long, the styles free to the ovary, 0.8-5.3 mm. long, papil-
lose for 0.3-1.0 mm. at the apex, the stipe 1-4.5 mm. long.

Fruit 5.3-7.7 mm. long, 6.7-8.8 mm. wide, globose, acutely pentag-
onal, slightly retuse at the apex, the sulci between the carpels rather
deep, extending a third to half way to the center of the fruit, tomen-
tulose, hairs stellate and sometimes also glandular and simple, 0.1-0.5
mm. long, the stipe 4.3-6.7 mm. long; dehiscence loculicidal all along
the dorsal suture; seeds 2.8-3.9 mm. long, 1.6-2.5 mm. wide, obovate,
trigonal or somewhat flattened, dark brown, 1 or 2 per cell.

Flowering specimens collected December—April, June, July, Septem-
ber, October; fruiting ones, December—July.

Type species: Melochia corymbosa (Presl) Meissn. ex Steud.

DistTRIBUTION: Sinaloa to Oaxaca, Mexico, in wet woods, on rocky,
shady canyon slopes in the short-tree forest, on wooded, grassy hills,
in shade along small streams, on hills covered with dry-season decidu-
ous forest, at 250-1,650 meters elevation.

346 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

52. Melochia adenodes A. Goldberg, sp. nov.

Frutex vel arbor parva; laminis foliorum asymmetrice ovatis, basi
rotundatis, truncatis vel cordatis, obliquis, apice acutis vel acuminatis,
juvenilibus utrinque dense stellato-tomentosis; inflorescentiis termi-
nalibus et axillaribus, corymbiformibus sed ramulis ultimis cymoso-
umbellatis, pedicellis calicibusque pilis glandulosis valde vestitis;
petalis 8-12 mm. longis, calycem per anthesin haud superantibus;
staminodiis 0.2—-1.2 mm. longis.

A shrub to small tree 2-10 meters high, bark ashy, the young stems
and petioles densely covered with yellow-brown stellate hairs 0.1—-0.3
mm. long, glandular hairs sometimes also present, internodes 1-4.3
cm. long; stipules soon deciduous, 2.6—-3.4 mm. long, 0.5-0.6 mm. wide,
lanceolate, densely covered with stellate hairs 0.1-0.3 mm. long; pet-
iole 1-3.5 em. long.

Lamina 4-14.8 em. long, 2.6-11.5 cm. wide, ovate, one side of the
leaf usually larger than the other, the base usually oblique, truncate,
cordate or rounded, the apex acute to acuminate, young leaves densely
covered with stellate hairs, 0.1-0.3 (-0.5) mm. long, sometimes also
with resin-colored granules, or pulverulent on the upper surface, woolly
to the touch, older leaves subglabrate, the margin irregularly crenate-
serrate, serrations 1.7-7.0 mm. wide, 0.5-2.5 mm. high, 6 or 7 pairs of
primary veins, 2 (—3) are basal.

Inflorescences 3-12.5 cm. long, in the axils of the upper 1-11 or more
leaves, also terminal extending 3-9 cm. beyond the first leaf, corymbi-
form, the ultimate divisions umbellate, 3-6 flowers per umbel, the
peduncle 0.6-2.8 em. long, with a primary branch 0.2-2.4 cm. long, a
secondary one 0-1.3 cm. long, and a tertiary one 0-0.7 cm. long, the
pedicels 6-12 mm. long at anthesis, 10-19 mm. long in the fruiting
stage, covered with glandular hairs 0.3-0.8 mm. long and stellate hairs
0.1-0.2 mm. long; bracts 1.5-3.7 mm. long, 0.2-0.5 mm. wide, lanceo-
late to nearly linear, acute, densely covered with stellate, and some-
times also glandular hairs 0.1-0.3 mm. long.

Calyx rose, rose purple or yellow, covered with glandular hairs
0.2-0.6 mm. long and fewer stellate hairs 0.1-0.2 mm. long, at anthesis
11.1-15 mm. long, 7.7-8.3 mm. wide, with teeth 2-4.5 mm. long,
2-3.5 mm. wide, accrescent, becoming 1.8-3.6 em. long, 1.1-1.9 cm.
wide, with teeth 5-12 mm. long, 6-10 mm. wide, the teeth deltoid-
acute or deltoid-acuminate, the sinus acute, inflated calyx acutely
pentagonal, reticulate-veined and with a midrib leading to each tooth
and a vein to each sinus; internode between the calyx and the corolla
0.7-1.2 mm. long; petals pink, red, yellowish red, or pinkish orange,
8-12 mm. long, 2-2.7 mm. wide, oblanceolate or obovate-oblong,
spatulate, attached to the staminal tube for 2.5 mm., the claw 2.5-6.5
mm. long, thickened, a few glandular hairs 0,1-0.2 mm. long, and

GOLDBERG—GENUS MELOCHIA 347

sometimes also fewer stellate hairs 0.1 mm. long, scattered over both
surfaces of the petal.

Longistylous form. Stamens 6.8 mm. long, the filaments free for
1.3 mm., the anther 1.7-1.8 mm. long, 1.1 mm. wide, oblong, slightly
emarginate at both ends; staminodes extending 0.2 mm. beyond the
staminal tube; pistil 7.5-7.8 mm. long, the styles 6 mm. long, free to
the ovary, papillose and slightly clavate for 1 mm. at the apex,
scattered simple and forked hairs 0.2 mm. long all along the styles,
the ovary globose, covered with glandular hairs 0.1 mm. long, the
stipe 4.3 mm. long.

Brevistylous form. Stamens 6.2-10.3 mm. long, the filaments free
for 1.5-5.0 mm., very few stellate hairs 0.1 mm. long at the base of
the staminal tube, the anther 2-2.6 mm. long, 1—-1.2 mm. wide, oblong,
emarginate at both ends, more so at the base; staminodes extending
0.2-1.2 mm. above the staminal tube; pistil 3.2-6.5 mm. long, the
styles 1.5-5.3 mm. long, free to the ovary, papillose and slightly
clavate for 0.3-1.0 mm., the ovary globose, densely covered with
glandular hairs, and sometimes also stellate hairs, less than 0.1 mm.
long, the stipe 2.7-4.5 mm. long.

Fruit 6.3-7.7 mm. long, 7.5-8.0 mm. wide, globose, rather acutely
pentagonal, slightly retuse at the apex, the sulci between the carpels
fairly deep, extending half way to the center of the fruit, covered with
stellate, glandular and simple hairs 0.1-0.3 mm. long, the stipe 6—6.7
mm. long; dehiscence loculicidal all along the dorsal suture; seeds
3-3.7 mm. long, 1.7-2.1 mm. wide, obovate, somewhat flattened,
brown.

Flowering specimens collected from December to April (dry season);
fruiting ones, from December to June.

Key to Varieties of M. adenodes

1. Calyx teeth acute, typically 5-6 mm. long in the fruiting stage.

52a. M. adenodes var. adenodes
1. Calyx teeth acuminate, typically 11-12 mm. long im the fruiting stage.

52b. M. adenodes var. acuminata

52a. Melochia adenodes A. Goldberg var. adenodes

The calyx is rose to rose purple, and the teeth are acute and typically
5-6 mm. long in the fruiting state. The petals are rose to red.

TypPE tocauity: The type was collected by G. B. Hinton, no.
16203, at Huizontla, Coalcoman, Michoacén, Mexico, and is cited
below.

DistriBuTIoNn: Jalisco to Oaxaca, Mexico, in woods, in shade along
small streams, on wooded grassy hills, on hills covered with dry-season
deciduous forest, at 200-1,500 meters elevation.

348 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

MEXICO: S. Carlos, Liebmann 5888? (US). Jauisco: San Sebastian and
vicinity, Nelson 4120 (US); Mexia 1485-a (MICH, US). Santa Cruz de Vallarta,
Mexia 1289 (US). Toward La Resolana 16 mi. SSW. of Autlin, R. L. & C. R.
Wilbur 1451 (MICH, US). Southwest of Autlin, toward Manzanillo, McVaugh
10231 (MICH, US). Along road from Barra de Navidad to Tequezquitlin,
Concepcién, and Autlin, McVaugh 11870 (MICH, US). Muicuoackn: cCoAL-
coMAN: Hinton 12957 (US). Huizontla, Hinton 16203 (US, type). Aquila,
Hinton 15815 (MICH, US). Co trma: Manzanillo, EH. Palmer 1372 in 1891
(MICH, US). GuErRReERo: El Naranjo, Nelson 6987 (US). Oaxaca: San
Bartolo Yauhtepec, C. & E. Seler 1653 (US). TEQquimua: vicinity of Tuchahengo?,
Conzatti 4552 (US).

This species has been misinterpreted by Standley (52) and others
as Physodium corymbosum Presl. I have seen the type of the latter
species. The calyx and pedicels do not have glandular hairs, those of
M. adenodes are covered with glandular hairs. In his description
Pres] did not mention staminodes and his illustration did not show
any, nor did I see any in the withered flower parts of the type, which
is in the fruiting stage. Staminodes are quite evident in M. adenodes.

52b. Melochia adenodes A. Goldberg var. acuminata (Rose) A. Goldberg, comb.

nov.
Physodium corymbosum Presl var. acuminata Rose, Contr. U.S. Nat. Herb.
1:309. 1895,

The calyx may be red or yellow and the teeth are acuminate and
typically 11-12 mm. long in the fruiting stage. The petals are red
or yellowish red to pinkish orange.

TypE bocauity: Jalisco, Mexico. The type, cited below, was
collected by E. Palmer, no. 86, in 1886.

DistriBuTion: Sinaloa to Mexico, Mexico, on rocky, shady canyon
slopes in the short-tree forest, and on hills, at 280-750 meters elevation.

MEXICO: Srnatoa: Ortega 5104 (US). Mazatlan, Ortega 5608 (US). Cerro
Pricto, Pueblo de Coyotitan, Montes & Salazar 120 (US). Near Bacubirito,
W. Palmer 1249 (US). BapraRGuato: Arroyo de Carrisal, Geniry 5800 (MICH).
Cerro Colorado, Gentry 5483 (MICH). Duranao: Sianori, Ortega 5396 (US).
Jauisco: Barranca of Guadalajara, Pringle 5882 (US); E. Palmer 86 in 1886
(MICH, US, type). Between La Venta de Nochitiltie and Barranquitas along
Tepic-Guadalajara highway, McVaugh 12045 (MICH, US). Mexico: TEMAs-

CALTEPEC: Palmar, Hinton 7294 (US). Micnoackn: Monte de Loreto near
Inguardn, Langlassé 17 (US).

53. Melochia rhodocalyx C. Koch & Bouché, Index Sem. Hort. Berl. 14. 1854.

PLATE 9
Physodium dubium Hemsl. Diag. Pl. Nov. 1:4. 1878.

A shrub to small tree 2-6 meters high, the young stems and petioles
woolly, densely covered with yellow-brown stellate hairs 0.3-0.7 mm.
long, internodes 1.5—5.5 cm. long; stipules 2.5-6.0 mm. long, 0.2-1.2
mm. wide, deltoid-acuminate to nearly linear, rather densely covered
with stellate hairs up to 0.6 mm. long; petiole 1-3.5 em. long.

Lamina 3.5-12.5 em. long, 3.2-10.0 em. wide, ovate, often slightly

GOLDBERG—GENUS MELOCHIA 349

oblique, the base rounded, truncate or cordate, woolly to the touch,
hairs stellate, 0.2-0.5 mm. long, usually not overlapping on the upper
surface, usually overlapping on the lower surface, the margin irregu-
larly crenate-serrate, serrations 1.5-7.5 mm. wide, 0.5-3.2 mm. high,
5-7 pairs of primary veins, 2 (-3) basal, second basal pair extending
about half way up the lamina.

Inflorescences terminal, extending 2-—6.5 cm. beyond the first leaf,
and axillary, 2-4 cm. long, in the axils of the upper 1-2 leaves,
corymbiform, the ultimate divisions umbellate, 3-9 flowers per umbel,
the peduncle 0.3-2.7 cm. long, with a primary branch 0.3-2.0 em.
long, and a secondary one 0.3-1.2 cm. long, the pedicels 5-13 mm.
long at anthesis, becoming 8-15 mm. long in the fruiting stage,
covered with stellate hairs 0.5 mm. long; bracts 3.2-6.2 mm. long,
0.2-0.7 mm. wide, lanceolate to nearly linear, densely covered with
stellate hairs up to 0.6 mm. long.

Calyx rose, densely covered with siellate hairs 0.2-0.5 mm. long,
at anthesis 10-13.3 mm. long, 6.2-8.5 mm. wide, with teeth 2.2-3.7
mm. long, 2.3-3.0 mm. wide, accrescent, becoming 1.5-2.3 cm. long,
1-1.4 cm. wide, with teeth 5 mm. long, 3 mm. wide, the teeth deltoid-
acute to deltoid-subacuminate, the sinus acute, inflated calyx acutely
pentagonal, reticulate-veined and with a midrib leading to each
tooth and a vein to each sinus; glandular tissue extending for 0.5 mm.
between the calyx and corolla and 0.8—1.0 mm. high between the bases
of the petals; petals light blue, red, mauve or violet, 16-17.4 mm.
long, 5.6-9.5 mm. wide, obdeltoid-spatulate, slightly emarginate at
the apex, the claw 2.5-4.0 mm. long, somewhat thickened, a few
simple, forked, and stellate hairs 0.1-0.2 mm. long on the abaxial
surface near the base, scattered glandular hairs 0.1 mm. long on the
adaxial surface.

Longistylous form. Stamens 5.3-6.8 mm. long, the filaments free
for 1.0-1.4 mm., a few forked hairs 0.2 mm. long at the base of the
staminal tube, staminodes none, or rarely extending 0.1 mm. beyond
the staminal tube, the anther 1.6-1.8 mm. long, 0.7-1.0 mm. wide,
oblong, slightly emarginate at both ends, slightly more so at the base;
pistil 6.2-6.7 mm. long, the styles 4.0-5.5 mm. long, free to the ovary,
papillose and slightly clavate for 0.7-1.0 mm. at the apex, forked and
stellate hairs 0.2 mm. long at the base of the styles, the ovary globose,
pentagonal, covered with stellate hairs 0.1-0.2 mm. long, the stipe
2.8-3 mm. long.

Brevistylous form. Stamens 8.3-8.5 mm. long, the filaments free
for 3.4-3.7 mm., the anther 1.6-1.9 mm. long, 0.7-0.8 mm. wide,
oblong, slightly narrower at the apex, emarginate at both ends or
only at the base, more so at the base, no hairs on the androecium,
staminodes none, or rarely extending 0.1 mm. above the staminal

227-051—67— 11

350 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

tube, dentiform; pistil 3.5 mm. long, the styles 0.8-1.0 mm. long, free
to the ovary, the ovary densely covered with stellate hairs 0.2 mm.
long, the stipe 2.5 mm. long.

Fruit 6.4-7.5 mm. long, 6.8-8.8 mm. wide, globose, rather acutely
pentagonal, slightly retuse at the apex, the sulci between the carpels
fairly deep, extending half way to the center of the fruit, covered with
stellate hairs 0.1-0.4 mm. long, the stipe 5 mm. long; dehiscence
loculicidal along the dorsal suture; seeds trigonal, or somewhat
flattened, 2.8-3.9 mm. long, 1.6—2.5 mm. wide, brown.

Flowering specimens collected in June, July, September and Octo-
ber; fruiting ones, in June and July.

Type Locatity: The species was described in 1854 from specimens
grown in the Berlin Botanical Garden. The native locality was
unknown to the authors. A specimen, cited below, labeled Melochia
rhodocalyr was received by the Riksmuseum, Stockholm, Sweden,
from the Berlin Botanical Garden in the above year. It may be an
isotype.

DistrisuTion: Mexico to Oaxaca, Mexico, in wet woods, on lava
beds, at the edge of a stream, on clay soil, at 1,200-1,650 m ters
elevation.

Cultivated in the Berlin Botanical Garden Andersson s.n. (S,
isotype, also US neg. 5610).

MEXICO: Morertos: Near Cuernavaca, Rose & Painter 6949 (US); Pringle
6335 (MICH, 8, US, W). Xochiltepec, Lyonnet 2164 (US); Lyonnet & Elcoro
1136 (US). Yautepec, Rose et al. 8548 (US). Xochicaleo, Kenoyer July 18,
19388 (MICH). Mexico: TemascaLtTEPsEc: Acatitl4n, Hinton 4338 (S, US).
GuERRERO: Limon Mt., Rusby 18 (US). Campo Morado, Mina, Hinton et al.
14472 (US). El Ocote, probably near Chapultepec according to MceVaugh (40),
Langlassé 1046 (US). Oaxaca: Jayacatlan, L. C. Smith 14 (US). Six miles
above Dominguillo, Nelson 1847 (US).

Hemsley (25) stated, ““Physodium does not appear to differ generi-
cally from Melochia; but J have followed Bentham and Hooker in
referring this plant to Presl’s genus.”” Hemsley used Andrieux no.
507 (K), collected between Acatlin and Chila, Puebla, Mexico, as
the type.

Schumann (48) considered Physodium Presl to be a synonym of
Melochia L.

The petals of this species exceed the calyx in length at anthesis,
those of M. adenodes A. Goldberg do not. The petals of M. corymbosa
(Presl) Meissn. ex Steud. are considerably shorter than those of
M. rhodocalyz.

54. Melochia corymbosa (Presl) Meissn. ex Steud. Nom. Bot. II, 2:123. 1841.
Physodium corymbosum Presl, Rel. Haenk. 2:150, pl. 72. 1835.

A sparsely branched frutex to small tree 4.5-6.0 meters high, the

GOLDBERG—GENUS MELOCHIA 351

young stems and petioles tomentulose, hairs yellowish brown, stellate,
).2-0.6 mm. long, internodes 1.5-3.5 cm. long; stipules 4 mm. long,
).7 mm. wide, deltoid-acuminate, densely pubescent, hairs stellate,
).3 mm. long; petiole 1-2 cm. long.

Lamina 4.8-9.0 cm. long, 3.3-7.0 cm. wide, broadly ovate, oblique,
‘he base rounded to subcordate, the apex acute to acuminate, scabrous
or woolly to the touch, the upper surface greenish, covered with
itellate hairs 0.2-0.5 mm. long, the lower surface lighter green, tom-
mtulose, hairs stellate, 0.3-0.6 mm. long, the margin irregularly
renate-serrate, serrations 1-6 mm. wide, 0.5-2.3 mm. high, 5-7
yairs of primary veins, 2 or 3 are basal.

Inflorescences terminal, extending 2.5-5.5 cm. beyond the first
leaf, corymbiform, the ultimate divisions umbellate, about 5 flowers
per umbel, sometimes also axillary, 1.5 cm. long, the peduncle 1 cm.
long with a primary branch 0.8-1.3 cm. long, a secondary one 0.4 cm.
long, and sometimes a tertiary one 0.2 cm. long, the pedicel 2.5-3.5
mm. long, becoming 5-13 mm. long in the fruiting stage; bracts
deciduous, 4.5 mm. long, 0.4 mm. wide, nearly linear, with stellate
hairs 0.2-0.3 mm. long.

Calyx 7 mm. long, 3.6 mm. wide, the teeth 1.7-1.8 mm. long,
1.5 mm. wide, deltoid-acute, accrescent, becoming membranous,
acutely pentagonal, 1.7-2.3 cm. long, 0.8-1.6 cm. wide, the teeth
3-5 mm. long, 5 mm. wide, densely pubescent, hairs stellate, 0.1-
0.8 mm. long, the sinus acute; petals 9.7 mm. long, 2.1 mm. wide,
oblanceolate, with scattered glandular hairs 0.1 mm. long on the
axial surface and a line of stellate hairs 0.2-0.7 mm. long on the
midrib, or a few simple hairs at the base, of the abaxial surface, the
claw 2.5 mm. long.

Longistylous form. Stamens 5.7 mm. long, the filaments free for
0.4 mm., staminodes none, the anther 1.4 mm. long, 0.6 mm. wide;
pistil developed to fruiting stage, the styles 3.5 mm. long, free to the
ovary.

Brevistylous form. Stamens 7 mm. long, the filaments free for
3.5-4.0 mm., the anther 1.7-1.9 mm. long, 0.7-0.8 mm. wide; pistil
3.2 mm. long, the styles 1.3 mm. long, free to the ovary, papillose for
0.6 mm. at the apex, the ovary densely covered with stellate hairs
0.2 mm. long, the stipe 1 mm. long.

Fruit 5.3-7.2 mm. long, 6.7-9.0 (-10) mm. wide, globose, acutely
pentagonal, the angles without points, the apex retuse, the sulci
between the carpels extending a third to half way to the center of the
fruit, tomentulose, hairs stellate, 0.3-0.5 mm. long, the stipe 4.3-6.0
mm. long; dehiscence loculicidal along the dorsal suture; seeds 3-3.7
mm. long, 1.6-2.4 mm. wide, dark brown, 1 or 2 per cell.

Flowering and fruiting specimens collected in July.

352 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

TypE LocaLity: The type, cited below, was collected by Haenke
in western Mexico.

DistrispuTion: ‘Western Mexico” and Oaxaca, Mexico, at 750
meters elevation.

MEXICO: ‘Western Mexico,” Haenke (PR 174400a, b, c, d, type and also US
neg. 5554); ?Haenke (W). Oaxaca: Tomellin Canyon, Pringle 4751 (S, US).

This species differs from Melochia adenodes A. Goldberg in the ab-
sence of glandular hairs on the calyx and pedicels, and in the absence
of staminodes. It differs from Melochia rhodocalyz C. Koch & Bouché
in having considerably smaller petals.

New Taxa, New Names, and New Combinations

Melochia L., section Physodium (Presl) A. Goldberg, comb. et stat. nov.

Melochia L., section Pyramis A. Goldberg, nom. nov.

Melochia adenodes A. Goldberg, sp. nov.

Melochia adenodes A. Goldberg var. acuminata (Rose) A. Goldberg, comb. nov.

Melochia chamaedrys St.-Hil. var. decumbens (R. E. Fries) A. Goldberg, comb. et
stat. nov.

Melochia thymifolia (Presl) A. Goldberg, comb. nov.

Melochia tomentosa L. var. speciosa (S. Wats.) A. Goldberg, comb. et stat. nov.

Melochia villosa (Mill.) Fawe. & Rendle var. regnellii (Schum.) A. Goldberg,
comb. nov.

Melochia villosa (Mill.) Fawe. & Rendle var. tomentosa (Schum.) A. Goldberg,
comb. nov.

Melochia werdermannii A. Goldberg, sp. nov.

Excluded and Doubtful Taxa

Altheria Madagascariensis DC. Prodr. 1:493. 1824.

The description is inadequate for a determination.
Antiphylla serrata Rafin. New Fl. Amer. 4:16. 1836.

Antiphylla Rafin. is attributed to Melochia in Index Kewensis (30) and by
Dalla Torre & Harms (7), but it probably belongs to the Capparidaceae or Cruci-
ferae.

Melochia acutangula Span. in Linnaea 15:174. 1841.

The description is inadequate for a determination.

Melochia Berteriana Balb. ex DC. Prodr. 1:492. 1824, is a Waltheria according
to Index Kewensis (30).
Melochia Borbonica Cav. Diss. 6:321. 1788.

The illustration is very reminiscent of Melochia nodiflora, but the description
does not correspond to that species.

Melochia capitata, Index Kewensis (30), erroneously attributed to Boj. Hort.
Maurit. 38. 1837.

Melochia corymbosa (DC.) Hemsl. Biol. Centr. Amer. Bot. 1:1380. 1879= Riedleia
corymbosa [Moc. & Sessé] ex DC., which see.

Melochia depressa L. Sp. Pl. 674. 1758.

I have examined a microfiche photograph, taken by the International Documen-
tation Centre, of a specimen in the Linnaean Herbarium so labeled by Solander.
The specimen appears to belong to the Malvaceae.

GOLDBERG—GENUS MELOCHIA 353

Melochia hirsuta Turez. Bull. Soc. Nat. Mose. 36(1):570. 18638.
The description is inadequate for a determination.
Melochia hispida Hook. & Arn. Bot. Beech. Voy. 60. 1841.
The description is inadequate for a determination and the type cannot be

located.
Melochia mollissima Desf. Tabl. ed. I, 148. 1804, nomem nudum; Pers. Syn.

2:218. 1807.

A small fragment ex P so labeled belongs to the Malvaceae. The description
in Persoon is inadequate for a determination.

Melochia nipensis Britt. Mem. Torr. Bot. Cl. 16:81. 1920, is a Waltheria.

Melochia ovata Desf. Tabl. ed. II, 172. 1815= Melhania abyssinica according to
Index Kewensis (80).

Melochia peruviana Desr. in Lam. Encyc. 4:83. 1797, based on Melochia depressa
L. in Cav. Diss. 6:320, pl. 173, fig. 1. 1788, appears to be close to Melochia
chamaedrys St.-Hil., but unlike the latter, it has only one flower per leaf axil
and the leaves are tomentose.

Melochia pulverulenta Miers, Proc. Roy. Hort. Soc. 4:185. 1865.

The description is inadequate for a determination.

Melochia rotundifolia Sessé & Moc. Pl. N. Hispan. 1:106. 1889, is a Plerostemon.

Melochia trigyna L. ex Jackson, Index Linn. Herb. 103. 1912, nomen nudum.

Melochia truncata Wall. Cat. No. 1196 K. 1929, is a Sida according to Index
Kewensis (30).

Melochia truncata Willd. Sp. Pl. 3:601. 1801=Melochia corchorifolia L. according
to Index Kewensis (30), but the obcordate, cuneate lamina, monopetalous
corolla, 10 stamens, and reticulate seeds of the description are not M. cor-
chorifolia characters.

Melochia varrontfolia Desf. Cat. Hort. Par. III, 252. 1829, nomen nudum.

Physocodon brachybotrys Turez. Bull. Soc. Nat. Mose. 31(1):2138. 1858.

The description is inadequate for a determination.

Riddelia antiphylla Rafin. New Fl. Amer. 4:16. 1836, is attributed to Melochia
hirsuta Cav. in Index Kewensis (30), but the fruit is too long to be that of
a Melochia and the seeds differ in shape from those of Melochia.

Riedleia Berteriana DC. Prodr. 1:492. 1824, is a Waltheria according to Index
Kewensis (30).

Riedleia Borbonica (Cav.) DC. Prodr. 1:491. 1824= Melochia Borbonica Cav.

Riedleia capitata Boj. Hort. Maurit. 38. 1837, nomen nudum.

Riedleia corymbosa [Moc. & Sessé] ex DC. Prodr. 1:491. 1824.

The description is inadequate for a determination. The petals are stated to
be twice the size of those of Riedleia odorata (L.f.) DC. Prodr. 1:491. 1824.
Very few species of Melochia have petals that large.

Riedleia depressa (l.) DC. Prodr. 1:491. 1824= Melochia depressa L.

Riedleia eriantha DC. Prodr. 1:492. 1824, belongs to the Malvaceae according to
Schumann (47).

Riedleia odorata DC. Prodr. 1:491. 1824= Melochia odorata L.f. and Melochia
umbellata (Houtt.) Stapf.

Riedleia Peruviana (Desr. in Lam.) DC. Prodr. 1:491. 1824= Melochia peruviana
Desr. in Lam.

Riedleia setulosa Miq. Hoev. & De Vriese Tijdschr. 10:80. 1843 probably is a
Sida.

304 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Riedleia truncata (Willd.) DC. Prodr. 1:491. 1824= Melochia corchorifolia L.
according to Index Kewensis (30), but De Candolle stated that the flower
is solitary and pedunculate whereas M. corchorifolia has a capitate inflores-
cence.

Riedleja hyptotdes Turcz. Bull. Soc. Nat. Mose. 31(1):210. 1858.

The description is inadequate for a determination.

Riedleja pteronema Turez. Bull. Soc. Nat. Mose. 31(1):208. 1858.

The description is inadequate for a determination.

Riedleja ramosissima Turcz. Bull. Soc. Nat. Moscow 36(1):571. 1863.

The description is inadequate for a determination.

Riedleja stphonandra Turez. Bull. Soc. Nat. Mose. 31(1):208. 1858.

The description is inadequate for a determination.

Turnera paniculata Willd. in Schultes Syst. Veg. 6:678. 1820=Melochia gram-
inifolia St.-Hil., according to Schumann (47), but the description in Schultes
is inadequate for a determination.

Visenia Berteriana (DC.) Spreng. Syst. 3:30. 1826, isa Waltheria according to
Index Kewensis (30).

Visenia corymbosa (DC.) Spreng. Syst. 3:29. 1826= Riedleia corymbosa [Moc. &
Sessé] ex DC.

Visenia depressa (L.) Spreng. Syst. 3:30. 1826= Melochia depressa L.

Visenia ferruginea Colla, Herb. Pedem. 1:432. 1833.

I have seen two specimens from the Colla Herbarium in a folder marked Visenia
ferruginea with descriptions on the specimen sheets which could be interpreted
as corresponding to the species description of V. ferruginea. One is a Waltheria
and the other belongs to the Turneraceae.

Visenia madagascarensis Spreng. Syst. 3:30. 1826.

The description is inadequate for a determination.

Visenta scabra Colla, Herb. Pedem. 1:431. 1833.

The description is inadequate for a determination.

Visenia spicata Colla, Herb. Pedem. 1:431. 1833, belongs to the Malvaceae.

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1959. Regnum vegetabile, vol. 15. Index herbariorum. lt. The herbaria
of the world, ed. 4, pp. 188-211. Utrecht.

5. Lemiée, A.

1952. Flore de la Guyane francaise 2:377-378. Brest.

. Leon, H., and Anarn, H.

1953. Flora of Cuba 3:279-282. Habana.

. Linnagus, C.

1753. Species plantarum, pp. 674-675. Stockholm.
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. Macsripg, J. F.

1956. Flora of Peru. Field Mus. Bot., vol. 13, part III A, No. 2, pp.
623-629.
McVauau, R.
1951. The travels and botanical collections of Eugene Langlassé in Mexico
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ManaGeEnot, S., and ManceEnot, G.
1958. Deuxitme liste de nombres chromosomiques nouveaux chez diverses
dicotylédones et monocotylédones d’Afrique occidentale. Bull.
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1923. An. enumeration of Philippine flowering plants 3:46-47. Manila.
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1947. Catalogo de la flora Venczolana 2:1386-137. Caracas,
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60.

61.

62.

63.

64.

GOLDBERG—GENUS MELOCHIA 357

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Index

(Synonyms in ttalics. Page numbers of priucipal entries in boldface.)

Aleurodendron, 207

album, 220

Altheria, 207

Madagascariensis, 352

Anamorpha, 192, 209, 240

melochioides, 259
waltherioides, 264, 266

Antiphylla, 352

serrata, 352

Buettnerieae, 202
Capparidaceae, 352
Corchorus javonicus, 304

olitorius, 191

Cruciferae, 352
Dombeyeae, 202
Eriolaeneae, 204
Eumelochia, 192, 308
Fremontieae, 202, 204
Geruma, subtriloba, 304
Glossospermum, 207

cordatum, 220
velutinum, 220

Helictereae, 202, 204
Hermannia, 191, 204
Hermannieae, 191, 202, 204
Hermanninae, 204
Hibiscus donii, 304

endlicheri, 304

Hypericum pentandrum, 220
Lasiopetaleae, 202
Lochemia, 209

corchorifolia, 304

Malvaceae, 352, 353, 354
Melhania abyssinica, 353
Melochia, 191, 192, 193, 194, 195, 197,

198, 199, 200, 202, 204, 205, 206,
207, 208, 210, 211, 212, 215, 279,
350, 352, 353
acutangula, 352,
adenodes, 196, 202, 208, 217, 346,
347, 348, 350, 352,
acuminata, 347, 348, 352
adenodes, 202, 347
affinis, 304

Melochia—Continued
anomala, 196, 202, 208, 217, 311,
315, 318, 319, 321
arborea, 220
arenosa, 196, 202, 208, 215, 295, 297
argentina, 196, 208, 214, 253
genuina, 253
lanceolata, 253, 255
arida, 329, 330, 332
aristata, 197, 208, 212, 226, 228,
229, 230
Benthami, 255, 257
bernoulliana, 196, 208, 213, 269
Berteriana, 352
betonicaefolia, 242
betonicifolia, 196, 208, 217, 322,
323, 324, 326
betsiliensis, 302, 304
Borbonica, 352, 353
bracteosa, 197, 202, 208, 215, 291
Burmanni, 304
capitata, 352
caracasana, 196, 208, 217, 324, 326,
327
carpinifolia, 271
carthaginensis, 255
cephalodes, 283, 286
chamaedrys, 196, 202, 208, 214,
246, 248, 250, 251, 353
decumbens, 251
typica, 250
chamaedrys, 202, 250
decumbens, 250, 251, 352
loretensis, 250
viscidula, 250
cinerea, 295, 297
discolor, 295
Clinopodium, 283
colombiana, 196, 208, 213, 261
compacta, 197, 208, 212, 236, 240
concatenata, 304
concinna, 293
conglobata, 271, 272
359

360

Melochia—Continued

corchorifolia, 191, 196, 197, 198,
202, 205, 208, 211, 215, 280, 302,
303, 304, 305, 308, 358, 354

cordata, 304

cordiformis, 328, 324

corumbensts, 295, 297

corymbosa, 196, 208, 217, 345, 350,
352

crenata, 196, 208, 211, 216, 329, 335

crinila, 291

decandra, 321

decumbens, 251

degeneriana, 197, 208, 211, 213,
231, 233, 234, 235

densiflora, 283

depressa, 352, 358, 304

diffusa, 335

domingensis, 338

erecta, 304

fasciculata, 315

frutescens, 332

gardneri, 196, 208, 214, 257, 258

glandulifera, 301, 302

globifera, 259, 260

gracilis, 242

graminifolia, 196, 208, 215, 299, 354

grayana, 197, 208, 212, 235

hassleriana, 196, 208, 215, 246, 248

hermannioides, 196, 208, 211, 216,
310, 311, 312, 315, 319

lacinulata, 310, 312
heterophylla, 310
typica, 310

lanceolata, 310, 312

hirsuta, 282, 353

calophylla, 287, 288

glabrata, 283

glabrescens, 283

grandiflora, 283

macrophylla, 287

paraguayensis, 283

Regnellii, 287

rotundifolia, 283

tomentosa, 287
albiflora, 287
ferruginea, 287
minus induta, 287

hirsulissima, 238

hispida, 353

Humboldtiana, 251, 253

hypoleuca, 329

illicioides, 196, 208, 216, 321, 322

INDEX

Melochia—Continued

indica, 220
deglabrata, 220
inflata, 264
interrupta, 242
Jamaicensis, 242
jurgensenii, 283
kerriifolia, 196, 208, 214, 251, 253
lacinulata, 310, 311
lanata, 196, 208, 216, 326
inclusa, 326, 327
lanceolata, 196, 197, 208, 215, 302,
304
Lessoniana, 238
leucantha, 196, 208, 213, 262
lilacina, 283, 286
littoralis, 295, 296, 297, 322
llanorum, 295, 296
longebracteolata, 304, 306, 308
longepetiolata, 197, 208, 212, 228
lupulina, 196, 208, 213, 261, 264,
267, 269, 271
macrophylla, 324, 326
manducata, 196, 208, 215, 295, 301
melissifolia, 196, 197, 208, 215, 292,
293
brachyphylla, 291
bracteosa, 291
microphylla, 291
mollis, 291
Welwitschti, 291
Missionum, 297, 299
mollipila, 197, 208, 213, 232, 233,
234, 235
mollis, 196, 208, 213, 258, 262, 263,
266, 267, 291
mollissima, 353
morongii, 196, 208, 216, 315, 316,
317, 318
denudata, 316, 317
nepetoides, 323, 324
nervosa, 242, 267
nipensis, 39d
nitidula, 242
nodiflora, 196, 208, 214, 269, 271,
276, 277, 279, 305, 352
nudiflora, 196, 208, 214, 269, 274,
277
oblonga, 297, 299
obovata, 313, 314
odorata, 197, 208, 212, 213, 223,
224, 226, 228, 230, 353
Schlechteri, 220

INDEX 361

Melochia—Continued

ovata, 358
paniculata, 242
parviflora, 245
parvifolia, 196, 208, 216, 311, 312,
3138, 315, 317, 318, 319
albiflora, 315
roseiflora, 310
Esperanzae, 313
fasciculata, 313, 315
microphylla, 315
transiens, 315
typica, 315
genuina, 313
induta, 313
Morongii, 316
genuina, 316
denudata, 316
intermedia, 316
tomentosa, 316
normalis, 313
parvifolia, 313
pauciflora, 304
peruviana, 353
pilosa, 196, 208, 215, 241, 246, 248,
250, 251, 253, 258
plicata, 329, 330
polystachya, 242
portoricensis, 329
pseudonodiflora, 267, 269
plerocarpa, 334, 335
pulverulenta, 353
pyramidata, 193, 196, 197, 208, 211,
217, 309, 332, 337, 338, 343
flava, 343
Grisebachii, 343
hieronymi, 338, 341, 343
foliis luteo variegatus, 343
transitoria, 343
normalis, 338
paraguariensis intermedia, 343
pseudotomentosa, 343
pyramidata, 338
ramuliflora, 255, 257
rhodocalyx, 196, 208, 217, 348,
350, 352
roseiflora, 197, 208, 212, 234, 235
rotundifolia, 353
savannarum, 196, 208, 215, 287, 288
scordiifolia, 313, 314
scutellarioides, 283
sericea, 242, 245
serrata, 283

Melochia—Continued
simplex, 196, 208, 215, 297, 301
sorocabensis, 297, 299
speciosa, 334, 335
splendens, 196, 208, 215, 287, 290,
292
stricta, 297, 299
Missionum, 297
subcordata, 287, 288
supina, 304
tenella, 2838
thymifolia, 196, 208, 214, 267, 352
tomentella, 193, 196, 208, 214, 277
tomentosa, 196, 197, 208, 217, 324,
326, 327, 329, 330, 332, 335, 336,
342, 343
Bahiensis, 329
frutescens, 329, 330, 332
mattogrossensis, 343
speciosa, 329, 334, 352
tomentosa, 329, 330
turpiniana, 629, 234
typica, 329
tragiifolia, 196, 208, 213, 276
trigyna, 353
truncata, 353
Turpiniana, 334
ulmarioides, 242, 246, 248
albo-villosa, 246, 248
brevipedunculata, 246, 248
Hassleriana, 246
sericea, 246, 248
ulmifolia, 196, 208, 214, 255, 257, 258
umbellata, 193, 197, 202, 208, 212,
219, 220, 223, 353
urticaefolia, 271
varronifolia, 353
velutina, 220
venosa, 241, 244
betonicifolia, 242, 245
polystachya, 242
flava, 242
sericea, 242
squarrosa, 242, 246
typica, 242
purpurea, 267
Widgrenii, 242
vestita, 283, 285, 286
villosa, 196, 208, 215, 280, 282, 290,
291, 292, 295
regnellii, 282, 287, 352
tomentosa, 282, 286, 287, 352
villosa, 282

362

Melochia—Continued
villosissima, 197, 208, 212, 238, 240
vitiensis, 197, 208, 213, 226, 228,
230
B, 235, 236
Welwitschit, 291
werdermannii, 196, 208, 216, 319,
352
yungasensis, 267, 269
Melochinae, 204
Meluchia, 207
Moluchia, 191, 207
domingensis, 338
frutescens, 332
fruticosa, 338
herbacea, 338
pyramidata, 338
tomentosa, 329
Mougeotia, 191, 192, 193, 205, 206, 207,
211, 218, 240, 255, 257, 258,
267, 312
caracasana, 324
corchorifolia, 304
hirsuta, 282
inflata, 264, 266
mollis, 258, 260
nodiflora, 271
polystachya, 241, 244
Pentaglottis tomentosa, 220, 222
Physocodon, 192, 209
brachybotrys, 353
macrobotrys, 264, 266
Physodium, 192, 193, 205, 206, 209,
211, 217, 344, 350, 352
corymbosum, 348, 350
acuminata, 348
dubium, 348
Polychlaena, 207
ramosa, 304
simplex, 304
Pterostemon, 353
Pyramis, 191, 193, 205, 206, 211, 216,
308, 312, 327, 352
Riddelia antiphylla, 353
Ridleia, 209
hypoleuca, 329
Riedlea, 191, 192, 207
glabrescens, 283
serrata, 282
villosisstma, 238
Riedleia, 191, 192, 207, 279
Aleurodendron, 220
argentea, 242

INDEX

Riedleia-——Continued
Berteriana, 353
Borbonica, 353
capitata, 353
Caracasana, 324
Chamaedrys, 250
concalenata, 304
concinna, 293
corchorifolia, 304
cordata, 220
corymbosa, 352, 353, 354
crenata, 335
depressa, 353
elongata, 283
ertantha, 353
graminifolia, 299
hirsuta,'283
inflata, 264
interrupla, 242, 244
J amaicensis, 241
littoralis, 295
lupulina, 264
melissaefolia, 264, 265
mollis, 258
multiflora, 258, 261
nodiflora, 271
odorata, 353
Peruviana, 353
polystachya, 241
radiata, 304
ramuliflora, 255, 257
sericea, 242
serrata glabrescens, 283
setulosa, 353
sparsiflora, 293
supina, 304
thymifolia, 267
tiliaefolia, 220, 223
tomentella, 277
truncata, 354
Ulmariae, 242
velutina, 220
venosa, 242
virgultosa, 242
Wallichtana, 220

Riedleja, 209
Cubensis, 283
dichotoma, 255, 256, 257
heterotricha, 283, 285
hyptoides, 354
Jurgensenit, 283
pteronema, 354
ramosissima, 354

INDEX

Riedleja—Continued
scutellarioides, 283
siphonandra, 354
tenella, 283
urticaefolia, 271, 273

Riedlia, 207

Sida, 353
pilosa, 241
rajoides, 329
Sabeana, 338
villosa, 282

Sterculiaceae, 202, 204

Sterculieae, 202, 204

Turnera paniculata, 354

Turneraceae, 354

Visenia, 191, 192, 206, 207, 212, 218,

229, 236, 255, 257
alba, 220
Berteriana, 354
caracasana, 324
concatenata, 304
corchorifolia, 304
corymbosa, 354
crenata, 335

Visenia—Continued
depressa, 354
ferruginea, 354
graminifolia, 299
hirsuta, 283
inflata, 264
Javanica, 220
lupulina, 264
madagascarensis, 354
mollis, 258
nodiflora, 271
polystachya, 242
scabra, 354
sericea, 242
serrata, 283
spicata, 354
supina, 304
tomentosa, 220
Ulmariae, 242
umbellata, 191, 220
velutina, 220
venosa, 242

Waltheria, 191, 204, 352, 353, 354

Wisenia indica, 220

U.S. GOVERNMENT PRINTING OFF.CE: 1967

363

CONTR. NAT. HERB. VOL. 34 GOLDBERG —PLATE 1

SL AA
AFL,

Melochia gardneri Sprague. Gardner 3608 (P, isotype).

CONTR. NAT. HERB. VOL. 34 GOLDBERG—PLATE 2

> UNIVERSITY OF CALIFORNIA
Second Botanical Garden Expedition to the Andes, 1058 30

Prev, Botivta, Cmte, | ay, AROENTINA

’ soe ’

Melochia leucantha Macbride. Stork & Horton 10908 (UC, type).

CONTR. NAT. HERB. VOL. 34 . GOLDBERG—PLATE 3

4

7 TYPUS

Melochia thymifolia (Presl) A. Goldberg. Haenke (PR 174403 a, lectotype)

CONTR. NAT. HERB. VOL. 34 GOLDBERG—PLATE 4

av

TYPUS

Melochia tomentella (Pres!) Hemsl. /7Taenke (PR 174404 ¢, isotype).

CONTR. NAT. HERB. VOL. 34 GOLDBERG—PLATE 5

a
Melochia villosa (Mill.) Fawe. & Rendle. Apical portion of flowering plant, approximately

twice natural size.

GOLDBERG—PLATE 6

CONTR. NAT. HERB. VOL. 34

a

ae
nen .

Flare PLORAP BRASTLEAR meridionatic
Mausiro Pavisiensi dedit Awg DE St. HILAIRE

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Melochia graminifolia St.-Hil. St.-/Tilaire 1589 (P, isotype).

CONTR. NAT. HERB. VOL. 34

*L; he ea Vos Lee

are

f
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grcteras Percberye Bee Mere n Cee
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Melochia anomala Griseb.

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nance

GOLDBERG—PLATE 7

i

fieke

Lorentz 503 (GOET, type).

CONTR. NAT. HERB. VOL.. 34 GOLDBERG—PLATE 8

Reise durch Ost-Bolivia Nr. C6C#

Dr. E. Werdermann

J

\.

Melochia werdermannii A. Goldberg. WW erdermann 2606 (S. type).

CONTR. NAT. HERB. VOL. 34 GOLDBERG—PLATE 9

Abelochow

Melochia rhodocalyx C. Koch & Bouché. Andersson s.n. (S, perhaps isotype).

BULLETIN OF THE UNITED STATES NATIONAL MUSEUM

ConTRIBUTIONS From THE Unitep States Natrona, HERBARIUM

Vo.LumE 34, Part 6

MEXICAN “CACALIOID” GENERA ALLIED TO
SENECIO (COMPOSITAE)

By Ricnarp W. Pippen

SMITHSONIAN INSTITUTION PRESS WASHINGTON, D.C. e 1968

MEXICAN “CACALIOID” GENERA ALLIED TO
SENECIO (COMPOSITAE) !

By Ricuarp W. Pieren ?

Introduction

This paper represents an investigation of a geographically distinct
group of North American species of Compositae occurring mainly
in the highlands of Mexico. In the past these species have been con-
sidered part of the genera Cacalia (DeCandolle, 1838; Gray, 1883;
Greenman, 1902), Senecio (Schultz 1845; Bentham & Hooker, 1873;
Hemsley, 1881), and several distinct, smaller genera (Rydberg, 1924;
Belcher, 1956; Cuatrecasas, 1960). In my study I worked out a taxo-
nomic system for these species, then compared the species individually
and as a group with superficially similar species of Senecio, and con-
sidered their relationships. It is now apparent that these species are
generically distinct from Senecio, from the species in eastern North
America and Asia usually referred to Cacalia, and also from the
type-species of Cacalia; instead, four separate genera are represented—
Digitacaha, Odontotrichum, Pericalia, and Psacalium.

The materials used in this study consisted primarily of specimens
obtained from the following herbaria: Field Museum of Natural His-
tory, Chicago; Gray Herbarium of Harvard University, Cambridge,
Massachusetts; Botanische Staatssammlung, Miinchen; University of
Michigan, Ann Arbor; New York Botanical Garden, New York;
University of Texas, Austin; and United States National Herbarium,
Smithsonian Institution. I wish to express my gratitude to the cura-
tors of these herbaria who made these specimens available. Abbrevi-
ations of herbarium names used hereafter are those of Lanjouw and
Stafleu (1964). The abbreviation “Field Mus. Neg.’ refers to Field
Museum of Natural History negatives of a series of type photographs
taken in European herbaria by J. F. Macbride during the period 1929
to 1939. “U. of MICH. Neg.” refers to a collection of negatives of
type specimens maintained at the University of Michigan Herbarium.

1 Part of a dissertation presented to the Horace H. Rackham School of Grad-
uate Studies, the University of Michigan, in partial fulfillment of the requirements
for the degree of Doctor of Philosophy.

2 Department of Biology, Western Michigan University.

365

366 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Observations were made in the field, and preserved cytological
materials and dried specimens were obtained during two seasons of
field work in Mexico (in the summer and autumn of 1958, and the
autumn of 1960), and from a survey trip through the southeastern
United States in the summer of 1962.3 Observations were also made
of living plants grown from seeds and rootstocks in the University
of Michigan Botanical Gardens.

All measurements were obtained from dried specimens. But flower-
ing material of all the species was studied both in the dry state and
after soaking, the latter state inducing no significant change in size.
Dimensions of the heads include from the base of the phyllaries to
the tip of the syngenesious anthers.

The chromosome counts were obtained by the standard aceto-
carmine squash technique on pollen spore mother cells killed and
fixed in Newcomer’s solution. Except for Odontotrichum decompositum
the chromosome numbers herein reported are new reports. Table 1
summarizes this information. Voucher specimens for these counts are
listed in table 1 and also indicated with an asterisk (*) in the list of
specimens examined under the species. Voucher specimens are pre-
served in the University of Michigan Herbarium.

For advice and assistance my sincere thanks and appreciation
are extended to Dr. Rogers McVaugh, who initially called my atten-
tion to this problem and who collected specimens and cytological
material for me on several occasions; to Dr. B. L. Turner, the Uni-
versity of Texas, for permitting me to cite unpublished chromosome
counts from his cytological investigations of the Mexican composites;
to the late Mr. N. Y. Sandwith, the Royal Botanic Gardens, Kew,
who sent me special information regarding the type of Cacalia cir-
stifolia Hook. & Arn.; to Dr. A. G. Norman, formerly Director of
the University of Michigan Botanical Gardens, for the use of the
facilities there for growing living plants; to Mr. J. R. Millar, formerly
Chief Curator of the Herbarium, Field Museum of Natural History,
for making available the botanical collections of Sessé and Mocifio
(then on loan to Chicago from Madrid); and to Dr. Charles Feddema,
U.S. Forest Service Herbarium, for the line drawings of flower types.

Morphological Considerations

Although these species, previously referred to Cacalia, do apparently
represent several genera, they have in common a number of morpho-
logical features and generalizations regarding these may be made.
Except for Digitacalia (here described as new) all the genera in the

3 Made possible by a grant from the Graduate Student Research fund of the
Horace H. Rackham School of Graduate Studies, the University of Michigan.

PIPPEN—MEXICAN “CACALIOID” GENERA 367

TABLE 1.—Metotic chromosome counts of some species of Mexican cacalioids

Species Voucher Specimens ! Number

Digitacalia tridactylitis McVaugh 21903 3 30
Odontotrichum amplum Pippen 4 2 30
Odontotrichum brachycomum McVaugh 21934 3 30
Odontotrichum cirsiifoliwm Pippen 52 2 30
Odontotrichum decompositum - 4 30
Odontotrichum multilobum Pippen 30 2 30
Odontotrichum palmeri Pippen 12 ca. ? 25
Odontotrichum pringlei Pippen 37 2 30

MecVaugh 21765 3 30
Odontotrichum sinuatum Pippen 9 3 30
Pericalia michoacana Pippen 63, 65 2 30
Pericalia sesstlifolia Pippen 61 2 30
Psacalium ertocarpum Pippen 62 2 30
Psacalium holwayana McVaugh 21953 3 30
Psacalium megaphyllum Pippen 8 3 30
Psacalium peltatum v. peltatum Pippen 49 2 30
Psacalium species MecVaugh 21913 3 30

1 Voucher specimens retained in the Herbarium of the University of Michigan,
Ann Arbor.

2 Determined by Pippen.

3 Determined by B. L. Turner, here published.

4 Determined by Kruckeberg in Ornduff et al. 1963.

Senecioneae, including the cacalioid ones, are contrasted by Belcher
(1956).

All of these species are herbaceous perennials, flowering from July
to December. The perennial caudex and/or roots survive the dry
season but the aerial portions wither soon after flowering. The first
season of growth is strictly vegetative. A basal rosette of leaves
forms in Odontotrichum and Psacalium; in Digitacalia and Pervealia
a nonflowering, leafy stem develops. Flowering may occur during the
second and succeeding years.

Leaves in all of the genera are petiolate (except certain specimens
of Pericalia sessilifolia). The leaves are usually fleshy or coriaceous
with prominently convex veins usually more pronounced on the
abaxial surface. In several species of Odontotrichum (O. pachyphyllum,
O. platylepis, and O. amplum) the two surfaces of the leaves are
similar with the venation equally pronounced on both surfaces.

The leaf shapes and patterns of lobing are relatively constant, al-
though wide ranges of variation may occur within the limits of some
species (e.g., Odontotrichum sinuatum) and occasional parallel leaf
forms between species (0. silphiifolium and O. goldsmithii). In general,
however, each species may be recognized by a characteristic leaf shape

368 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

and pattern of lobing. The venation is mostly pinnate, subpalmate,
or radiate (the latter in Psacalium only).

In Odontotrichum the form of the leaves exhibits a series ranging
from entire or toothed (as in O. amplum, O. pachyphyllum, and 0.
silphiifolium) to lobed (O. sinwatum, O. pringlei, and O. multilobum)
and pinnatisect (0. decompositum). This series in general parallels a
reduction in the size of heads and size of inflorescence bracts.

The inflorescences are monochasial (Rickett, 1955) with at least 5
heads. They may be corymbose, paniculate, or less frequently race-
mose. The bracts subtending the branches of the inflorescence and the
calyculate bracts are relatively constant within a species and either
small, linear-subulate or larger and leafy. The leafy inflorescence bracts
are usually vaginate or naviculate and clasping the stem or in some
species, petiolate.

Heads are always discoid with perfect and fertile flowers. The size
of the heads, as determined by the number of phyllaries and flowers, is
relatively constant within each species.

The flowers are nearly white or cream colored, often with a greenish
or purplish tint but never yellow or orange. The corolla tube is narrow-
ly cylindrical and usually as long as or longer than the lobes of the
limb. The limb is divided into 5 equal lobes that are usually free to the
base of the limb or, in some species, connate for part of their length
into an ampliate, campanulate, or funnelform throat. The throat is
usually shorter than the lobes or, less frequently, as long as the lobes
(longer than the lobes only in Periealia michoacana).

This condition of the corolla is referred to as the “cacalioid condi-
tion” to distinguish it from that found in Senecio where the corolla
throat is much longer than the lobes, the latter forming only five small,
deltoid, ascending teeth above the throat—referred to as the “sene-
cioid condition”’ (fig. 1).

The achenes vary from ellipsoid, obovoid and dorsally compressed,
elliptic or ovate in cross section (Odontotrichum and Psacalium) to
oblong, subcylindrical, and subterete in cross section (Digitacalia and
Pericalia). The surface is usually narrowly striated or often promi-
nently 10- to 12-ribbed. In several species the achenes are pubescent
with simple unicellular hairs but none of the species has hygroscopic
trichomes like those found on achenes of several species of Senecio.

The pappus consists of a single row of many, persistent (deciduous
in Odontotrichum cirsiifolium), white or cream colored, stiff, barbellate,
capillary bristles (as in most of the Senecioneae). They are usually as
long as the corolla or occasionally much shorter and stiffer (O. bra-
chycomum and O. palmeri), or even entirely lacking (Psacaliwm calvum
and P. nanum).

A study of pollen grains of representative species (mounted in lactic

PIPPEN—MEXICAN “CACALIOID” GENERA 369

Ficure 1.—Types of discoid corollas (all X10): a. Pericalia sessilifolia (Pippen 64, MICH);
b. Psacalium peltatum var. typ. (Pippen 49, MICH); c. Odontotrichum sinuatum
(Pippen 22, MICH); d. Digitacalia jatrophoides (L. C. Smith 123, MICH); e. Senecio

angulifolius (sect. Palmatinerott) (McPaugh 9949, MICH); f. Senecio callosus (sect.
Mulgedifolii) (Hurd 36, MICH).

370 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

acid or Hoyer’s medium) revealed a similarity in all four genera. The
grains are spherical, tricolpate, 42-45 microns in diameter. The surface
is ornamented with simple spines 7 to 9 microns long.

Several species exhibit various degrees of pubescence, the presence
or absence of which on organs such as leaves, achenes, and phyllaries
is relatively constant for each species. The trichomes are mostly non-
glandular, conical-attenuate, white or hyaline, multicellular, frequently
with purple crosswalls. They vary in length, number of cells, stiffness,
and density. Occasionally the distal end of the hair is prolonged into
a caudate tip as long as or longer than the body. A minute stipitate
eland, ca. 0.5 mm. long, consisting of a stalk of several cells in a sin-
ele row, topped with a multicellular, globose, purple gland occurs
mixed among the longer, nonglandular hairs in several species of
Psacalvum.

Distribution and Ecology

The genera Digitacalia, Odontotrichum, Pericalia, and Psacalium
are all endemic to Mexico, except that one species, Odontotrichum
decompositum, extends northward into the mountains of southern
Arizona. The species of all four genera are restricted mostly to the
oak-pine zone, which is extensively developed in the higher mountain
regions of Mexico. Within this zone the species of Odontotrichum and
Digitacalia are usually associated with the grasslands or oak savannahs
and Pericalia and Psacalium with the open oak-pine woods or their
borders. But species of the different genera do not usually grow to-
gether in the same community.

The annual seasonal pattern in the oak-pine woodland zone is one
of alternating wet and dry periods. The rainy season often begins
in May and continues as late as October in certain areas, the peak
reached during July and August. The dry season lasts from about
mid-November until mid-May. The highest temperatures are usually
reached during the months just prior to the peak of the rainy season,
April through June (Contreras Arias, 1942). The active growing
period corresponds to the rainy season with little plant growth
during the dry season when the plants die or remain dormant.

Most of these taxa occur at altitudes between 1,000 and 3,000
meters, although a few specimens have been collected at elevations
as low as 350 meters. These plants seem to thrive in disturbed sec-
ondary associations such as grazed grasslands, burned-over wood-
lands, or roadsides. Originally they probably grew along the margins
of the oak-pine woods or in natural openings within the woods. The
majority of the species grow in well-drained rocky, clay soils but a
few species do well in poorly drained, similar soils.

The region of greatest concentration of these species corresponds
closely to the Trans-Mexican volcanic belt (Clausen, 1959) or Neo-

PIPPEN—MEXICAN “CACALIOID” GENERA 371

volcanic Plateau (Raisz, 1959). This zone extends from the region
of Orizaba, in Veracruz, through the region of Mexico City to west-
ern Michoacin, thence curving northward toward Guadalajara and
again northwestward toward Tepic. In all directions away from this
zone the number of species diminishes rapidly. Other centers of
concentration for Digitacalia, Odontotrichum, and Psacalium are in
southern Durango and central Oaxaca, in the latter region are several
endemic taxa.

The general distributional patterns within Mexico are illustrated
in figures 2 and 3.

Generic Relationships and Evolution

The genera Digitacalia, Odontotrichum, Pericalia, and Psacalium are
apparently natural genera, possibly having closer relationships to sep-
arate elements within the genus Senecio than to each other. Still, these
segregate cacalioid genera may be distinguished as a group from
Senecio by a number of relatively constant characters if the characters
are applied collectively (table 2). Odontotrichum is the largest genus
in the complex. It may be distinguished from the other cacalioid
genera by the subscapose habit, the erect, nonpeltate basal leaves, and

TABLE 2.—Comparison of cacalioid genera and Senecio

Cacalioid Genera Senecio

(a) Heads always discoid, the flowers (a) Heads radiate or discoid, the ray
all regular and perfect. flowers pistillate; the discoid

flowers regular and perfect.

(b) Corollas always creamy or nearly | (b) Corollas mostly yellow or purple,
white, never yellow. occasionally creamy or nearly

white.

(ec) Corollas deeply lobed, the lobes (ec) Corollas mostly shallowly lobed,
extending to the base of the limb the lobes usually less than half as
or connate into a throat rarely as long as the throat.
long as the free lobes.

(d) Achenes mostly obovate or ellip- (d) Achenes mostly cylindric, subterete
soid, elliptic in cross section, less in cross section; glabrous or pu-
frequently cylindric and subterete bescent, the hairs in many species
in cross section; glabrous or pu- hygroscopic.
bescent, the hairs never hygro-
scopic.

(e) Involucre mostly campanulate or (e) Involucre cylindrical, less fre-
turbinate, less frequently quently campanulate or turbinate.
cylindrical.

(f) Plants all herbaceous perennials. — (f) Plants annuals, biennials, herba-

ceous perennials, shrubs or trees.

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

372

SCALE
Del 700 foot
™ Bs TD

may
(\4 1000 FOOT CONTOUR
BONNE'S LQUAL-ARLA PROJECTION

BONNIE'S LQUAL-ARELA PROUECTION

t

f the Mexican cacalioid genera

3 2. Odontotrichum.
f one or more spec

ion Maps o

buti
1. Digtiacalia

istri

FicurE 2,—D

Collect

1es.

10n oO

PIPPEN—MEXICAN “CACALIOID” GENERA 373

se tel tonne
e ee eco ke TE
#44 1900 FOOT CONTOUR
BONNE'S EQUAL-AREA PROJECTION

aeoneet

. - oa ee
Fx.# 10 FOOT CONTOUR
BONWE'S CQUAL-AREA PROJECTION

Lt ———1

Ficure 3.—Distribution maps of the Mexican cacalioid genera:
3. Pericalia; 4. Psacalium.
@=Collection of one or more species.

374 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

the corolla lobes free to the base of the limb (fig. 1). Taxa of this de-
scription apparently have no counterpart in Senecio unless it is in S.
sect. Mulgedifolii. The species in this section have discoid heads usu-
ally of purple flowers (although some white-flowered forms do occur)
but the corolla throat is much longer than the lobes, the lobes forming
only five small, deltoid teeth above the throat (fig. 1). The leaves are
mostly membranaceous, narrowly ovate or oblong, regularly callose-
dentate, and often runcinate or lyrate-pinnatifid. Several of these
species have been described as cacalias (sensu latu), e.g., S. callosus
Greenm., S. coulteri Greenm., S. runcinatus Less., S. purpurascens
Klatt, and S. deformans Klatt.

Psacalium is the most distinctive genus of the complex because of
the centrally peltate leaves. It is further distinguished by the frequent
occurrence of stipitate, glandular trichomes in the inflorescence and
the fusion of the corolla lobes into a short throat ca. 1 mm. long (fig. 1).
No group of species in Senecio is known to be comparable to the species
of Psacalium, although several species in S. sect. Palmatinervii appear
closely allied. Senecio subpeltatus Schultz Bip. and S. cordobensis Hemsl.
have peltate leaves but the flowers are typically senecioid and the
plants are woody shrubs. Senecio angulifolius DC. is similar in several
respects to P, peltatum; the two have leafy inflorescence bracts, heads
of approximately the same size, glandular-pubescent stems, and a
chromosome number reported asn=30. However, S. angulifolius is a
woody shrub, the leaves are not peltate, the heads are usually radiate
(occasionally discoid), and the flowers are always yellow.

Since this group of senecios appears to be somewhat transitional
between Senecio and Psacalium, it is possible that the latter was de-
rived from an ancestral stock that also gave rise to species like those
in S. sect. Palmatinervii.

In Digitacalia the plants are leafy stemmed and the upper leaves
reduced but not bracteiform. There is neither a basal rosette of
leaves nor subterranean tubercles. The corolla is mostly deeply cleft
(fig. 1), but the achenes are subcylindric, subterete in cross section,
and thus tending toward the senecioid type. Senecio roldana DC.
and S. jaliscana S. Wats. resemble the species of Digitacalia in habit,
although the leaves are less deeply lobed, but in floral characters
they are distinctly senecioid; S. jaliscana has discoid heads of orange
flowers and S. roldana radiate heads of yellow or white flowers. In
both species the corollas are senecioid.

Conceivably Digitacalia may have been derived from the same
ancestral stock that also gave rise to species in Senecio like those just
discussed.

Finally, Pericalia is unique in the cacalioid complex because of its
leafy-stemmed habit, the distal bracteiform leaves, the fleshy sub-

PIPPEN—MEXICAN “CACALIOID” GENERA 375

terranean tubercles, the numerous (8-15) calyculate bracts, and the
well developed, long corolla throat with relatively long, recurved
corolla lobes. This genus contains only two species and these have
no apparent counterpart in Senecio. But S. hartwegu Benth. and S.
seemanit Schultz Bip. (sect. Palmatinervit) have certain similarities
in vegetative structure to the species of Pericalia, although the heads
of these senecios are much smaller and distinctly senecioid.

A Central American species, Senecio coopert Greenm. (sect. Multi-
nervii), bas disk florets similar to the discoid flowers of the cacalioid
genera, with the corolla lobes much longer than the throat. The heads,
however, are radiate, the flowers yellow, the phyllaries narrowly
oblong and barely overlapping, and the leaves lyrate-pinnatifid. This
combination of characters definitely associates the species with
Senecio.

The hypothetical relationship of the cacalioid genera to each other
and to the closely allied species of Senecio is summarized in figure 4.

It is also apparent that the Mexican cacalioid taxa are not the same
as those in the eastern United States (Cacalia sect. Conophora DC.)
or those in eastern Asia (C. sect. Hucacalia DC.). Further studies of
these groups are necessary to determine their proper relationships
and taxonomic position.

The 7 or 8 species usually referred to Cacalia sect. Conophora DC.
form a natural group endemic to the eastern United States and
southern Canada. These are easily distinguished from the taxa in-
cluded in this paper as well as from Senecio by: The presence of a
prolonged chaffy scale forming a beak up to two millimeters long in
the center of the receptacle; the consistently small heads with 5
phyllaries and 5 or 6 flowers; the entire or dentate but not lobed,
glabrous or glaucous leaves; and the haploid chromosome number of
usually 25.

The 8 to 10 species that most earlier workers referred to C. sect.
Eucacalia comprise a natural group mostly endemic to eastern Asia,
with the exception of C. suaveolens L. which is found only in the
eastern United States. These differ from the Mexican members of
this complex in: The lack of tufts of hair at the base of the basal
leaves; the narrowly paniculate inflorescence; the short corolla tube;
and the throat of the corolla equaling or exceeding the corolla lobes
in length.

The genus Cacalia proper, a small group of central European
species typified by C. alpina L. (see discussion to follow) differs from
the taxa discussed in this paper primarily as follows: The style
branches are long-attenuate, the abaxial surface uniformly densely
glandular-papillose; the corolla is usually 4-parted; the chromosome
number is reported as n=19 (Langlet, 1936); and these taxa, under

376 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

[3 seen HARTWEGI|I

S. SEEMANNII
| PERICALIA SESSILIFOLIA
; 7 MICHOACANA
[| —
| | HETEROIDEA
| | DIGITACALIA
| | / CHIAPENSIS _
ly of JATROPHOIDES PACHYPHYLLUM
I | j NAPEIFOLIA AMPLUM
| | / | TRIDACTYLITIS SILPHIIFOLIUM
lj GOLDSMITHI!
l | S. JALISCANA AMPLIFOLIUM
/ y S. ROLDANA GLOBOSUM
it! ; PURPUSI|I
[| s
Zz BRACHYCOMUM
A __-— S$. SECT. MULGEDIFOLII PAL MERI
\em
ANCESTRAL PLATYLEPIS
| ODONTOTRICHUM PRINGLE!
PROTOTYPES k PAUCICAPITATUM
\ NELSONI MULTILOBUM
~~ PELTATUM
—Y \ \ PSACALIUM
.\ LAXIFLORUM RADULIFOLIUM
V\ HOLWAYANUM TUSSILAGINOIDES
\ \ s. SECT. ERIOCARPUM NAPELLIFOLIUM
sp,
\ PALMATNERVII CERVINUM
\ TABULARE FILICIFOLIUM
\ MEGAPHYLLUM CIRSIIFOLIUM
\ POCULIFERUM DECOMPOSITUM
\ PELTIGERUM —
S. ANGULIFOLIUS | yanuM
S. COOPERI CALVUM

Ficure 4.—Hypothetical relationships of the species of the cacalioid genera in Mexico to
each other and to certain closely allied species of Senecio. Vertical gaps and positions
represent relative morphological diversity among species and groups of species.

the name Adenostyles, have been referred to the tribe Eupatorieae

(Hoffmann, 1890).
Nomenclature of Cacalia

One of the major points of difficulty concerning the generic nomen-
clature of the taxa considered in this paper as well as that of Cacalia
has been the typification of the latter, since no type-species was desig-
nated by Linnaeus. The problem of the typification of Cacalia has

PIPPEN—-MEXICAN “CACALIOID” GENERA 377

been reviewed several times (Rydberg, 1924; Hitchcock & Green, 1947;
Shinners, 1950; Cuatrecasas, 1960) and therefore does not need to be
repeated in detail here. It seems clear that C. alpina L. is the most
logical lectotype of Cacalia. This species, named C. alpina by Linnaeus
(1753), embodied the Linnaean and pre-Linnaean concept of Cacalia
in that essentially all of the species of Cacalia described by the pre-
Linnaean botanists (L’Obel, 1581; Clusius, 1601; Bauhin, 1623;
Morison, 1699; Tournefort, 1700) actually represented the same
species.

Linnaeus’s concept of Cacalia was an inclusive one, and a number
of the species he considered in this genus were transferred to other
genera (Haworth, 1812; Cassini, 1816, 1826, 1827). Nomenclatural
difficulty began when Cassini (1816) created a new genus, Adenostyles,
for a small group of central European plants including C. alpina L.
as the type-species. Adenostyles was actually superfluous when pub-
lished since the name Cacalia should have been used for the genus
containing C. alpina.

The name Adenostyles has been generally accepted (DeCandolle,
1838; Bentham and Hooker, 1873; Hoffmann, 1890; Hegi, 1917; Fiori,
1933) for C. alpina and four or five other species endemic to central
Europe.

DeCandolle (1838) summarized the work of Cassini and the other
post-Linnaean authors. He completely revised the cacalioid complex
and in effect retypified Cacalia, although he designated no type-species.
He divided Cacalia into four sections: Hucacalia and Conophora and
two others that he considered doubtfully valid, Aulacophora and Cis-
sampelopsis (the latter two comprised of several shrubby paleotropical
yellow-flowered species now mostly referred to Senecio). In the
emended genus Cacalia, he retained only three Linnaean species, C.
hastata and C. suaveolens in Eucacalia, and C. atriplicifolia in Cono-
phora. DeCandolle recognized Cassini’s genus Adenostyles, thereby ex-
cluding from Cacalia the Linnaean type of the genus; he used the
name Psacalium Cass. for the Mexican species with peltate leaves;
and he referred to Cacalia various species now properly referred to
the genera Digitacalia, Odontotrichum, Pericalia, and Psacalium.

In the sense of DeCandolle, therefore, Cacalia was to be typified
by one of three Linnaean species (but not the logical generic type).
It included a variety of species, occurring in Asia, eastern North
America, and Mexico, that differed consistently from Senecio in a
number of respects and was regarded by most authors (Gray, 1883;
Hoffmann, 1890; Greenman, 1902; Fernald, 1950; Blake, 1960; Makino,
1958; Cronquist and Gleason, 1963) as comprising a single widespread
genus. On the other hand, several authors (Schultz, 1845; Bentham &
Hooker, 1873; Hemsley, 1881) did not accept the Candollean Cacalia,

378 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

preferring to relegate all of the species to Senecio. Some authors (e.¢.,
Rydberg, 1924; Cuatrecasas, 1960) took the position accepted in this
paper, that of recognizing several valid genera in the Candollean
Cacalia.

Systematic Treatment

Key to Mexican ‘‘Cacalioid” Genera Allied to Senecio

1, Basal rosette present, the cauline leaves reduced upward, the uppermost
usually bracteiform.

2. Basal leaves centrally peltate . . . . . . 4. Psacalium
2. Basal leaves not centrally peltate, the petiole attached. to the blade at the
edge (subpeltate in O. radulifoltum) . .. . . . . . 2. Odontotrichum

1. Basal rosette lacking, the plants with only cauline leaves, the uppermost leaves
reduced but not bracteiform.

3. Plants with underground tubercles attached to the base of the stem; heads

subtended by 8-15 linear bracts as long as or longer than the phyllaries.

3. Pericalia

3. Plants lacking underground tubercles; heads subtended by 3-5 subulate

bracts shorter than the phyllaries . . ....... 2... J. Digitacalia

1. Digitacalia
Digitacalia Pippen, gen. nov.

Plantae caulescentes, tuberculis subterraneis nullis, caulibus
foliisque glabris vel pubescentibus, foliis omnibus caulinibus, alternis,
numerosis, laminis profunde lobatis; inflorescentia corymbosa ampla
30 cm. longa vel longior; capitula numerosa, breves, corollis profunde
5-lobatis, lobis saepe basin versus connatis; achaenia oblonga, sub-
teretia; pappus sordidus.

Tyerus: Cacalia jatrophoides H.B.K.

Perennial, leafy-stemmed herbs to 3 m. tall; roots fibrous, fleshy;
stems annual, erect, stout, terete or less frequently angulate, glabrous
or pubescent, not tuberculate at base; leaves cauline, not forming a
basal rosette, alternate, numerous, uniformly distributed along the
stem, not clustered toward the middle, the uppermost not bracteiform;
blades shaped variously, deeply pinnately subpalmately lobed; in-
florescence corymbose, large, at least 30 cm. long and broad, the
branches subtended by small, linear-subulate or large, leafy bracts;
heads many, small (except D. heteroidea), calyculate, the bracts 3-7,
usually shorter than the phyllaries; involucre campanulate or turbi-
nate, the phyllaries 5-8, biseriate appearing uniseriate, imbricate, the
overlapped margins subscarious, the exposed surface glabrous or
pubescent, the tips fringed with minute, nonglandular trichomes up
to 0.3 mm. long; receptacle flat or nearly so, alveolate, not paleaceous;
flowers all discoid, perfect, fertile; corollas glabrous, creamy or nearly
white, occasionally greenish or purplish-tinted but never yellow, the

PIPPEN—MEXICAN “CACALIOID” GENERA 379

tube narrowly cylindrical, ca. 0.5 mm. in diameter, the limb divided
into 5 equal, 3-nerved, narrowly triangular or linear, ascending or
spreading lobes, the lobes free to the tube or connate into an ampliate
throat up to 1.5 mm. long and broad; anthers 1.5-3 mm. long, exserted
from the corolla throat, the bases rounded or slightly sagittate, the
terminal appendages trullate or ovate, 0.5-0.7 mm. long; style
branches 0.7-2 mm. long, slightly flattened, recurved or spreading,
stigmatic over most of the abaxial surfaces, the adaxial surfaces
glabrous or hispidulous below, the tips truncate, rounded or obtuse,
fringed with unicellular, nonglandular “pollen producing” trichomes,
unappendaged; achenes narrowly oblong, subterete in cross section,
ribbed; pappus a single row of creamy white, stiff, barbellate, capil-
lary bristles.

This genus differs from Psacalium and Odontotrichum in that the
leaves are strictly cauline with no basal rosette formed and the
achenes are essentially cylindrical. It differs from Pericalia in that the
leaves are not concentrated toward the middle of the stem but more
uniformly distributed, there are no underground tubercules and the
corolla lobes are not recurved.

Key to Species of Digitacalia

1. Heads large, 1.5-2 em. high at maturity; flowers 40 or more; corolla tube at
least twice as long as the lobes. . .. . ..... 2D. heteroidea
1. Heads small, 1 em. high or less; flowers 13 or “fewer; corolla tube and lobes
about equal.
2. Achenes pubescent; corollas short, 3-3.5 mm. long . . 1. D. chiapensis
2. Achenes glabrous; corollas 5-7 mm. long.
3. Leaves 3-lobed, the lobes forming an angle, with the midrib, of less than
45 degrees... 1 ee ee ee ee ee . . 5. D. tridactylitis
3. Leaves 5- to 7-lobed, the lobes or at least some of the lobes forming an
angle with the midrib greater than 45 degrees.
4, Phyllaries and flowers 5 or 6 per head. .... . 4. D. napeifolia
4, Phyllaries 7; flowers 8-10 per head... . . . 3. D. jatrophoides

1. Digitacalia chiapensis (Hemsl.) Pippen, comb. nov.
Senecio chiapensis Hemsl. Biol. Centr. Amer. Rot. 2:238. 1881. Holotype:
Chiapas & c., Ghiesbreght 537 (K, not seen; isotype GH!).
Cacalia chiapensis (Hemsl.) A. Gray, Proc. Amer. Acad. 19:53. 1883.
Odontotrichum chiapensis (Hemsl.) Rydb. Bull. Torrey Bot. Club 51:418. 1924.
Lower portion of plant not seen; stems terete, glabrous; leaves
presumably all cauline, petiolate, oblong-ovate in outline, 10-14 cm.
long, 9-14 cm. wide, pinnately veined; blades deeply pinnately
5-lobed ca. two-thirds the distance to the midrib, the lobes ovate
or oblong-acute, 4-6 em. long, 2-3 cm. wide, about half as wide as
long, the lowermost pair of lobes rounded proximally or with two
smaller deflected lobules, the margins entire, the abaxial surface
266-83

380 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

puberulent mostly along the veins, the adaxial surface glabrous;
petioles short, to 4 cm. long, glabrous, terete; inflorescence broadly
corymbose-paniculate, to 30 em. long and broad, the branches gla-
brous, subtended by sessile, ovate bracts; calyculate bracts ca. 3,
subulate, 1.5-2 mm. long; heads many, 7-8 mm. long; phyllaries
(6 -) 8, glabrous, narrowly ovate, 3.5-5 mm. long, 1-1.5 mm. wide;
flowers 9; corolla 3-3.5 mm. long, the tube 1.5-2 mm. long, the lobes
linear or narrowly elliptic, 1.5 mm. long, spreading or reflexed, the
throat none; anthers 1.5 mm. long; style branches 0.7-1 mm. long,
the tips truncate or rounded with a terminal fringe of trichomes;
mature achenes not seen, the immature achenes brown, sparsely
pubescent; pappus bristles 2-3 mm. long, white.

DistTriputTion: Known only from Chiapas. Apparently flowering
in the fall but the exact time not known.

This plant has been very poorly collected. It is known from only
two collections in Chiapas, one is the type.

CutApas: Nelson 3467 (GH, US).

2. Digitacalia heteroidea (Klatt) Pippen, comb. nov.

Senecio heteroideus Klatt, Leopoldina 24:125. 1888. Holotype: Mexico; be-
tween San Andrés and San Miguel, Liebmann 178 (C, not seen; tracing of
type in Klatt’s herb, GII!).

Cacalia laevigata Schultz Bip. Leopoldina 24:125. 1888, in synonymy.

Cacalia longipetiolata Rob. & Greenm. Amer. Journ, Sci. III. 50:157. 1895.
Holotype: Oaxaca; Sierra de San Felipe, 7,000 ft., Oct. 10, 1894, C.C.
Pringle 5828 (GH!).

Plants to ca. 2 m. (Pringle 6176) tall; lower portion of the plant
not seen; stems glabrous, glaucous; leaves pubescent on both surfaces
mostly along the veins, presumably all cauline, long-petioled, sub-
palmately veined with 5-7 principal veins, transversely broadly
elliptic in outline, the larger leaves 6-13 cm. long from the basal sinus
to the apex, 9.5-22 cm. wide; blades cordate-based, deeply sub-
palmately 5- to 7-lobed, the lobes oblong or triangular, the larger
ones 5-6 cm. long, 1-3 cm. broad, the margins toothed or lobulate;
petioles glabrous, terete, 8-12 cm. long; inflorescence paniculate
corymbose, the major branches subtended by leafy, narrowly elliptic,
petiolate bracts, 3-12 cm. long, 0.5-3 cm. broad, glabrous; pedicels
5-13 cm. long; calyculate bracts 5-7, subulate, 5-7 mm. long; heads
12-40, 16-18 (-20) mm. long; phyllaries 7 or 8, ovate, 11-12 (-14)
mm. long, 3-4 (-6) mm. wide, glabrous; receptacle ca. 7 mm. across;
flowers ca. 40; corolla 9-10.5 mm. long, the tube 5.5-6 mm. long, the
throat short, campanulate, 1-1.5 mm. long, ca. 1.5 mm. across at the
apex, the lobes 2-3 mm. long, narrowly triangular, 0.5 mm. wide,
loosely reflexed; anthers 3 mm. long; style branches ca. 1 mm. long,

PIPPEN—MEXICAN “CACALIOID” GENERA 381

minutely hispid on the abaxial surface, the tips obtuse with a sparse,
subterminal fringe of trichomes; mature achenes ellipsoid, 3.5-5.5
mm. long, 1.5 mm. wide, 1 mm. thick, glabrous, brown, with 10-12
prominent, paler ribs; pappus 7-8 mm. long, white.

DistrisuTion: Known only from central Oaxaca. Presumably
flowering in September or October.

This species is placed in Digitacalia because its general habit is like
that of the other species in this genus, although the inflorescence
with few, large heads of 30 or more flowers is more like the species in
Pericalia. It differs from Pericalia in the short corolla throat and the
fewer calyculate bracts. D. heteroidea appears to be somewhat inter-
mediate between this genus and Pericalia.

Oaxaca: Conzattt 725 (GH); Pringle 6176 (GH, NY, US); C. L. Smith 287
(MICH, NY, US).

3. Digitacalia jatrophoides (H.B.K.) Pippen, comb. nov.

Cacalia jatrophoides H.B.K. Nov. Gen. et Sp. 4:169. 1820. (Folio ed. 4:132.
1820.) Holotype: Michoacdn; nr Lake Cuitzeo, 900 hex., Humboldt &
Bonpland 4298 (P, not seen; Field Mus. Neg. 37869!).

Senecio jatrophoides (H.B.K.) Schultz Bip. Flora 28:498. 1845.

Cacalia digitata Sessé & Moc. Pl. N. Hisp. ed. 1. 132. 1889. Lectotype
without locality (presumably “in Purulandiri montibus’’) Sessé & Moc.
2826 (MAI; Field Mus. Neg. 42288).

Odontotrichum jatrophoides (H.B.K.) Rydb. Bull. Torrey Bot. Club 51:419.
1924.

Plants to 3 m. tall; stems to 1 cm. thick at the base, mostly hollow,
often purplish, glabrous, glaucous; leaves sparsely pubescent on both
surfaces mostly along the veins and margins, numerous, long-petiolate,
subcircular in outline, 6-10 cm. long and broad, pinnate or subpalmately
veined with 5-7 principal veins fused into a common trunk above the
base of the blade; blades deeply 5- to 7-lobed, the lobes subpinnate or
subpalmate, the central one larger, linear or narrowly ovate, 6-8 cm.
long, 1-1.5 cm. wide, the margin subentire or distantly toothed (lobes
occasionally larger and the leaves similar to those of D. napeifolia) ;
petioles terete, glabrous, purplish, to 9 cm. long; inflorescence broadly
corymbose, loose spreading, usually more than 30 cm. long and broad,
the branches sparsely pubescent or tomentose, the proximal branches
in the axils of the 3-lobed upper leaves, the distal branches subtended
by petiolate, narrowly ovate bracts to 2 cm. long; calyculate bracts
3-7, subulate, 2-5 mm. long, heads many, 1-1.5 cm. long; phyllaries
7 or 8, narrowly ovate, 4-5.5 mm. long, 1-1.5 mm. wide, glabrous;
flowers 8-10; corolla 6-S mm. long, the tube greenish, 3-5 mm. long,
the lobes 3 mm. long, 0.5 mm. wide, linear or narrowly elliptic, the
throat none; anthers ca. 2 mm. long; style branches 1 mm. long, re-
curved, the tips rounded or obtuse with a subterminal fringe of tri-
chomes; mature achenes narrowly oblong appearing subterete in cross

382 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

section, 4-5 min. long, 1.5 min. wide, glabrous, pale tan, strongly 10- to
12-ribbed; pappus 5-7 mm, long, creamy, becoming tawny in age.
DisTrIBuTION: Southern Zacatecas and Jalisco, eastward through
Michoacén and Guanajuato and south to Oaxaca. Rocky places in
highland pastures and grasslands. Flowering in July and August.
The specimens collected in Zacatecas and one from Oaxaca have
much broader leaf lobes than the others. These leaf forms are similar
to those of D. naperfolia DC., but these specimens are considered with
D. jatrophoides because the floral characters are more like the latter.

Guanasuato: Pringle 4262 (GH, NY, US). Jauisco: McVaugh 17160 (MICH).
MicuoacAn: Arsene, s.n., Sept. 9, 1909 (US); Pippen 46(MICH). Oaxaca: L. C.
Smith 123 (GH); C. L. Smith 388 (MICH, NY, US); Pringle 5741(GH). Zacatr-
cas: Rose 3040 (GH, US).

4. Digitacalia napeifolia (DC.) Pippen, comb. nov.

Cacalia napeaefolia DC. Prod. 6:328. 1838. Holotype: Oaxaca; summit of

San Felipe, in 183-, Andrteux 280 (G, not seen; Field Mus. Neg. 33809!).
Senecio napeaefolium (DC.) Schultz Bip. Flora 28:498. 1845.
Odontotrichum napeaefolium (DC.) Rydb. Bull. Torrey Bot. Club 51:418. 1924.
Plants to 2.5 m. tall (Pringle 4778); lower portion of the plant not
seen; stems terete, purplish, sparsely pubescent; leaves presumably all
cauline, numerous, petiolate, pinnately veined, oblong-ovate, 15-29
cm. long, 16-23 cm. wide, progressively smaller upward; blades deeply
5- to 7-lobed to within 0.5-1 em. of the midrib, the lobes oblong-
attenuate, narrowly ovate, to 14 cm. long, 4 em. wide, the margins
finely or coarsely toothed, the proximal pair of lobes usually deflected
or at right angles to the midrib, the adaxial surface of the blade
glabrous or sparsely pubescent mostly along the veins, the abaxial
surface pale, sparsely pubescent or tomentose; petioles subterete,
elabrous below becoming pubescent toward the blade; inflorescence
corymbose, to 45 cm. long and broad, the branches tomentose, the
proximal ones subtended by the uppermost leaves, the distal branches
subtended by linear-subulate bracts, 1.5 cm. long; calyculate bracts
subulate, 1.5-3 mm. long; heads many, 1 em. long; phyllaries 5 (rarely
6), oblong or narrowly ovate, 3.5-5.5 mm. long, 1.5-2 mm. wide,
glabrous or very sparsely puberulent; flowers 5 or 6; corolla 6-7 mm.
long, the tube 3-8.5 mm. long, the lobes 3-4 mm. long, linear or
narrowly elliptic, 0.5 mm. wide, the throat none; anthers 2-3 mm.
long; style branches 1-2 mm. long, minutely hispid on the abaxial
surface, the tips obtuse with a subterminal fringe of trichomes; mature
achenes narrowly oblong, appearing subterete in cross section, 4-5
mm. long, 1-1.5 mm. wide, glabrous, purplish brown, strongly 10-
ribbed; pappus 6-7 mm. long, creamy, becoming tawny in age.
DisrrirputTion: Known mostly from central Oaxaca with one col-

PIPPEN—MEXICAN “CACALIOID” GENERA 383

lection from Michoacin. Openings in oak-pine woodlands. Flower-
ing in September and October.

Micuoachn: Hinton et al. 12572 (MICH, NY). Oaxaca: McVaugh 21823
(MICH); Nelson 1133 (GH, US); Pringle 4778 (GH, MICH, NY, US).

5. Digitacalia tridactylitis (Rob. & Greenm.) Pippen, comb. nov. PuaTE 1
Cacalia tridactylitis Rob. & Greenm. Amer. Journ. Sci. III. 50:159. 1895.
Holotype: Oaxaca; Sierra de San Felipe, 6,000 ft., Nov. 19, 1894, C. G.
Pringle 5841 (GH!).
Odontotrichum tridactylitis (Rob. & Greenm.) Rydb. Bull. Torrey Bot. Club
51:419, 1924.

Plants to at least 2.5 m. tall; stems to slightly over 1 cm. thick at
the base, angulate or terete, sparsely pubescent below becoming denser
above; leaves cauline, numerous, pinnately veined, obtrullate or
rhombic in outline, the larger ones 14-20 cm. long, from two-thirds
as broad to as long as broad, sparsely pubescent adaxially, gray
tomentose on the abaxial surface; blades deeply 3-lobed at least half
way to the midrib, the lobes ovate, narrowly ovate or narrowly tri-
angular, variable in size from 2- to 6-times longer than broad, the
angle of the lateral lobes to the midrib less than 45 degrees, the margin
of the lobes usually entire, occasionally toothed, the base of the blade
decurrent along the petiole for a short distance; petioles 2-3.5 cm.
long, triangular in cross section, pubescent; inflorescence corymbose,
ca. 30 cm. long and broad, the branches tomentose, subtended by the
uppermost cauline leaves; pedicels 3-7 mm. long; calyculate bracts
3-7, subulate, 1.5-3.5 mm, long; heads many, 9 mm. long; phyllaries
7 or 8, narrowly ovate, 3.5-4 (-5) mm. long, 1-1.5 mm. wide, glabrous;
flowers (7—) 8-10; corolla 5-6 mm. long, the tube 2.5-3 mm. long, the
throat usually obsolete or very short, 0.5 mm. long and wide, occa-
sionally to 1.5 mm. long, ca. 1 mm. wide (McVaugh 21903), the lobes
narrowly elliptic, 2-3 mm. long, 0.5 mm. wide; anthers 2-3 mm. long;
style branches 1-1.5 mm. long, the tips rounded with subterminal
fringe of trichomes; achenes narrowly oblong, appearing subterete or
slightly compressed in cross section, 3.5-4 mm. Jong, ca. 1 mm. wide,
glabrous, pale tan, 10- to 12-ribbed; pappus (3.5—-) 4-5.5 mm. long,
creamy, slightly clavate distally; chromosome number, n=30 (Turner,
ined.).

DistriBution: Morelos, Oaxaca, and Guerrero. Growing in open-
ings in the pine-oak zone. Flowering in October and November.

The specimens from Morelos tend to have deeper, more narrowly
lobed leaves and less pubescence on the abaxial surface than the other
specimens.

GuERRERO: McVaugh 21903* (MICH). Moretos: Pringle 6164 (GH, MICH,
NY, US); Pringle 9877 (GH, MICH, NY, US). Oaxaca: Nelson 2080 (US);
C. L. Smith 380 (MICH, NY); L. C. Smith 966 (GH).

384. CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

2. Odontotrichum

Odontotrichum Zuec. Abhandl. Baier. Akad. Wiss. 1:311. 1832.

Perennial, subscapose herbs, 0.5-2 m. tall; roots fleshy, fibrous,
caudex tough, to 5 cm. long, 3 cm. thick, the nodes and terminal
buds densely covered with tufts of tawny, long, multicellular hairs;
stems usually solitary, annual, erect, slender, terete or less frequently
angulate, glabrous or pubescent; radical leaves forming a basal
rosette of two to several, the petiole attached to the base of the
blade, not centrally peltate; blades pinnately or subpalmately veined,
variously lobed or occasionally subentire, glabrous or pubescent;
cauline leaves similar to the basal but progressively smaller upward,
the uppermost bracteiform; inflorescence paniculate or corymbose
often pubescent but not glandular, the branches subtended by small,
linear-subulate or large, leafy, often vaginate bracts; heads few to
many, calyculate, the bracts usually 1-5, usually as long as or shorter
than the phyllaries; involucre cylindrical, campanulate or turbinate,
the phyllaries 5-17, imbricate, biseriate appearing uniseriate, the
overlapped margins subscarious, the exposed surfaces glabrous or
pubescent, the tips fringed with minute trichomes up to 0.3 mm.
long; receptacle flat or nearly so, alveolate, occasionally sparsely
pubescent, not paleaceous; flowers all discoid, perfect, fertile, 5-80
per head; corollas glabrous, creamy or nearly white, occasionally
ereenish or purplish tinted but never yellow, the tube narrowly
cylindrical, ca. 0.5 mm. in diameter, the limb divided into 5 equal,
3-nerved, narrowly elliptic, narrowly triangular or linear, ascending
or spreading, not recurved lobes, the lobes usually free to the tube,
occasionally connate into a short, ampliate throat up to 1 mm. long;
anthers 2-4 mm. long, exserted above the corolla, the bases rounded
or slightly sagittate, the terminal appendages trullate or ovate, 0.5-
0.7 mm. long; style branches 0.7-2 mm. long, slightly flattened,
recurved or spreading, stigmatic on most of the abaxial surface, the
adaxial surface glabrous or hispidulous below, the tips unappendaged,
rounded or obtuse, fringed with minute, unicellular, nonglandular,
“pollen producing” trichomes; achenes oblong, ellipsoid or obovoid,
somewhat dorsally compressed, elliptic or broadly elliptic in cross
section, glabrous or pubescent, striate or prominently 10- to 12-
ribbed; pappus a single row of white or creamy, stiff, barbellate,
capillary bristles; chromosome numbers 25, 30.

TypeE-spEciES: Odontotrichum cirsiifolium Zuce.

This is the largest genus treated in this paper, with 21 species.
It is distinguished from Psacalium by the lack of centrally peltate
leaves, the blades being attached at their bases to the petiole. In
most of the species the leaves are vertically oriented, standing erect.

PIPPEN—MEXICAN “CACALIOID” GENERA 385

Key to Species of Odontotrichum

1. Pappus short, 2 mm. long or less, about half as long as the corolla tube.

2. Petioles flattened, winged, the margin up to 1 em. wide; basal leaves lobed
about one-half the distance to the midrib, lobes usually longer than
broad. ..... see ee we ww ew ew we 6 OW brachycomum

2. Petioles subterete, not winged; basal leaves coarsely toothed or lobulate,
but not lobed, lobules broader thanlong. . ..... . 13. O. palmeri

1. Pappus at least 3 mm. long, as long as or longer than the corolla tube.

3. Basal leaves toothed or subentire, not lobed (some species with two auriculate
basal lobes formed by the deeply cordate basal sinus).

4. Inflorescence very compact, globose, 2.5-3 em. long and broad; phyllaries

5 or 6 per head . ve ee ee ee ee . 8. O. globosum

4. Inflorescence tightly or loosely corymbose, not globose, at least 4 cm.

long and broad; phyllaries at least 7.

5. Heads large with 30 or more flowers; phyllaries 13 or 14.

6. Bracts subtending the involucre oblong or obovate, distally laciniate,
3 mm. wide; corolla tube 9-10 mm. long, twice as long as the lobes
(5-6 mm. long); phyllaries 12-15 mm. long. . . . 1. O. amplum

6. Bracts subtending the involucre linear-subulate, tapering to a point
distally; corolla tube 5-6 mm. long, slightly longer than the lobes
(3.5-4 mm. long); phyllaries 7-9 mm. long. 12. O. pachyphyllum

5. Heads small with 15 or fewer flowers; phyllaries 8 or fewer.

7. Basal leaves deeply cordate-based; petioles of the cauline leaves
broadly leafy-margined, the base auriculate, clasping the stem;
corolla tube 5-6 mm. long. ....... . 19. O. silphiifolium

7. Basal leaves truncate- or obtuse-based, rarely slightly cordate;
petioles of the cauline leaves usually terete, the base dilated but
not auriculate or clasping the stem (occasionally the petiole nar-
rowly leafy-margined or the leaves sessile); corolla tube 3-4 mm.

long.
8. Plants 0.6 m. tall or less; basal leaves (including the petiole)
11-17 cm. long. ...... » «+ « « « LZ. O. purpusii

8. Plants at least 1 m. tall; basal leaves. (including the petiole) (15-)
20-39 cm. long.

9. Phyllaries 5 or 6; flowers 6 or 7; leaves sparsely puberulent.
9b. O. goldsmithii var. rowellii

9. Phyllaries 7 or 8; flowers 9-13; leaves glabrous.
9a. O. goldsmithii var. goldsmithii
3. Basal leaves pinnately or subpalmately divided into several lobes, the lobes

as long as broad or longer.

10. Leaves deeply dissected or pinnatisect into linear segments, the ultimate
segments linear or narrowly triangular, at least 5 times longer than
broad.

11. Median and distal cauline leaves broadly auriculate-based, the
margin laciniate; proximal inflorescence bracts ovate, navicular,
1.5-3 cm. long, 2-3 cm. broad, the margin distally laciniate; pappus
deciduous before maturity of the achenes . . . .5. O. cirsiifolium

11. Median and distal cauline leaves often dilated at the base but not
broadly auriculate-based or laciniate-margined; proximal inflores-
cence bracts linear subulate, up to 1.5 cm. long, 1 mm. wide or less;
pappus persistent.

12. Achenes glabrous.

3386 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

13. Leaves 3- to Epinnatisect heads with 6-8 flowers and 5 or 6
phyllaries. . . . ... . .6. O. decompositum
13. Leaves deeply dissected or pinnatifid with the primary divisions
again 3- to 5-lobed but not cleft to the midrib; heads with 10
flowers and 8 phyllaries. . . . . . . . . 10. O. multilobum
12. Achenes pubescent.

14. Heads with 12-25 flowers; phyllaries sparsely pubescent.

11. O. napellifolium
14. Heads with 10 or fewer flowers; phyllaries glabrous.

15. Pappus 7 mm. long, about as long as the corolla (lobes plus

tube); phyllaries 5or6.....2.. =. . 7. O. filicifolium
15. Pappus 3.5-4 mm. long, about as long as the tube of the corolla;
phyllaries 7... . . 2... 4 O. cervinum

10. Leaves shallowly or deeply lobed but not finely dissected or pinnatisect
into linear segments, the ultimate segments triangular or deltoid,
usually less than 3 times longer than broad.

16. Heads large, 1.5-2.5 cm. high with 40 or more flowers; corolla tube

7-10 mm. long, twice as long as the lobes.

17. Bracts subtending the involucre leafy, oblong or obovate, distally
laciniate, 3-5 mm. wide; leaves and stem sparsely puberulent;
basal leaves 18-75 em. long and broad . . . . 15. O. platylepis

17. Bracts subtending the involucre linear-subulate, simple, entire-
margined, 1 mm. wide or less, not leafy; leaves and stem denscly
covered by an interwoven mass of gray white, long trichomes;
basal leaves 14-16 cm. long, ca. 10 em. wide.

14. O. paucicapitatum
16. Heads small, 1-1.2 em. high or less; flowers 14 or fewer; corolla tube
about equaling the lobes (ca. 5 mm, long).

18. Achenes pubescent.

19. Basal leaves subpeltate, the petiole attached to the blade 2-10
(-30) mm. above the base, the blades 10-12 (-18) cm. long.

18. O. radulifolium

19. Basal leaves attached at base to petiole, not peltate, the blades
20-30 cm. long.

20. Heads with 9-14 flowers and 7 or 8 phyllaries; cauline leaves
reduced, bracteiform, narrowly ovate, laciniate-lobed, 8 em.
long or less, 1-3 em. broad, the base not auriculate.

16. O. pringlei

20. Heads with 5 or 6 flowers and 5 phyllaries; cauline leaves broadly
ovate, leafy, auriculate-based, 11-20 cm. long, 11-20 cm.
broad, the margin dentate or lobed . . 21. O. tussilaginoides

18. Achenes glabrous.

21. Leaves shallowly lobed, the lobes as long as broad or broader
(3-7 cm. broad), rounded at the tip, the margin regularly
toothed; petiole of the cauline leaves leafy-margined, broadly
ovate, auriculate-based; lower inflorescence bracts leafy, 4-9 cm.
long, 1-3 cm. wide, ovate-naviculate. . . . 2. O. amplifolium

21. Leaves deeply lobed, the lobes longer than broad (1-3 em. broad),
oblong or obovate, each of the lobes irregularly 3- to 5-lobed;
petiole of the cauline leaves dilated or often clasping the stem
but not broadly leafy-margined or auriculate-based; lower
inflorescence bracts narrowly ovate or linear-subulate, 1-1.5 cm.
long, 3mm.wide. ........... . 20.0. sinuatum

PIPPEN—MEXICAN “CACALIOID” GENERA 387

1. Odontotrichum amplum Rydb. Bull. Torrey Bot. Club 51:417. 1924. Holotype:
Jalisco; road between Huejuquilla and Mesquitic [Mezquitic], Aug. 25,
1897, J. N. Rose 3575 (US)).

Plants 0.7-1 m, tall, glabrous; stem usually one, 5-8 mm. thick at
the base, terete, glaucous; basal leaves 2 or 3, coriaceous, long-
petioled, deltoid or transversely triangular, the apex rounded, 9-10
em. long, (11-) 18-19 (—22) cm. wide; blades sagittate-based, coarsely
dentate or lobulate, the teeth or lobules many, deltoid or 3-angled,
5-8 (-12) mm. long, 8-10 (-15) mm. wide, ciliate, the abaxial and
adaxial surfaces very similar; petioles to 28 (-40) cm. long, terete;
cauline leaves 5-6, the lowermost one-half to two-thirds as large as
the basal ones and similar to them, the petiole proximally dilated,
clasping the stem; upper cauline leaves gradually smaller upward,
the petioles broadly leafy-margined, the uppermost forming leafy
inflorescence bracts; inflorescence corymbose, 9-23 cm. long, 15-20
cm. broad, glaucous; pedicels 2-6 cm. long; calyculate bracts 3-5,
leafy, 1-1.5 cm. long, 3-6 mm. wide, distally 5- to 9-toothed; heads
9-15, 18-25 mm. long; phyllaries 13 (-15), ovate or narrowly ovate,
12-15 mm. long, 4-5.5 mm. wide, glabrous; receptacle 10-12 mm.
across; flowers 40-50 or more; corolla 14-16 mm. long, the tube
9-10 mm. long, ca. 0.5 mm. wide, the lobes oblong, acute-tipped,
5-6 mm. long, the throat none; anthers 3.5 mm. long; style branches
1.5 mm. long, the tips truncate with a terminal fringe of trichomes;
mature achenes ellipsoid, somewhat dorsally compressed, 6-7 mm.
long, 2 mm. wide, ca. 1.5 mm. thick, glabrous, tan, 20- to 22-ribbed;
pappus creamy, 8-11 mm. long, chromosome number, n= 30 (Pippen).

DistrispuTion: Western Aguascalientes and southwestern Zacate-
cas. Grazed grassland areas. Flowering in September.

Known only from four collections. This species is very similar to
O. pachyphyllum but differs in having larger heads with more flowers
and leafy calyculate bracts.

AGUASCALIENTES: McVaugh & Koelz 57 (MICH); Pippen 2, 3, 4 (MICH).
Zacatecas: McVaugh 17726 (MICH).

2. Odontotrichum amplifolium (DC.) Rydb. Bull. Torrey Bot. Club 51:416. 1924.
Cacalia amplifolia DC. Prod. 328. 1838. Holotype: Oaxaca; Sierra de San
Felipe, on the summit, July 1832, G. Andrieux 293; (G, not seen; Field

Mus. Neg. 33808)).

Senecio amplifolius (DC.) Schultz Bip. Flora 28:498. 1845.

Senecio rumicifolius Klatt, Leopoldina 24:126. 1888. Holotype: Mexico;
Cumbre de Estepa, September 1842. Liebmann 237 (C, not seen; tracing of
type in Klatt’s Herb., GH!).

Cacalia rumicifolia Schultz Bip. Leopoldina 24:126. 1888, in synonymy.

Odontotrichum rumicifolium (Klatt) Rydb. Bull. Torrey Bot. Club 51:416.
1924.

388 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Plants to 1.5 m. tall; stem 0.5-1 cm. thick at the base, terete
arachnoid pubescent, the hairs multicellular, 0.5 mm. long, white,
interwoven; leaves sparsely pubescent adaxially, densely pubescent
abaxially; basal leaves 2-5, occasionally more, long-petioled, pin-
nately veined, ovate-cordate or oblong-cordate, (13-) 17-27 (-30)
cm. long or longer, (12—) 13-19 (—24) cm. wide or wider, usually ca.
two-thirds longer than wide; blades shallowly lobed, the lobes 12-16,
rounded, usually broader than long, (1.5-) 3-4 (-5) cm. long, (2.5-)
3-5 (-7) cm. wide, each lobe often shallowly 2- to 3-lobed or coarsely
toothed; petioles usually longer than the blade, 20-35 cm. long;
cauline leaves 3 or more, the lowermost similar to the basal ones
but the petiole leafy-margined and more dilated and clasping at the
base; upper cauline leaves progressively smaller upward, the upper-
most leaves bracteiform, forming leafy, ovate-naviculate inflorescence
bracts, (3-) 4-9 cm. long, 1-3 cm. wide, distally toothed; inflorescence
corymbose, (6-) 16-30 cm. long, 10-20 cm. broad, the branches
tomentose; pedicels 0.5-1.5 cm. long; calyculate bracts 1-3, subulate,
3-6 mm. long; heads 50 to over 100, 6-10 mm. long; phyllaries 7 or 8
ovate or narrowly ovate, 4-5.5 mm. long, 1-2 mm. wide (specimens
from Oaxaca with phyllaries 6-7 mm. long, 2-3 mm. wide), glabrous;
flowers (8-) 10-13, corolla (5-) 6.5-7.5 mm. long, the tube (2.5-)
3.5-4 (-5.5) mm. long, the lobes linear, (2-) 3-3.5 mm. long, the
throat none; anthers 2-3 mm. long; style branches 1-1.5 mm. long,
hispid on the abaxial surface, the tips truncate or obtuse with a
terminal or subterminal fringe of trichomes; mature achenes ellipsoid
or obovoid (often the marginal ones faleate), 5-5.5 mm. long, 2-3
mm. wide, 1.5-2 mm. thick, glabrous, light brown, striate; pappus
creamy, becoming fuscous with age, 4-6 mm. long.

DistRIBUTION: Oaxaca, Guerrero, and México. Grassy openings 1n
the oak-pine-fir woodlands. Flowering from August to October.

This species is closely related to O. stlphiifolium and O. goldsmitha.
The specimens from Guerrero and México differ from those in Oaxaca
in that the size of the heads (phyllaries and flowers) tend to be smaller,
the inflorescence more open and spreading, and the plants less
pubescent.

Distriro Feperau: Matuda 26624 (NY). Guerrero: Sharp 441465 (NY);
Nelson 2243 (GH, US, this specimen has unusually large leaves); Rzedowski 15974,
16015 (MICH). M&xico: Matuda 27730, 29313, 29609 (NY); Hinton, et al. 8349
(MICH). Oaxaca: Purpus 3139 (GH, NY, US); Conzattt & Gonzalez 397 (GH);
L. C. Smith 124 (GH, this specimen has large cauline leaves and small inflorescence
bracts) ; Nelson 1061 (US); C. L. Smith 391 (MICH, US); McVaugh 21824 (MICH),

3. Odontotrichum brachycomum (Blake) Rydb. Bull. Torrey Bot. Club 51:415.
1924,
Cacalia brachycoma §. F. Blake, Contr. Gray Herb. II. 52:58. 1917. Lectotype:
Michoacdn; wet ravines near Uruapan, 1,525 m., Nov. 13, 1905, C. G.
Pringle 10126 (GH!).

PIPPEN—MEXICAN “CACALIOID” GENERA 389

Plants 1-2 m. tall; stems terete, 7-9 mm. thick at the base, densely
villous below, becoming subglabrous upward, the hairs 0.5-1 mm.
long; leaves minutely puberulent mostly along the veins and margins
with short, white hairs ca. 0.25 mm. long; basal leaves 3 or 4, cori-
aceous, long-petioled, mostly deltoid or ovate, 13-19 cm. long or
longer and broad, pinnately veined; blades sinuately lobed about
one-half the distance to the midrib, the lobes 3-6 cm. long, 2-3.5 cm.
wide at the base, each distally divided into 3 lobes, the secondary
obes deltoid, 3-angulate, the base of the blade truncate, decurrent
along the entire length of the petiole; petiole as long as or longer than
the blade, 13-19 cm. long, leafy-margined, the margin up to 5 mm.
wide on each side; cauline leaves 3 or 4, the lowermost usually smaller
than the basal ones but similar (occasionally larger, more deeply
dissected, up to 30 cm. long, 20 cm. wide), the petiole broadly leafy-
margined, ovate, dentate, clasping the stem; upper cauline leaves
progressively smaller upward, the uppermost bracteiform forming
ovate, naviculate inflorescence bracts, 5-8 cm. long, 3-5 cm. wide;
inflorescence corymbose, 13-28 cm. long and broad; pedicels to 1
em. long; calyculate bracts 1-3, subulate, 1-2.5 cm. long, 1-5 mm.
wide; heads 50-100, 10-12 mm. long; phyllaries 9, narrowly ovate
or oblong, 6-7 mm. long, 1.5-2 mm. wide, glabrous; flowers 10-12
(-14); corolla 6-8 (-10) mm. long, the tube 3-4 mm. long, the lobes
Jinear or narrowly elliptic, 3.5-4 (-6) mm. long, the throat none;
anthers 2-3 mm. long; style branches 1 mm. long, the tips truncate
with a terminal fringe of trichomes; mature achenes oblong or ellip-
soid, (3.5-) 4-6 mm. long, 1 mm. wide, glabrous, brown, 10- to 12-
ribbed; pappus 1.5-1.6 mm. long, about half as long as the tube of
the corolla, creamy, the bristles unequal; chromosome number, n=30
(Turner, ined.).

DistRIBUTION: Known only from the mountains of western central
Michoacan. Grasslands, openings and wet ravines in the oak-pine-
woodland zone. Flowering in October.

Blake cited both the Pringle 10126 collection at the Gray Herbarium
and at the U.S. National Herbarium as the type. Pringle 10126 (GH)
is chosen as the lectotype because it is annotated in what appears to be
Blake’s writing, and he was associated with the Gray Herbarium at the
time this species was named.

As the name suggests this species has a very short pappus. In
Odontotrichum palmeri the pappus is as short as in O. brachycomum but
O. palmeri differs in having fewer phyllaries and flowers per head and
toothed, but not lobed, basal leaves.

MicHoacikn: Hinton et al. 12146 (MICH, this specimen has an unusually large
leaf, NY, US); Hinton et al. 15288 (GH, NY, US); Hinton et al. 15624 (GH, NY,
TEX, US); King & Soderstrom 4904 (MICH); McVaugh 21934, 21959 (MICH).

390) CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

4. Odontotrichum cervinum Rydb. Bull. Torrey Bot. Club 51:420, 1924. Holo-
type: Nayarit; between Pedro Paulo and San Blascito, Aug. 4, 1897,
J. N. Rose 1986 (US!).

Plants ca. 0.5 m. tall; stem slender, 3-4 mm. thick at the base, sub-
elabrous proximally, becoming sparsely pubescent to tomentose to
densely tomentose upward, the hairs white, multicellular; leaves glabrous,
the abaxial surface paler; basal leaves 3-5, long-petioled, rhombic or
ovate or subcircular in outline, the larger ones about 6-9 cm. long and
wide, pinnately veined; blades deeply pinnately dissected into linear
divisions, 3-4 mm. wide, the major divisions subdivided several times,
the secondary divisions also subdivided; petioles 20-25 cm. long,
slender, glabrous, purplish; cauline leaves 3 or 4, the lowermost similar
to the basal ones; upper cauline leaves reduced, bracteiform, the upper-
most forming inflorescence bracts; inflorescence corymbose, to 10 cm.
long and broad, the branches tomentose; pedicels subtended by a lin-
ear-subulate bract to 1 em. long; calyculate bracts linear-subulate, 4-6
mm. long; heads ca. 25 or fewer, 9-10 mm. long; phyllaries 7, oblong-
ovate, 5-6 mm. long, 1.5-2.5 mm. wide, glabrous; flowers 8-10; corolla
7-8 mm. long, the tube 3.5—4 mm., the lobes 3.54 mm. long, linear or
narrowly elliptic, the throat essentially none; anthers dark brown or
black, 3 mm. long; style branches 0.7-1 mm. long, the tips truncate
with a terminal fringe of trichomes; mature achenes not seen, the
immature achenes greenish, sparsely pubescent with white hairs;
pappus 3.5—4 mm. long, creamy.

DisrrinutTion: Known only from Nayarit. Grasslands and borders
of fields. Flowering in July.

This taxon is known only from one collection besides the type. It
is unusual because of the small, almost subternate leaves. It is related
to O. filicifolium.

Nayarit: McVaugh 15313 (MICH).

5. Odontotrichum cirsiifolium Zucc. Abhandl. Baier. Akad. Wiss. 1:311. 1832.
Holotype: A specimen of a garden plant grown from seeds sent by Kar-
winski from Mexico (M!; Mich. Neg. 1892). Not Cacalia cirsiifolia Hook.
& Arn.

Cacalia cervariifolia [‘cervariaefolia’] DC. Prod. 6:328. 1838. Holotype:
Oaxaca; San Bartolo, Andrieux specimen in the DC. herb. (G, not seen;
Field Mus. Neg. 28888!).

Senecio cervariaefolius (DC.) Schultz Bip. Flora 28:498. 1845.

Sciadoseris vaginata Kunze, Bot. Zeit. 349. 1851. Holotype: specimen grown
in the gardens at Herb. Berol. (not seen).

Mesoneuris bipinnatifida A. Gray, Proc. Amer. Acad. 8:661. 1873. Holotype:
Mountain region of Chiapas, Ghiesbreght 805 (GH!).

Odontotrichum bipinnatifidum (A. Gray) Rydb. Bull. Torrey Bot. Club 51:4138.
1924.

PIPPEN—MEXICAN “CACALIOID” GENERA 391

Plants to 1.5 m. tall; stem 4-7 mm. thick at the base, arachnoid-
pubescent or tomentose (often flocculent), the hairs white, to 2 mm.
long or longer; leaves sparsely pubescent on the adaxial surface,
densely pubescent on the abaxial surface; basal leaves 2 or 3, petiolate,
elliptic or ovate in outline, 18-25 cm. long, 14-22 cm. wide, usually
2-5 (-10) cm. longer than wide; blades 2 or 3 times pinnatifid, the
primary segments 17-21, narrow, opposite proximally becoming alter-
nate distally, 8-12 cm. long, ca. 0.5 cm. wide at the base, each divided
into 5-9 lobes, the secondary lobes often lobed or toothed, the ulti-
mate segments narrowly triangular, ca. 1-2.5 em. long, 1-3 (-7) mm.
wide, the rachis narrowly leafy-margined, the margin ca. 1-3 mm.
wide; petiole and midrib of the blade not clearly distinguished, the
petiole 8-15 cm. long, often narrowly leafy-margined almost to the
base, occasionally lobed; cauline leaves 2 or 3, the lowermost similar
to the basal ones; upper cauline leaves progressively smaller upward,
the bases broadly leafy-margined, auriculate and sheathing the stem,
the blades pinnatisect or lobed; inflorescence corymbose, 20-30 em.
long, 10-20 cm. wide, the branches subtended by leafy, ovate-
naviculate bracts, 1.5-3 cm. long, 1-2 em. wide, distally laciniate;
pedicels 1.5—-4 em. long; calyculate bracts linear, 3-5, up to 1 cm. long;
heads (10—) 30-40 (-50), 10-12 mm. long; phyllaries 11-13, narrowly
ovate, (5-) 6-7 (-9) mm. long, 2-3 mm. wide, glabrous; receptacle ca.
3 mm. across; flowers 20-30 (-40); corolla ca. 7 mm. long, the tube
3-3.5 mm. long, the lobes narrowly oblong, (3.5-) 4 (-5) mm. long, the
throat none; anthers 3 mm. long; style branches 1-1.5 mm. long, hispid
on the abaxial surface, the tips truncate or rounded with a subterminal
or terminal fringe of trichomes; mature achenes obovoid, 5.5-6 mm.
long, 2-3.5 mm. thick, glabrous (in specimens from Oaxaca and
Chiapas) or pubescent with short, unicellular hairs (in specimens from
México, Guerrero, and Michoacan), light brown; pappus creamy or
tawny, 3-3.5 mm. long, the bristles stiff, deciduous; chromosome
number, n=380 (Pippen).

Distripution: Michoacdén, México, Guerrero, and east to Chiapas.
Habitat variable: roadside ditches, grasslands, and openings in the
oak-woodlands. Flowering in August and September.

This species has two forms distinguished by the presence or absence
of pubescence on the achenes and apparently by their somewhat
distinct geographical ranges. The specimens from Oaxaca and Chiapas
have glabrous achenes whereas those specimens from the north and
west of Oaxaca have pubescent achenes. Since no other correlative
differences are evident I do not feel that these forms are worth nomen-

clatural status.

392 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Cutapas: Ghiesbreght 805 (NY, the leaves of this specimen are not as deeply
dissected as the other specimens); Nelson 3117 (GH, US). Guerrero: Hinton
et al. 9435 (MICH, NY). Mé&x1co: Gilly et al. 81 (MICH); Hinton et al. 8211
(MICH, NY); Matuda 28821 (NY); Karwinski s.n., September 1827 (M, probable
parent of type of Odontotrichum cirsiifolium Zucc.). Micnoackn: Pringle 5272
(GH); Pippen 52* (MICH). Oaxaca: Pringle 4984 (GH, NY, US); Nelson 1420
(US); Galeotti 2491 (US).

6. Odontotrichum decompositum (A. Gray) Rydb. Bull. Torrey Bot. Club
51:414, 1924.
Cacalia decomposita A. Gray, Pl. Wright. 2:99. 1853. Holotype: Sonora;
mountains east of Santa Cruz, September 1851, Charles Wright 1286 (GH1]).
Senecio grayanus Hemsl. Biol. Centr. Amer, Bot, 2:241. 1881. Name
illegitimate.
Mesadenia decomposita (A. Gray) Standley, Contr. U.S. Nat. Herb. 19:749.
1915.

Plants up to 1-1.5 m. tall; stem slender, 3-5 mm. thick at the
base, terete, densely pubescent below the lowest leaf with long,
multicellular hairs in flocculent patches, becoming subglabrous or
sparsely pubescent above the lowest leaf; leaves mostly glabrous or
sparsely pubescent, the hairs multicellular, 0.4-1 mm. long, mostly
along the veins and margins; basal leaves 2-4, long-petioled, pin-
nately veined, ovate or elliptic in outline, 20-30 cm. long, 12—27 cm.
wide, about as wide as long or usually 2-5 cm. longer than wide;
blades 3 or 4 times pinnatisect, the primary divisions 10-15 (-17),
opposite below, becoming alternate distally, each 2 or 3 times pin-
natisect, the ultimate segments narrowly triangular or narrowly
oblong and acute-tipped or falcate, 0.5-2 cm. long, 2-5 mm. wide,
the laminar portion of the blade reduced to a narrow margin ca. 1
mm. wide on each side of the main veins; petioles subterete or tri-
angular in cross section, 17-25 cm. long, pubescent below becoming
subglabrous or sparsely pubescent distally; cauline leaves 3 or 4, the
lowermost one-half as large or as large as the basal ones and similar;
upper cauline leaves reduced upward, bracteiform, 4-9 cm. long,
1-4 cm. wide, 1-3 times pinnatisect into linear segments, the upper-
most cauline leaf bracts 1-6 cm. long, 5 mm. wide; inflorescence
paniculate or corymbose, 11-20 cm. long, 12-22 em. broad, the
branches densely pubescent, the major branches subtended by nar-
rowly ovate or linear-subulate bracts ca. 1.5 cm. long; pedicels 1-1.5
mm. long; calyculate bracts 1-3, 24 mm. long, linear-subulate;
heads ca. 100 or fewer, 10-12 mm. long; phyllaries 5 or 6 (rarely 8),
narrowly ovate, obovate or oblong-acute, (4—-) 5-6 (-7) mm. long,
(1-) 1.5-2 mm. wide, mostly glabrous or sparsely pubescent; recep-
tacle 1.5 mm. across; flowers (5-) 6-7 (-8); corolla 7-8 (-9) mm.
long, the tube 3.5-4 (-5) mm. long, the lobes linear-acute, 3.5-4 mm.
long, 0.6 mm. wide, the throat none; anthers 2-3 mm. long; style
branches 1-1.5 mm. long, the tips truncate or obtuse with a terminal

PIPPEN—MEXICAN “CACALIOID” GENERA 393

fringe of trichomes (or rarely a cluster of trichomes covering the
entire tip); mature achenes ellipsoid, often falcate, 4-5 mm. long,
1.5-2 mm. wide, 1 mm. thick, glabrous, dark brown, contrasted
with paler ribs, the ribs 15-18; pappus white or creamy, (4-) 5.5-6
(-7) mm. long; chromosome number, n=30 (Kruckeberg in Ornduff
et al., 1963).

Distrisution: Northwestern Durango, northward into Sonora,
Chihuahua, and Arizona. Its southern limit appears to be northern
Durango as it has not been found farther south. Grasslands, pastures,
and borders of oak-pine woodlands. Flowering in August and
September.

This species is unusual because of its several times pinnatisect
leaves which have the appearance of being compounded. Of the
species of Odontotrichum growing in Mexico this one is the most
northern in range, extending into southern Arizona.

Odontotrichum decompositum, or ‘‘Matarique,” as it is called by
the natives, is the only species of Odontotrichum of known economic
importance. It is used medicinally in the treatment of diabetes and
other ailments (Martinez, 1959).

Arizona: Barkley 14A624 (TEX); Darrow et al. 1235 (NY); Eggleston 10765
(US); Blumer 1378 (GH, NY, US); Kruckeberg 4644 (MICH); Darrow, s.n.,
July 4, 1937 (NY); Harrison & Kearney 6194 (US); 6045 (GH); Kearney &
Peebles 10092 (US); Goodding, s.n., Aug. 28, 1912 (NY); Goodding 784 (GH, NY);
Jones 25053 (GH, NY); Palmer s.n., July 1890 (GH, US); Wilcoz s.n., in 1891
(US). Curnvanva: Gentry 1959, 2815 (GH, US); Hewitt 68 (GH); Knobloch
1293 (MICH); Pringle 767 (GH, MICH, NY, US); Mearns 2219 (NY, US),
527, 531 (US, appear to be stunted plants, collected near the summit); Muller
3420 (GH); Nelson 6095 (GH, US); Townsend & Barber 158 (GH, NY, US);
Harde LeSueur 145 (GH, TEX); Tucker 2456 (US); Townsend s.n., in 1909 (US);
Gentry 536M (MICH). Durango: Nelson 4758 (NY, US). Sonora: Phillips
651 (GH, MICH); White 2741, 3289, 4353 (GH MICH).

7. Odontotrichum filicifolium Rydb. Bull. Torrey Bot. Club 51:414, 1924.
Holotype: Nayarit; between Santa Gertrudis and Santa Teresa, Aug. 8,
1897, J. N. Rose 2101 (US!).

Plants up to at least 0.5 m. tall; stem glabrous proximally, sparsely
pubescent to tomentose distally; leaves glabrous; basal leaves 2-4,
long-petioled, pinnately veined, deltoid or ovate-cordate in outline,
4-5 cm. long and broad; blades deeply and finely dissected, the major
divisions several, each divided into smaller segments or lobes, the
ultimate segments narrow, linear-attenuate, laminar portion of the
blade only a narrow margin, ca. 3-5 mm. wide on each side of the
major veins; petioles slender, 11-12 cm. long; cauline leaves 3 or 4,
the lowermost smaller than the basal ones but similar; upper cauline
leaves bracteiform, dilated at the base, distally laciniate; inflorescence
corymbose, the branches tomentose, subtended by narrowly ovate

394 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

bracts; calyculate bracts 2 or 3, linear-subulate, 2-4 mm. long;
heads 15-25, about 10 mm. long; phyllaries 5 or 6, narrowly ovate,
6-7 mm. long, 2 mm. wide, glabrous; flowers 6-8; corolla ca. 7.5 mm.
long, the tube 4 mm. long, the lobes 3.5 mm. long, the throat none;
anthers ca. 3 mm. long; style branches 0.7—1 mm. long, hispid on
the abaxial surface, the tips obtuse with subterminal fringe of tri-
chomes; mature achenes not seen, the immature achenes densely villous;
pappus fuscous, 7 mm. long.

Distrisution: Known only from the type specimen.

This taxon is quite similar to Odontotrichum cervinum and was
collected in the same region. It differs from 0. cervinum in having
smaller heads and the larger leaves. Field observations and study of
future collections may indicate that these two taxa should be con-
sidered as one species.

8. Odontotrichum globosum (Rob. & Fern.) Rydb. Bull. Torrey Bot. Club 51:418,
1924,

Cacalia globosa Rob. & Fern. Proc. Amer. Acad. 30:119. 1894. Holotype:
Chihuahua; in a moist meadow, Guachuchic, June 25, 1892, Hartman 522
(GH!).

Odontotrichum scabrum Rydb. Bull. Torrey Bot. Club 51:416. 1924.

Cacalia pratensis Stand]. Publ. Field Mus. Bot. 22:61. 1940, Holotype:
Chihuahua; wet meadows near San Juanito, 2,430 m., July 26, 1937, F.
Shreve 8031 (F!, isotype MICII!).

Odontotrichum pratense (Standl.) J. Cuatr. Brittonia 8:157. 1955,

Piants 0.5-0.7 (-1.2) m. tall; stem slender, 5 mm. thick or less at
the base, mostly subglabrous below becoming sparsely pubescent to
arachnoid-tomentose above; leaves glabrous or very sparsely puber-
ulent mostly along the veins; basal leaves 2-4, long-petioled, pinnately
veined, cordate or cordate-ovate, 4-7 (-8) cm. long and broad; blades
shallowly dentate or entire, the teeth rounded or triangular, the
marein of the blade ciliate; petioles glabrous, slender, 12-21 (—30)
cm. long; cauline leaves 3 or 4, the lowermost similar to and as large
as the basal ones; upper cauline leaves progressively smaller upward,
the uppermost bracteiform, leafy, ovate, clasping the stem; inflores-
cence globose, very compact, 2.5-3 cm. long and broad, the branches
arachnoid-tomentose; pedicels 5 mm. long or less; calyculate bracts
1-5, narrowly ovate or linear-subulate, 4-5 mm. long, | mm. wide or
less; heads about 20, 7-10 mm. long; phyllaries 5 or 6, narrowly ovate
or ovate, 5-7 mm. long, 1.5-2 mm. wide, glabrous; flowers (4—-) 5-6
(-8); corolla 6-7 (-10) mm. long, the tube 3-3.5 (-5) mm. long, the
lobes 3-3.5 (-5) mm. long, 0.5 mm. wide; anthers 2-3 (-3.5) mm.
long; style branches ca. 1 mm. long, the tips truncate or obtuse with
a subterminal fringe of trichomes; mature achenes pale brown, gla-
brous; pappus creamy, 3-3.5 (-5) mm. long.

PIPPEN

MEXICAN “CACALIOID” GENERA 395

Distrisution: Chihuahua and Durango. Wet marshy areas in
pastured grasslands. Flowering in July and August.

This species is unique because of its habitat—most of the other
species in this genus grow in dryer situations—and the very tightly
globose inflorescence.

Comparison of the type of Cacalia pratensis Standl. with the type
of Odontotrichum globosum indicates that they are definitely con-
specific because they are similar in all respects.

Evidence of natural hybridization between 0. globosum and 0.
stnuatum was observed by Dr. McVaugh in southern Durango in 1962
and a sampling of these populations was made (voucher specimens,
MeVaugh 21732-21739, MICH). The hybrid population showed
various degrees of ecological and morphological intergradation be-
tween the two very dissimilar parents.

Odontotrichum scabrum, described by Rydberg from a specimen
collected in 1897 (Rose 3471, US), is one of these intermediate hybrid
forms, resembling O. globosum in leaf shape but with a more open
inflorescence like O. sinuatum. It is not distinct enough to deserve
nomenclatural status.

Cuinvuanua: Knobloch 1273 (MICH); Shreve 8031 (US); Gentry 2785 (MICH,
GH, isoparatype of C. pratensis Standl.). Durango: Graham 99 (MICH);
McVaugh 21732, 21734-21739 (MICH, intermediate forms between O. globosum
and O. sinuatum); Pennell 18532 (NY); Nelson 4769 (NY); Rose 3471 (US, holo-
type of O. scabrum Rydb.).

9. Odontotrichum goldsmithii (B. L. Rob.) Rydb. Bull. Torrey Bot. Club 51:416.
1924,
Cacalia goldsmithii B. L. Rob. Proc. Amer. Acad. 43:45. 1907.

Slender plants to 1 m. tall; stems up to 5 mm. thick at the base,
mostly glabrous or sparsely pubescent above; leaves mostly glabrous
or sparsely puberulent along the veins and margins; basal leaves
2 or 3, erect, long-petioled, pinnately veined, variable in shape from
ovate to broadly ovate, 7-14 (-19) cm. long, 6-12 (-16) cm. wide,
as long as wide to twice as long as wide; blades mostly truncate or
obtusely based, occasionally slightly cordate, the margins sinuately
double-dentate or lobulate, the teeth deltoid-rounded, in two alter-
nating series of large and small respectively; petioles 8-14 (-20) cm.
long, slender; cauline leaves ca. 3, the lowermost usually elliptic,
as large as or one-half as large as the basal ones and similar; upper
cauline leaves greatly reduced, the uppermost bracteiform, leafy,
up to 6 cm. long, 0.5-1 cm. wide, the base of the petiole dilated;
inflorescence loosely corymbose, 18-23 cm. long, 9-11 cm. broad,
the branches densely pubescent, subtended by subulate or narrowly
ovate bracts 1-3 cm. long, 1-5 mm. wide, smaller distally; pedicels
5-15 mm. long; calyculate bracts 3 or more, subulate, 1.5-4 mm.

266—-836—68——-3

396 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

long; heads 50-75, 10-11 mm. long; phyllaries (5-) 7 or 8, usually
ovate-oblong or narrowly ovate, 5-6 (-7) mm. long, (1-) 2-3 mm.
wide, glabrous; receptacle up to 1.5 mm. across; flowers 7-13; corolla
mostly 6-8 mm. long, the tube 3-4 mm., the lobes 3-4 (-7, Hinton
9202) mm. long, narrowly elliptic, strongly 3- (-5-) nerved; anthers
2-3 mm. long; style branches 1 mm. long, usually hispid on
the abaxial surface, the tips truncate with a terminal fringe of tri-
chomes; mature achenes ellipsoid, 4-5 mm. long, 2 mm. wide, ca. 1
mm. or less thick, brown, glabrous, 10- to 12-ribbed; pappus creamy
white, becoming fuscous in age, 3.5-5 (-7) mm. long.

Disrrispution: Eastern Jalisco to Michoacdn and western Guerrero
and México. Grasslands and pastures. Flowering in July (and
August?).

This species is very closely related to O. silphiifolium, and further
field and garden study may indicate that they should be considered
as the same species. In the present treatment, however, they are
recognized as two species with O. goldsmithi differing from OQ. sil-
phiifolium in its slender, mostly glabrous habit, open, loose inflores-
cence with shorter heads, truncate or obtuse leaf bases, and smaller
cauline leaves that are not auriculate-based.

Key to Varieties of O. goldsmithit

a. Heads with 5 or 6 phyllaries, 6 or 7 flowers; leaves sparsely puberulent.
9b. O. goldsmithii var. rowellii
a. Heads with 7 or 8 phyllaries, 9-13 flowers; leaves glabrous.
9a. O. goldsmithii var. goldsmithii

9a. Odontotrichum goldsmithii var. goldsmithii
Cacalia goldsmithii B. L. Rob. as to Holotype: Jalisco; Hacienda San Marcos,
350 m., July 12, 1905, P. Goldsmith 8 (GH!).

Phyllaries mostly 8; flowers 9-13 per head; basal leaves ovate or
elliptic, glabrous, the blades truncate or obtuse-based, rarely cordate.

DistriBuTion: Jalisco, Michoacén, Guerrero (Mina Distr.), and
México.

Guerrero: Hinton et al. 9202 (MICH, NY, this specimen has larger heads
than the other specimens). Jatisco: McVaugh 15460, 16132 (MICH). México:
Hinton 1266 (MICH); Hinton et al. 8066, 7984 (MICH, NY). Micnoacin:
Hinton et al. 5078 (GH, NY, US).
9b. Odontotrichum goldsmithii var. rowellii Pippen var. nov. PLATE 2

A varietas goldsmithii floribus phyllariisque paucis, foliis radicalibus,
puberulibus differt.

Heads with 5 or 6 phyllaries, 6 or 7 flowers; basal leaves ovate or
narrowly ovate, the blades obtuse or cordate-based, sparsely
puberulent.

PIPPEN—MEXICAN “CACALIOID” GENERA 397

Ho.otype: Guerrero; 1 mi. N. of Agua del Obispo, 3,300 ft., July
1, 1952, C. M. Rowell, Jr. 2991 (MICH!).

Distrisution: Known only from central Guerrero.

This variety is distinguished from the other in the smaller heads
with fewer phyllaries and flowers and the somewhat puberulent leaves.

GuERRERO: Wilkinson & Rowell 3381 (MICH); Rhymes & Rowell 3886 (MICH);
Hicks & Rowell 3604 (MICH); Rzedowski 16073 (MICH); Sharp 441431 (NY).
10. Odontotrichum multilobum Pippen, sp. nov. PLATE 3

Plantae tenues ca. 1 m. altae, caule foliisque glabris vel sparse
pubescentibus, foliis radicalibus erectis, latis ellipticis 18-30 cm. longis
10-17 cm. latis, profunde pinnatifidis, sezmentis principalibus 12-15
linearibus; petiola tenus glabra 10-30 cm. longa; folia culaine bractei,
forma 1-2; inflorescentia corymbosa, 16-30 cm. long, 11-16 cm. lata-
bracteis 10-20 mm. longis, linearis vel anguste triangularis; capitula
50-100, ca. 12 mm. longa; phyllaria 8, 5-6 (-7) mm. longa, 1.5—2 mm.
lata, anguste ovata, glabra; flores 10, corollis 7 mm. longis, tubo lobis-
que 3.5 mm. longis, achaenis ellipsoidea vel obovata, (8-) 3.5-4 mm.
longa, 1.5-2 mm. lata, glabra, pallide fulva; pappus sordidus (5-) 6
(-7) mm. longus.

Plants ca. 1 m. tall; stem and leaves glabrous or sparsely pubescent,
becoming sparsely or densely pubescent in the inflorescence; flowering
stem 4-6 mm. thick at the base; basal leaves 3 or 4, long-petioled,
erect, pinnately veined, broadly elliptic in outline, 18-30 cm. long or
longer, 10-17 cm. broad; blade deeply pinnatifid, the laminar portion
narrow, ca. 3-6 mm. wide on each side of the vein, the primary lobes
12-15, linear, 7-11 cm. long, each divided into 3-5 major lobes, the
secondary lobes either 1- to 2-lobed or coarsely toothed, the ultimate
segments linear-acute; angle of the lobes with the midrib less than 90
degrees; petioles ca. 10-30 cm. long, mostly glabrous, triangular in
cross section; cauline leaves 1 or 2, similar to the basal leaves but
much smaller; inflorescence loosely corymbose, 16-30 cm. long, 11-16
cm. broad, the branches subtended by linear or narrowly triangular bracts
10-20 mm. long, 1-4 wide; calyculate bracts 3-5, subulate, 3-5 mm.
long; heads 50-100, ca. 12 mm. long; phyllaries 8, narrowly ovate,
5-6 (-7) mm. long, 1.5-2 mm. wide, glabrous; receptacle 2 mm.
across; flowers 10; corolla 7 mm. long, the lobes as long as the tube,
3.5 mm. long, linear, the throat none; anthers 3 mm. long; style
branches 1 mm. long, the tips truncate or rounded with a subterminal
fringe of trichomes; mature achenes (3-) 3.5-4 mm. long, ellipsoid or
obovoid, 1.5-2 mm. wide, 1 mm. thick, glabrous, light brown, ca.
20-striate; pappus (5—-) 6 (-7) mm. long, creamy or tawny; chromo-
some number, n=30 (Pippen).

398 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Houortype: Jalisco; ca. 15 mi. SSE. of Autlin by way of Chante,
oak-dominated slopes E. of Mamantlan, ca. 4,500 ft., Aug. 1, 1949,
R.L. & C. R. Wilbur 2094 (MICH!).

The type specimen is somewhat smaller in stature than the other
specimens of the same taxon, but it is representative.

This species is related to O. pringlet by the shape of the leaf, but
differs in that the achenes are glabrous; and to O. sinuatum, differing
in the more cylindrical heads, rather than turbinate, and in the lower
pair of lobes on the leaves not deflected.

This species is sympatric with O. pringlet and O. palmeri, possibly
intergrading with the latter. Two specimens from western Jalisco
(Pippen 29, sheets 1 and 2, MICH) are intermediate between 0.
multilobum and O. palmeri, suggesting that hybridization may be
occurring. At this locality both O. multilobum and O. palmeri as well
as the intermediate forms were growing together, the latter plants
mistaken for a third species at the time of collection. Future field
studies and examination of additional material are necessary to
determine if integration has occurred.

Jatisco: Pippen 25, 30*, 29 (possible hybrid between O. multilobum and O.
palmert), 40 (MICH).

11. Odontotrichum napellifolium (S. Schauer) Rydb. Bull. Torrey Bot. Club.
51:419. 1924.
Cacalia napellifolia S. Schauer, Linnaea 19:732. 1847. Holotype: Mexico;
about 1840, Aschenborn 312 (B, not seen; tracing of type in Klatt’s Herb.,
GH)).

Plants 0.5-0.6 m. tall; stem slender, 2-4 mm. thick at the base,
densely pubescent below, becoming sparsely pubescent upward;
leaves mostly glabrous, the margin ciliate; basal leaves 4-7, long-
petioled, subpalmately veined with 5-7 principal veins, subcircular
in outline, the base cordate, 2.5-5 em. long from the basal sinus to
the tip, 3-8 cm. wide; blades deeply lobed, the primary lobes 3-5
(-7), each deeply divided into 2 or 3 (-4) lobes, the secondary lobes
again divided into 3-5 lobules, the ultimate segments narrowly tri-
angular or deltoid, 3-10 mm. long, the proximal pair of lobes de-
flected, parallel to the petiole, the base of the blade decurrent on the
petiole for a short distance; petioles 7-15 em. long, slender; cauline
leaves 1 or 2, the lowermost similar to the basal ones; upper cauline
leaves bracteiform, reduced, 2-3 cm. long, 0.5-1 em. wide, 5- to 7-
lobed; inflorescence corymbose, the branches tomentose, the pedicels
0.5-3 cm. long; calyculate bracts about 3, subulate, ca. 4 mm. long;
heads 5-8, 10-14 mm. long; phyllaries 10, narrowly ovate, 5-6 mm.
long, 1.5-2 mm. wide, sparsely pubescent, the tips purplish; flowers
12-25; corolla 6-7 mm. long, the tube 3-3.5 mm. long; anthers 2.4-3
mm. long; style branches 0.7-1 mm. long, hispid on the abaxial

PIPPEN—MEXICAN “CACALIOID” GENERA 399

surface, the tips truncate or obtuse with a subterminal fringe of
trichomes; mature achenes ellipsoid, 5 mm. long, 1.5 mm. wide,
1 mm, thick, pale tan or creamy, 10- to 12-ribbed, the ribs pubescent
with ascending hairs; pappus creamy, 3-5 mm. long.

DistRiBpuTION: Guerrero and Hidalgo. Grasslands and openings
in oak-pine woodlands. Flowering in June and July.

This taxon is distinct because of the small, deeply lobed basal
leaves. It is closely related to O. filicifolium and O. cervinum but differs
from both in the larger number of flowers per head.

GUERRERO: Kubicek & Rowell 3779 (MICH); Rhymes & Rowell 3896 (MICH).
HipauaGo: Pringle 10024 (MICH, NY, TEX, US); Salazar s.n., Aug. 1, 1918 (US).

12. Odontotrichum pachyphyllum (Schultz Bip.) Rydb. Bull. Torrey Bot. Club
51:417, 1924.
Cacalia pachyphylla Schultz Bip. ex Seem. Bot. Voy. Herald. 310. 1856.
Holotype: Northwestern Mexico; Seemann 1999 (K, not seen; MICH Neg.
6221).
Senevio sclerophyllus Hemsl. Biol. Centr. Amer. Bot. 2:247. 1881. Based on
Cacalia pachyphylla Schultz Bip. not S. pachyphyllos Remy.

Plants 0.5-0.7 m. tall; stem 3-4 mm. thick at the base, pubescent,
the hairs white, multicellular, often flocculent; leaves mostly glabrous
or sparsely pubescent along the veins; basal leaves 2 or 3, coriaceous,
long-petioled, reniform or rounded-deltoid, rarely subcircular in out-
line (6-) 8-10 (-11) em. long from the basal sinus to the apex, (10—)
13-18 (-20) cm. wide, the tip broadly rounded, the base truncate or
usually shallowly cordate; blades variable, shallowly repandly dentate
or subentire, the teeth rounded or obtuse, the adaxial and abaxial
surfaces very similar; petioles (4—) 8-18 (-35) cm. long; cauline leaves
2 or more, the lowermost smaller than the basal ones but similar;
upper cauline leaves becoming bracteiform upward with the petiole
broadly leafy-margined, clasping the stem; inflorescence corymbose,
7-20 cm. long, 7-15 cm. broad, the branches tomentose or villous
(glabrous in the type and isotype); pedicels 2-6 cm. long; calyculate
bracts 3-5, subulate, 9-13 mm. long, 1 mm. wide, sparsely pubescent;
heads 9-15, 1-1.5 cm. long; phyllaries 13 or 14, narrowly ovate, 7-9
mm. long, 1-3 mm. wide, glabrous; flowers 30-35; corolla (7.5—) 8-10
mm. long, the tube (4—) 5-6 mm. long, the throat short, ampliate, up
to 0.5 mm. long, 1 mm. wide at the apex, occasionally lacking, the
lobes 3.5-4 mm. long; anthers 2-3 mm. long; style branches 1.5 mm.
long, the tips truncate or obtuse with a terminal or subterminal fringe
of trichomes; mature achenes ellipsoid or oblong, 4-5 mm. long, 2 mm.
wide, 1.5 mm. thick, glabrous, brownish, 15- to 16-ribbed; pappus
white or tawny, 6-8 mm. long.

DistrisuTion: Known only from southern Durango. Openings in
pine-oak woodlands. Flowering in August and September.

400 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

The type is mostly glabrous with linear inflorescence bracts and
large basal leaves with acute basal sinuses. These characteristics are
somewhat different from the other specimens studied but the speci-
mens are all similar enough in size of heads, number and size of
flowers, texture and general shape of the leaves to be considered
part of the same taxon.

This species is closely related to O. amplum, differing from it in
having smaller heads with narrower inflorescence bracts and basal
leaves rounded at the base.

Duranao: Gentry 6969 (MICH, US); McVaugh 21659 (MICH); Pennell 18332
(NY); Waterfall 12668 (MICH); McVaugh 21731 (shows possible intergradation
with O. globosum), 21729 (MICH); Ownbey & Ownbey 1874 (MICH, US); Maysilles

7784 (MICH). Locauiry UNKNowN: Seemann s.n. (probably isotype of Seemann
1999, GH).

13. Odontotrichum palmeri (Greene) Rydb. Bull. Torrey Bot. Club 51:417. 1924.
Cacalia palmert EX. L. Greene, Pittonia 1:219. 1888. Holotype: Jalisco;
Rio Blanco, in 1886, #. Palmer 168 (ND!, isotypes GH!, US!, NY!).

Plants 0.7-1 (-1.5) m. tall; stems 3-5 (-7) mm. thick at the base,
sparsely or densely pubescent, rarely subglabrous, hairs 1.5-3 mm.
long, stiff, white; leaves sparsely or densely pubescent, the pubescence
denser on the abaxial surface; basal leaves 2-3 (-5), pinnately veined,
coriaceous, broadly ovate or occasionally subcircular in outline, 9- 17
(-23) em. long and broad (broadest below the middle); blades coarsely
dentate or shallowly lobulate, the lobules up to 1 em. long, 2 em.
wide, each 3-angulate, mucronate; petioles (5-) 6-15 (-30) cm. long;
cauline leaves several, the lowermost as large as or one-half as large
as the basal ones and similar, the petiole dilated and clasping the stem
at the base, often leafy-margined; upper cauline leaves reduced and
bracteiform, the petiolar portion broadly leafy-margined and clasping
the stem, the blade portion ovate, dentate; inflorescence corymbose,
9-12 (-18) em. long, 10-15 em. broad, the branches sparsely pubes-
cent, the distal branches subtended by narrowly ovate or subulate
bracts up to 1.5 mm. long, the proximal branches in the axils of the
uppermost leaves; calyculate bracts usually 3, subulate, 5 mm. long
or less; heads up to 100, ca. 1 em. long; phyllaries 5 or 6 (-7), narrowly
ovate, 5-7 (-8) mm. long, 1.5 (-3) mm. wide, glabrous or sparsely
pubescent, tightly imbricated forming a narrow cylindrical involucre,
the tips attenuate; receptacle often sparsely pubescent with long,
mult cellular hairs; flowers 5 or 6 (-8); corolla 6.5-9 mm. long, the
tube (3—) 3.5-4 mm. long, the lobes 3.5-5 (—5.5) mm. long, the throat
none; anthers 2-3 mm. long; style branches 1-1.5 mm. long, the tips
truncate or rounded with a terminal fringe of trichomes; mature
achenes 3.5-4 mm. long, 1.5-2 mm. wide, ellipsoid, glabrous, promi-
nently 10- to 12-ribbed, the ribs dark red purple, sharply contrasted

PIPPEN—MEXICAN “CACALIOID” GENERA A401

by pale golden-tan bands separating them; pappus creamy or white,
(1-) 1.5 (-2) mm. long, about one-half as long as the corolla tube;
chromosome number, n=about 25 (Pippen).

DistriBuTion: Southern Durango, Nayarit, Jalisco, and Micho-
acén. Common in grasslands, pastures, and borders of oak-pine
woodlands. Flowering in August and September.

This species is similar to 0. brachycomum in having an extremely
short pappus but differs in having smaller heads with fewer flowers
and phyllaries, and the leaves not lobed.

Rydberg described Odontotrichum nephrophyllum from a specimen
collected in southern Durango (Rose 2248, US). It differs from 0.
palmeri in that the pappus is as long as the corolla tube and the
petioles of the basal leaves are densely pubescent. Examination of
future collections may indicate that this is distinct from O. palmert,
but at the present I do not feel that it is significantly different.

Duranao: Rose 2248 (US). Jauisco: Diguet s.n., July 1912 (MICH); Graham
124 (MICH); McVaugh 20481, 12929 (MICH); McVaugh & Koelz 441 (MICH);
King 3659 (MICH, NY, TEX, US); Powell & Edmondson 869 (TEX); Pringle
2304 (GH, NY, US); Pringle 11500 (GH, MICH, US); Pippen 10, 20, 27A, 32,
28 (MICH). Micnoackn: Arsene 5132 (GH, NY, US); Feddema 183 (MICH);
Leavenworth 539 (NY, MICH); Pippen 43 (MICH); McVaugh 21960 (MICH).
Nayarit: Feddema 308 (MICH); Pippen 12* (MICH).

14, Odontotrichum paucicapitatum (Rob. & Greenm.) Rydb. Bull. Torrey Bot.
Club 51:414. 1924.

Cacalia paucicapitata Rob. & Greenm. Amer. Journ. Sci. IIT. 50:158. 1895.
Holotype: Oaxaca; Sierra de Clavellinas, 7,000 ft., Oct. 25, 1894, C. G.
Pringle 6018 (GH!, isotypes GH!, US!).

Plants to 1 m. tall; caudex apparently fleshy, globose, at least 2
em. long, 2.5 em. wide; stem ca. 5 mm. thick at the base, densely
covered with a flocculent interwoven mass of long, grayish-white
hairs persisting through the inflorescence; leaves glabrous on the
adaxial surface, the abaxial surface arachnoid or tomentose; basal
leaves 2 or 3, petioled, pinnately veined, oblong in outline, 14-16 cm.
long, ca. 10 cm. wide; blades deeply pinnately lobed, the primary
lobes 9 or 10, at right angles to the axis but ascending distally, extend-
ing ca. four-fifths of the distance to the midrib, each irregularly 3- to
5-lobed; petioles 5-12 cm. long, slender; cauline leaves ca. 3, the
lowermost about one-half as large as the basal ones and similar;
upper cauline leaves reduced and bracteiform, 4-6 cm. long, ca. 1 cm.
wide; inflorescence subracemose; pedicels (2—) 4-6 cm. long; calycu-
late bracts 3-5, linear-subulate, 1-2 cm. long; heads 5-7, 2-2.5 cm.
long; phyllaries 12 or 13, oblong or narrowly ovate, 10-12 mm. long,
3-4.5 mm. wide, densely pubescent; flowers 40 or more; corolla 11-14
mm. long, the tube 7-10 mm. long, the throat narrowly campanulate,
1-1.5 mm. long, equally broad at the apex, the lobes ca. 3 mm. long,

402 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

linear; anthers 3 mm. long; style branches 1.5 mm. long, hispid dis-
tally on the abaxial surface, the tips truncate with a terminal fringe
of trichomes; mature achenes ellipsoid, obovoid or 3-angulate, 5-6
mm. long, 2-3.5 mm. wide, 1.5 mm. thick, densely villous, the hairs
ascending, grayish-white or creamy, up to 1 mm. long; pappus
creamy, 7-8 mm. long, clavate distally.

Distripution: Known only from the type specimens.

This species is most unusual because of the dense, grayish-white,
flocculent pubescence covering the entire plant except the adaxial
surface of the leaves. The leaves are very much like those of Q.
sinuatum but the heads are much larger and the flowers have a short,
but evident throat. The apparently fleshy caudex is also unique.

15. Odontotrichum platylepis (Rob. & Seat.) Rydb. Bull. Torrey Bot, Club 51:
417, 1924.

Cacalia platylepis Rob. & Seat. Proc. Amer, Acad. 28:110. 1893. Lectotype:
Jalisco; Rio Blanco, October 1886, F. Palmer 689 (GH!).

Cacalia coriacea M. I, Jones, Extracts from Contr. West. Bot. 18:80. 1933.
Holotype: Jalisco; Orendain, Nov. 28, 1930, M. E. Jones s.n. (POM, not
seen; isotype GH!).

Plants to 1.5 m. tall; stem 6-12 mm. thick at the base, densely
shaggy-villous below, becoming less dense upward; leaves sparsely
pubescent mostly along the veins, the abaxial surface densely pubes-
cent; basal leaves 3 or 4, long-petioled, erect, coriaceous, broadly ovate
in outline, 18-35 (—75) cm. long and broad or slightly broader; blades
deeply pinnately lobed two-thirds to three-fourths the distance to the
midrib, the primary lobes ca. 11, the proximal pair (6—-) 10-16 (—22)
em. long, 2-6 em. wide at the base, the distal lobes progressively
smaller upward, each lobe irregularly divided into 2-4 lobes, the major
secondary lobes often again lobed, the ultimate segments triangular or
deltoid or oblong-obtuse, the base of the blade decurrent on the petiole
for a short distance forming a leafy margin, the abaxial and adaxial
surfaces very similar; petioles 16-32 cm. long, triangular in cross sec-
tion, densely pubescent below becoming sparsely pubescent distally;
cauline leaves several, the lowermost smaller than the basal ones but
similar; upper cauline leaves bracteiform, the petiolar portion leafy-
margined, clasping the stem, the blade portion sinuately lobed, the
uppermost forming inflorescence bracts, the larger ones 3-6 cm. long,
1.5-4 cm. wide, distally laciniate; inflorescence corymbose, 16-27 cm.
long, 16-18 cm. broad, the branches sparsely or densely arachnoid
pubescent, the hairs frequently with purple crosswalls; pedicels 1.5-3
(-9) em. long; calyculate bracts 5-8, narrowly obovate, 1.5—-2 cm.
long, 5-10 mm. wide, laciniate-lobed; heads 15 or fewer, 1.5-2 cm.
long; phyllaries 13 or 14 (-17), narrowly ovate or elliptic, 13-15 (-16)
mm. long, 4-5 mm. wide, mostly glabrous, often sparsely puberulent;

PIPPEN—MEXICAN “CACALIOID” GENERA 403

receptacle to 1 cm. across; flowers 40-80 or more per head; corolla 13-
15 (-16) mm. long, the tube 10-11 mm. long, the lobes (3.5-) 4—5 (-6)
mm. long, the throat none; anthers 3.5-4 mm. long; style branches
1-1.5 mm. long, the tips truncate with a subterminal fringe of tri-
chomes; mature achenes ellipsoid, 6-7 mm. long, 3 mm. wide, 1.5-2
mm. thick, pale with ca. 10 contrasting dark ribs, the intervals between
the ribs pubescent with hyaline, ascending hairs; pappus white, (8-)
10-11 mm. long, clavate distally.

Distrisution: Mostly in the central grassland regions of Jalisco
around Guadalajara and extending northward to Aguascalientes.
Open, rocky grasslands, scrub and pastured areas. Flowering from
mid-October to December, the flowers emitting an unpleasant odor.

In the original description two specimens were cited as syntypes:
Jalisco; plains of Guadalajara, Nov. 26, 1888, C. G. Pringle 1816
(GH!) and Jalisco; Rio Blanco, October 1886, H. Palmer 689 (GH1!).
The latter was designated the lectotype because it is a more complete
specimen than Pringle 1816. The lectotype specimen consists of a
basal leaf ca. 18 cm. long, a stem in three pieces, with 6 cauline leaves
below the 8-headed inflorescence (Mich. Neg. 1253). There is also
a second sheet at the Gray Herbarium (an isolectotype) that has a
basal leaf ca. 28 cm. long, the base of the plant with the roots, and
an inflorescence with ca. 15 heads, but no portion of the stem.

This species is spectacular because of its extremely large basal
leaves, more than one-half meter long, that stand erect.

AGUASCALIENTES: McVaugh & Koelz48 (MICH). Jatisco: Jones 27692 (NY);
Palmer 689 (US, 3 sheets, isotypes); McVaugh 22123 (MICH); Pringle 1816
(US, 2 sheets, NY, MICH, isoparatypes); Pringle 9873 (GH, US, NY); Safford
1432 (US); Jones s.n., Nov. 27, 1930 (GH, isotype of C. coriacea Jones); Pippen
23, 58 (MICH); Jony s.n., in 1892 (US).

16. Odontotrichum pringlei (S. Wats.) Rydb. Bull. Torrey Bot. Club 51: 415.
1924, PLATE 4
Cacalia cirsitfolia Hook. & Arn. Bot. Beech. Voy. 436. 1841. Holotype:
Nayarit; between Tepic and San Blas, Sinclair s.n., in Hooker Herb.

(K, not seen; Mich. Neg. 623!). Not Odontotrichum cirsiifolium Zucc.

Senecio cirsiifolia (Hook. & Arn.) Schultz Bip. Flora 25:499. 1845.

Cacalia pringlei 8. Wats. Proc. Amer. Acad. 25:156. 1890. Lectotype: Jalisco;
grassy slopes of the barranca near Guadalajara, November 1888, C. G.
Pringle 1749 (GH!).

Plants to 1.5 m. tall; stem 4-10 mm. thick at the base, densely
villous, flocculent below the lowest leaf and sparsely puberulent or
becoming glabrous above; leaves sparsely pubescent with 8- to 10-
celled hairs 0.3-0.5 mm. long, slightly longer and denser along the
veins and on the abaxial surface; basal leaves 2-4, erect, coriaceous,
long-petioled, pinnately veined, oblong or elliptic in outline, 20-35
(-40) cm. long, 13-30 cm. wide, as long as broad to twice as long as

404 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

broad; blades deeply pinnately lobed, the primary lobes 10-16, (4-)
5-10 (-15) cm. long, 3-4 cm. wide at the base, each usually divided
into 3 major lobes and 2 or 8 lesser lobules or teeth, the latter usually
narrowly triangular, the secondary lobes triangular or 3-angulate or
again divided into 2 or 3 triangular lobules, the lobes, including the
proximal pair, forming an angle with the midrib of less than 90
degrees; blades obtuse or attenuate, not cordate at the base, decurrent
along the petiole for 1-2 cm. below the proximal lobes; petioles 20-30
(-40) em. long, triangular in cross section, villous proximally, be-
coming sparsely pubescent distally; cauline leaves 3-7, much reduced
in size, bracteiform, the lowermost leaflike, sinuately lobed, the
petiole dilated at the base and clasping the stem, up to 15 cm. long,
3 em. broad; inflorescence loosely paniculate or corymbose, 18-25
cm. long, 16-20 cm. broad, the branches subglabrous to sparsely
pubescent, subtended by narrowly ovate bracts (2-) 3-5 (-7) em.
long, 0.5-1.5 cm. wide; calyculate bracts 5-8, linear-subulate, 3.5-5.5
mm. long; heads ca. 100, 10-12 mm. long; phyllaries 7 or 8 (9),
ovate or elliptic, 5-7 mm. long, 2-3 (-4) mm. wide, glabrous; recep-
tacle 1.5-2 mm. across; flowers 11-14; corolla 6-8 (-10) mm. long,
the tube 3-4 (-5) mm. long, the lobes 3-4 (-5) mm. long, linear, the
median nerve strong, the throat none; anthers 2-3 mm. long; style
branches 1 (-1.5) mm. long, usually hispid on the abaxial surface, the
tips truncate or obtuse with a subterminal or terminal fringe of
trichomes; mature achenes ellipsoid (3-) 4-5 mm. long, 2 mm. wide,
1-1.5 mm. thick, pubescent with minute, creamy or tawny hairs
between the ribs, the ribs about 20, darker than the area between
the ribs; pappus (4-) 6-7 mm. long, white, the tips slightly clavate;
chromosome number, n=30 (Pippen; Turner, ined.).

Two specimens were cited in the original description as syntypes:
Jalisco; grassy slopes of the barranca near Guadalajara, November
1888, C. G. Pringle 1749 (GH) and 1811 (GH). Pringle 1749 is chosen
as the lectotype because it is a more complete specimen.

DistriBuTion: Nayarit, Jalisco, and Colima. Grasslands, pastured
areas, roadsides, and less frequently rocky slopes in oak-pine wood-
lands. Flowering from October to early December.

This species is closely related to O. platylepis and O. sinuatum,
differing from the former in having smaller heads; from the latter in
having pubescent achenes and differently shaped leaves. Cacalia
cirsiifolia Hook. & Arn., known only from the very incomplete type
specimen consisting of just a portion of the inflorescence and one
upper cauline leaf, is undoubtedly conspecific with O. pringleit because
of the similarity of the heads and the pubescent achenes.

Couima: Palmer 1234 (GH, NY, US). Jauisco: McVaugh & Koelz 1253, 1104
(MICH); McVaugh 13616, 21765* (MICH); Pippen 37*, 41, 59 (MICH); Pringle

PIPPEN—MEXICAN “CACALIOID” GENERA 405

1811 (US, isoparatype of C. pringlet 8. Wats.); Pringle 11499 (GH, MICH, US).
Nayanit: Ferris 5881 (US); Mexia 845 (GH, US); McVaugh & Koelz 601 (MICH);
Palmer s.n., Jan, 5-Feb. 6, 1892 (US); Barclay s.n. (GH, probably collected in late
December 1837 at the same time and place as the type of Cacalia cirsiifolia
Hook. & Arn., since Barclay and Sinclair were together at that time, attached to
the H.M.S. Sulphur [McVaugh, manuscript ined.]).

17. Odontotrichum purpusii (Greenm.) Rydb. Bull. Torrey Bot. Club 51:418.
1924,
Cacalia purpusii Greenm. Univ. Calif. Publ. Bot. 4:95. 1910. Holotype:
Puebla; Cacalote, August 1909, C. A. Purpus 3845 (F!, isotype NY!).

Plants to 0.6 m. tall, pubescent with white, stiff, multicellular hairs;
stem slender, ca. 3 mm. thick at the base; basal leaves 2-4, pinnately
veined, petioled, subcoriaceous, ovate or broadly elliptic, to 8 em.
long, 6 cm. wide, the margin of the blade doubly dentate with two alter-
nating series of large and small teeth, respectively, the base of the
blade truncate or obtuse; petioles terete, 3-9 cm. long, about as long
as the blade; cauline leaves ca. 4, the lowermost smaller than the basal
ones but similar to them; upper cauline leaves reduced, bracteiform;
inflorescence compact or spreading, corymbose, 3-14 cm. long, 3-19
cm. broad, the branches arachnoid-pubescent or tomentose, subtended
by linear-subulate bracts ca. 5 mm. long; calyculate bracts 3-7,
linear-subulate, 3-4 mm. long; heads 15-30, 10-12 mm. long; phyl-
laries 7 or 8, oblong or ovate, 5.5-7 mm. long, 1.5-2 mm. wide, gla-
brous; flowers 10-13, corolla ca. 7 mm. long, the tube (3.5-) 4 mm.
long, the lobes (3-) 3.5 mm. long; anthers 2.5 mm. long; style branches
1.5 mm. long, the tips truncate with a terminal fringe of trichomes;
mature achenes oblong, ca. 3.5 mm. long, 1 mm. wide, glabrous,
brown; pappus 4—6 mm. long, creamy or tawny.

DistTRIBUTION: Puebla and Oaxaca. Known only from three collec-
tions. Flowering in June to August.

This taxon is quite closely related to the O. silphiifolium-goldsmithii
group but is much smaller and does not occur within the present known
geographical range of either of the others. When more material of
this taxon becomes available and can be studied it may indicate that
O. purpusii should not be considered as a distinct species. It is con-
sidered distinct here because of the general small habit, distinct geo-
graphical distribution, and lack of correlative evidence to combine it
with any other taxon.

Oaxaca: Conzatti 4015 (US); Conzattt & Gonzales 1237 (GH, this specimen has
no leaves).

18. Odontotrichum radulifolium [‘‘radulaefolium”] (H.B.K.) Rydb. Bull. Torrey
Bot. Club 51:415. 1924.

Cacalia radulifolia [‘‘radulacfolia’’] H.B.K. Nov. Gen. et Sp. 4:169. 1820.
(Folio ed. 4:132. 1820). Holotype: Guanajuato; between the village of
Temascatio and Guanajuato, no date, 1050 hex., Humboldt & Bonpland
s.n. (P, not seen; Field Mus. Neg. 37875!).

406 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Senecio radulaefolius (H.B.K.) Schultz Bip. Flora 28:498. 1845.

Cacalia schaffnert A. Gray, Proc. Amer. Acad. 19:53, 1883. Lectotype: near
San Luis Potosi in 1876, Schaffner 294 (GH)).

Odontotrichum schaffnert (A. Gray) Rydb. Bull. Torrey Bot. Club 51:415.
1924,

Plants to 1.5 m. tall; entire plant pubescent except the phyllaries,
the hairs white, 5- to 7-celled; stem 3-5 (-S8) mm. thick at the base,
green or purplish; basal leaves 2 or 3, long-petioled, pinnately veined,
ovate-cordate or subcircular in outline, (6-) 10-12 (-18) cm. long,
(S—) 12-14 (22) em. wide, usually 3-5 cm. broader than long or as
long as broad, subpeltate, the petiole attached to the blade 2-10 (—30)
mm. above the base; blades pinnately lobed three-fifths to four-fifths
the distance to the midrib, the primary lobes 7-9 (-11), each divided
into usually 3 (2-5) lobes, the secondary lobes often lobed, 3-angulate
or coarsely dentate, the lowermost pair of primary lobes deflected,
parallel to the petiole; petioles (7—) 12-21 (-30) em. long; cauline
leaves 2-4, the lowermost about as large as the basal ones and similar,
but the base of the petiole more dilated and often leafy-margined ;
upper cauline leaves progressively smaller upward, bracteiform, auric-
ulate-based, elliptic or ovate, distally sinuately 7- to 9-lobed; in-
florescence loosely corymbose, (8-) 12-30 cm. long and broad, the
branches subtended by linear-subulate bracts (5-) 7-10 mm. long;
calyculate bracts 3-5, subulate, 2-5 mm. long; heads (25—) 50-75,
10-11 mm. long; phyllaries 5, oblong or narrowly ovate, 5-7 mm.
long, 1-2 (-3) mm. wide, glabrous; flowers 5 or 6; corolla 6.5—7.5 mm.
long, the tube 3-3.5 (-4) mm. long, the throat very short, ampliate,
0.5-0.7 mm. long, 0.7-1 mm. wide at the apex, the lobes 3-3.5 mm.
long; anthers 2-3 mm. long; style branches 1-1.5 mm. long, hispid on
the abaxial surface, the tips truncate or obtuse with a subterminal
fringe of trichomes; mature achenes ellipsoid, 4 mm. long, 1.5 mm.
wide, brown, ca. 14-ribbed, pubescent, the hairs 0.5 mm. long, greenish
or tawny; pappus creamy or fuscous, 4—5.5 mm. long.

DistriBuTion: Sinaloa, Durango, San Luis Potosf and Guanajuato
to Tamaulipas. Grassy openings in oak-pine woodlands and pastures.
Flowering in July and August.

Comparison of the type specimen of Cacalia schaffneri A. Gray
with the photograph of the type of C. radulifolia H.B.K. and the
similarity of the original descriptions of these two taxa indicate that
they are conspecific, even though the subpeltate condition of the
leaves was not mentioned in the original description of C. radulifolia.
Gray mentioned in his description of C. schaffneri that possibly this
was the little known C. radulifolia of H.B.K. This is the only species
of Odontotrichum with subpeltate leaves.

Durango: Graham 100 (MICH); Feddema 1821 (MICH); Maysilles 7587
(MICH). GuaNnasuato: Kenoyer 2110, 2300 (GH); Schumann 148 (US). San

PIPPEN—MEXICAN “CACALIOID” GENERA 407

Luis Porosf: Manning & Manning 53560 (GH); Mickel 501, 561 (MICH);
Parry & Palmer 543 (NY); Pringle 3566 (GH, US); Pringle 4095 (NY, US);
Schaffner 294 (NY, US, isotype of C. schaffnert A. Gray; the US specimen also
bears the number 724 assigned to the sheet by A. Vigener who purchased many
of the Schaffner collections [J. Rzedowski, pers. comm.]). S1rnaLtoa: Powell &
Edmondson 929 (MICH, TEX). Nuevo Leon: Meyer & Rogers 2608 (MICH)
Tamavuuipas: Stanford, Retherford & Northcraft 818 (GH); Stanford, Taylor, &
Lauber 2421 (MICH, NY, TEX, US).
19. Odontotrichum silphiifolium (Rob. & Greenm.) Rydb. Bull. Torrey Bot.
Club 51:416. 1924.
Cacalia silphiifolia Rob. & Greenm. Amer. Journ. Sci. III. 50:158. 1895.
Holotype: México; Sierra de las Cruces, Aug. 21, 1892, C. G. Pringle
5251 (GH!).

Plants to ca. 1.5 m. tall; roots exuding an amber-colored fluid
when cut; stem 1-1.5 cm. thick at the base, subglabrous or puberu-
lent upward; leaves minutely puberulent especially along the veins;
basal leaves 2 or 3, erect, long-petioled, pinnately veined, ovate-
cordate, (13-) 15-20 (-45) em. long from the tip to the basal sinus,
(11-) 14-21 cm. wide, usually 2-4 cm. longer than wide, occasionally
as wide as long; blades sinuately double-dentate, the teeth in two
alternating series of large and small, respectively, the base of the
blade cordate or sagittate, rarely truncate, the basal lobes 4-7 cm.
long, 5-9 cm. wide, rounded; petioles (15-) 20-30 cm. long; cauline
leaves 2-4, the lowermost similar to the basal ones but smaller,
occasionally deltoid and truncate-based with the petiolar portion
broadly leafy-margined, auriculate-based and clasping the stem;
upper cauline leaves smaller, bracteiform; inflorescence tightly
corymbose, usually 6-11 em. long and broad, occasionally up to 28
cm. long, ca. 20 cm. broad, the branches tomentose, subtended by
linear or narrowly ovate bracts 1.5 cm. long, 1 mm. wide; pedicels
0.3-3 em. long; calyculate bracts 1-5, linear-subulate, (3.5-) 5-7
mm. long; heads 50-75; 10-15 mm. long; phyllaries 8, narrowly
ovate or ovate, (6-) 7-8 mm. long, 1.5-2 (-3) mm. wide, glabrous;
flowers 9-13; corolla 8.5-9 (-11) mm. long, the tube 5 (-6) mm.
long, the lobes 3.5-4 (-5) mm. long, the throat none; anthers 2-3
mm. long; style branches 1—-1.5 mm. long, the tips truncate with a
sparse terminal fringe of trichomes; mature achenes ellipsoid, 4-5.5
mm. long, 1.5-2 mm. wide, 1-1.5 mm. thick, glabrous, light brown,
inconspicuously 16- to 18-ribbed; pappus creamy, (5—-) 7-8 mm. long.

Distripution: Michoacan, east to México and Hidalgo, south to
Morelos. Grasslands, pastures, borders, and openings in oak wood-
lands. Flowering in August and September.

This species is very closely related to O. goldsmithit and may be
distinguished from it by its coarser habit, larger, puberulent, cordate
leaves and more compact, tight inflorescence with larger heads.

408 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Distrito FepEraL: Matuda 19184 (US). Hrpaueo: Rose et al. 9187 (GI, NY,
US). México: Matuda 19417 (NY); Pringle 6453 (GH, MICH, NY, US).
Micuoacin: King 3615 (MICH, NY, TEX, US); Pippen 54 (MICH). Moretos:
Moore 3421 (GH); Orcwlt 3746 (GH, US); Rose & Hay 5311 (GH, US); Pringle
9080 (GH, US).

20. Odontotrichum sinuatum (Cerv.) Rydb. Bull. Torrey Bot. Club 51:415. 1924.
Cacalia sinuata Cerv. in Llave & Lex. Nov. Veg. Desc. fac. 1:29. 1824.
Holotype: Distrito Federal; mountains of the desert of San Angel, near
Mexico City, Cervantes s.n. (not seen).
Senecio albo-lutescens Schultz Bip. Flora 28:498. 1845. Based on Cacalia
sinuata Cerv. not S. sinwatus H.B.K.
Senecio calophyllus Hemsl. Biol. Centr. Amer. Bot. 2:237. 1881. Name
illegitimate.

Slender plants to 1.5 (-2) m. tall; stem 4-6 mm. thick at the base,
light green, pubescent below becoming subglabrous above; leaves
sparsely puberulent, mostly along the veins and margins; basal leaves
3-5 (-7), erect, long-petioled, oblong-cordate in outline, (8-) 12-19
em. long, (8—) 12-17 em. wide, usually 2-3 em. longer than broad or as
long as broad; blades deeply lobed to within 1-3 cm. of the midrib, the
primary lobes 8-11 (-13), 1-3 cm. broad at the base, mostly at right
angles to the midrib, each lobe distally divided into 3 major secondary
lobes and 2 or 3 lesser lobules or coarse teeth, the proximal pair of
primary lobes usually deflected, more or less parallel to the petiole;
petioles 1040 cm. long, usually longer than the blade, slender, gla-
brous, green, often mottled with purple; cauline leaves 3-5 or more,
the lowermost similar to the basal ones and about one-third as large,
the petiole more clasping at the base; upper cauline leaves greatly re-
duced, bracteiform, the petiolar portion leafy-margined and clasping
the stem, distally lobed; inflorescence corymbose (10-) 15-27 em. long,
(6—-) 12-21 em. broad, the branches sparsely to arachnoid-pubescent,
subtended by narrowly ovate bracts 1-1.5 cm. long, 3 mm. wide,
occasionally larger; pedicels 3-5 mm. long; calyculate bracts 3-7, lin-
ear-subulate, 3-7 mm. long; heads 75-100, 10-12 mm. long; phyllaries
mostly 8 (7-9), elliptic or narrowly ovate, 5-6 (-7) mm. long, 2-3 mm.
wide, glabrous, receptacle often slightly pubescent, up to 2 mm. across;
flowers (9-) 10-11 (-13); corolla 6-7 mm. long, the tube 3-3.5 (-4)
mm. long, the lobes 3-3.5 mm. long with a strong median nerve;
anthers 2-3 mm. long; style branches 1-1.5 mm. long, the tips truncate
with a terminal fringe of trichomes; mature achenes ellipsoid, 3.5—4
mm. long, 1.5-2 mm. wide, ca. 1-1.5 mm. thick, glabrous, light brown,
ca. 20-striated; pappus white, 4-5.5 (-7) mm. long; chromosome
number, n=30 (Turner, ined.).

DistrispuTion: From southern Chihuahua south and east to
México. Common in grasslands, pastures, and borders of oak-pine

PIPPEN—MEXICAN “CACALIOID” GENERA 409

woodlands, occasionally in poorly drained areas. Flowering from
August to October.

This is one of the most common species in western and central
Mexico and exhibits considerable morphological variation. The basal
leaves on the same plant as well as within the same population may
vary in size, shape, and depth of lobing.

The holotype of this species has not been located. McVaugh and
Rzedowski searched in Mexico City and McVaugh in Madrid but no
Cervantes specimen was located. It is possible that no type specimen
exists, as few other La Llave and Lexanza specimens are known. The
original description, however, is sufficiently diagnostic to accept this
name.

AGUASCALIENTES: McVaugh & Koelz 49 (MICH); Pippen 1, 7 (MICH). D1s-
TRITO FEDERAL: Bourgeau 716 (GH, US); Smyth 172 (US). Durango: Gould 8989
(MICH); King 3734 (NY, TEX); Mc Vaugh 21733 (MICH, part of hybird popula-
tion between O. sinuatum and O. globosum); Maysilles 7572, 7793, 7880, 8448,
(MICH), 8485 (MICH, NY); Palmer 651 (GH, NY, US); Powell & Edmondson
948 (TEX); Gentry 8562 (MICH). Guanasuato: Duges 451 (GH). Jauisco:
Hartweg 125(GH, NY); Mc Vaugh 12810 (MICH); McVaugh & Koelz 276 (MICH);
Pippen 9*, 22, 57 (MICH). México: Matuda 19706, 21808, 26882 (NY); Pippen
56 (MICH); Pringle 4272, 9875 (GH, NY, US); Rose & Painter 7808 (GH, US).
MicHoackn: Arséne 5740 (US); Feddema 10 (MICH); Pippen 45, 50 (MICH).
Pursua: Arsene 93A, 334, 93, 1181 (US). Wirnout Locauity: Vischer 58 (NY).

21. Odontotrichum tussilaginoides (H.B.K.) Rydb. Bull. Torrey Bot. Club
51:416. 1924.

Cacalia tussilaginoides H.B.K. Nov. Gen. et Sp. 4:168. 1820. (Folio ed.
4:132. 1820.) Holotype: Hidalgo; Real del Monte, 1420 hex., no date,
Humboldt & Bonpland s.n. (P, not seen; Field Mus. Neg. 37878!).

Senecio tussilaginoides (H.B.K.) Schultz Bip. Flora 28:498. 1845. Not S.
tussilaginoides Walt. 1788.

Senecio farfarus Hemsl. Biol. Centr. Amer. Bot. 2:239. 1881. Based on
Cacalia tussilaginoides H.B.K., not S. tussilaginoides Walt. 1788.

Cacalia ampullacea Greenm. Proc. Amer. Acad. 34:577. 1899. Holotype:
Hidalgo; Sierra de Pachuca, 9,000 ft., July 17, 1898, C. G. Pringle 1617
(GH).

Odontotrichum ampullaceum (Greenm.) Rydb. Bull. Torrey Bot. Club 51:416.
1924.

Roots and lower portion of the stem not seen; erect herbs up to
1.3 m. tall (ace. Balls 5428); stem pubescent; leaves puberulent on
the adaxial surface, the abaxial surface gray-tomentulose, the tomen-
tum often flocculent; basal leaves presumably 2 or more, pinnately
veined, ovate-cordate in outline, 20-30 (-35) cm. long from the apex
to the basal sinus, 17-35 cm. wide, usually 2-4 em. longer than broad
or occasionally as broad as long; blades lobed one-half the distance
to the midrib (at least on the distal portion of the blade), the pri-
mary lobes 10-12, 5-6 (-14) cm. long, 4-5 (-11) em. broad at the

410 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

base, each sinuately and irregularly divided into at least, 3 lobes, the
secondary lobes usually coarsely dentate; petioles at least as long as
the blades, usually longer (to 0.5 m.); lower cauline leaves not seen;
upper cauline leaves leafy, bracteiform, ovate, naviculate, clasping
the stem, 11-20 em. long, 10-12 em. wide, the margin toothed or
sinuately lobed; inflorescence tightly corymbose, 17 (—21) cm. long,
12-17 cm. broad, the branches tomentose, the lowermost branches
in the axils of the uppermost leaves, the distal branches subtended
by linear-subulate bracts 1.5-4 cm. long, 1-2 (4) mm. wide; calyeu-
late bracts 4 or 5, linear-subulate, 3-5 mm. long; heads 100 or more,
7-10 mm. long; phyllaries 5, narrowly ovate, 5-7 mm. long, 1.5—2
(—3) wide, glabrous, often purplish; flowers 5 or 6, corolla 6-7 mm.
long, the tube 3-3.5 mm. long, the lobes 3-3.5 mm. long, the throat
none; anthers 2 (-2.5) mm. long; style branches 1 mm. long, hispid
on the abaxial surface, the tips truncate or obtuse with a subterminal
fringe of trichomes; mature achenes not seen, immature achenes
3.5-4 mm. long, pale tan, densely pubescent; pappus creamy or
tawny, 4-5 (—7) mm. long.

DistriputTion: Known only from Hidalgo and Veracruz. In wet
mountain meadows. Flowering in August (?) (Kunth stated in the
description that the plants flowered in May. The specimens I have
examined flowered in August). This is one of the few species of Odon-
totrichum found in eastern Mexico.

This species is somewhat similar to 0. amplifolium, differing mostly
in the smaller heads with fewer flowers and the pubescent achenes.
Comparison of the description and type specimen of Cacalia ampul-
lacea Greenm. with the description and type photograph of O. tus-
silaginoides indicates that they are conspecific because of their
similarity in appearance and their many common characters. Green-
man apparently reached this conclusion after the publication of C.
ampullacea because the type specimen is annotated as C. tussilagi-
nodes H.B.K., in what appears to be Greenman’s writing.

Hipauteo: Rose & Hay 5574 (US); Rose & Painter 6738 (GH, US); Pringle
9868 (GH, NY, US). Veracruz: Balls 5428 (US).

3. Pericalia
Pertealia (Cass. Dict. Sci. Nat. 48: 459. 1827, not validly published) Rydb. Bull.
Torrey Bot. Club 51: 376. 1924.

Perennial, leafy-stemmed herbs, 1.5—-2 m. tall; roots fibrous, fleshy;
stems usually solitary, erect, annual, terete to often 6- to 8-angulate
distally, pubescent or glabrous, tuberculate at base, the clustered
subterranean tubercles each up to 2.5 em. thick, 5 em. long, pubescent
with minute, multicellular, tan hairs; leaves cauline, not forming a
basal rosette, succulent when fresh, coriaceous when dry, petiolate or

PIPPEN—MEXICAN “CACALIOID” GENERA 411

sessile, alternate, clustered toward the middle of the stem but con-
tinuing upward to the inflorescence, becoming smaller and widely
separated distally, not usually becoming bracteiform; inflorescence
paniculate or corymbose; heads usually few, nodding, 15-20 mm.
long, subtended by 10-15, linear or narrowly triangular, calyculate
bracts as long as or longer than the phyllaries and loosely surrounding
them, especially in bud; involucre campanulate to turbinate, the
phyllaries subequal, biseriate appearing uniseriate, imbricate, the
overlapped margins subscarious, the exposed surfaces glabrous, the
tips fringed with minute trichomes up to 0.3 mm. long; receptacle
flat or nearly so, alveolate, not paleaceous; flowers all discoid, perfect,
fertile, 25-50 or more per head; corollas glabrous, creamy or nearly
white but never yellow, the tube narrowly cylindrical, ca. 0.5 mm. in
diameter, the limb divided into 5, equal, narrowly triangular, 3-
nerved, recurved lobes connate into a campanulate throat, 2-5 mm.
long, as long as or twice as long as the lobes; anthers 3-4 mm. long,
exserted beyond the corolla throat, the bases rounded or slightly
sagittate, the terminal appendages trullate, rounded or ovate, 0.5-0.7
mm. long; style branches 1-2 mm. long, slightly flattened, recurved,
the stigmatic area covering most of the adaxial surface, the abaxial
surface glabrous or hispidulous below, the unappendaged tips truncate,
to obtuse, fringed with nonglandular ‘‘pollen-producing’’ trichomes;
achenes narrowly cylindrical, subterete in cross section, glabrous,
purplish brown, 10-ribbed; pappus creamy or white, distally clavate.

Tyre specigs: Pericalia sessilifolia (Hook & Arn.) Rydb.

Cassini suggested that Cacalia cordifolia H.B.K. should be con-
sidered as either a separate genus or subgenus under the name Peri-
calia but he did not validly publish the name as either. DeCandolle
cited Pericalia as a synonym of C. cordifolia H.B.K. and Rydberg
used Pericalia as a genus in 1924.

Microscopic examination of freehand sections of the unique tubercles
revealed their stemlike nature by the presence of discrete collateral
vascular bundles. The parenchyma cells contain needle-shaped crys-
tals. The tubercles function not only in storage but also in vegetative
reproduction, since in some circumstances they give rise to new
flowering stems.

Key to Species of Pericalia

1. Flowers 40—50 per head; lobes of the corolla as long as the throat, 2-3 mm. long;
leaves broadly ovate or elliptic, the margin coarsely toothed or lobulate, the
lobules broader than long .........2.. 2468+. 2. P. sessilifolia

1. Flowers 25-35 per head; lobes of the corolla less than half as long as the throat,
1-1.5 mm. long, the throat 4-5 mm. long; leaves deltoid or subcireular, 3- to
7-lobed, the lobes longer than broad. ......... 1. P. michoacana

266-836—68-——4

412 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

1. Pericalia michoacana (B. L. Rob.) Rydb. Bull. Torrey Bot. Club 51:377.
1924. PLATE 5
Cacalia michoacana B. L. Rob. Proc. Amer. Acad. 43:46. 1907. Holotype:
Michoacin; Uruapan, on pine covered crater cone, 1,680 m., Oct. 31,
1905, C. G. Pringle 10117 (GIT).
Cacalia trigonophylla 8. F. Blake, Journ. Wash. Acad. 19:280. 1929. Holotype:
Jalisco; San Sebastian, trail to mine La Sabala, Feb. 10, 1927, Ynex Mexia
1656 (USI).

Slender herbs 0.6-1.5 (-2) m. tall; underground tubercles 1.5-2.5
em. thick, to 5 em. long; stem 2-5 (-7) mm. thick at the base, greenish,
often mottled with purple, glabrous to sparsely or densely pubescent
proximally, becoming glabrous and often glaucous above; leaves
10-15 or more, long-petioled, subpalmately veined with 3-5 principal
veins with the central vein stronger, densely to sparsely pubescent
to glabrous; blades variable in shape from subcircular-cordate-based
with 5-7 deltoid or triangular lobes, to broadly deltoid-hastate with
3 major, narrowly triangular lobes, 4-7 (-10) cm. long from the
basal sinus to the tip, 5.5-10 (-16) em. wide, usually 2-3 cm. wider
than long, progressively smaller upward, the margins minutely or
coarsely dentate; petioles 5-10 (-15) mm. long, purplish; inflorescence
racemose or paniculate, 10-20 cm. long, 8-12 em. broad or larger, the
distal branches subtended by linear-subulate bracts 1-1.5 cm. long,
the proximal branches often in the axils of the uppermost leaves;
pedicels 2-5 (-7) em. long; calyculate bracts 10-15, linear or narrowly
triangular, 4-5 (-7) mm. long; heads mostly 6-12 (-50), nodding, 15-
20 mm. long; phyllaries (10—) 12 or 13, narrowly ovate or narrowly
oblong, (7-) 10-13 mm. long, 1-2 mm. wide; receptacle 4-5 mm.
across; flowers 25-35; corolla 9-12.5 mm. long, the tube (3.5-) 4-4.5
(-5.5) mm. long, the throat funnelform or campanulate, 4-5.5 mm.
long, 1.5 mm. wide at the apex, the lobes 1-1.5 mm. long, triangular,
strongly recurved; anthers 3.5-4 mm. long; style branches ca. 1.5
mm. long, the tips truncate or rounded with a sparse subterminal
fringe of trichomes; achenes narrowly oblong or narrowly obovoid,
terete in cross section, 2.5 mm. long, 0.5 mm. thick, glabrous, dark
brown with 10 pale ribs; pappus (8—) 9-10 (-11) mm. long, white,
slightly clavate distally; chromosome number, n=30 (Pippen).

DistripuTion: From northwest Jalisco eastward to México.
Growing mostly on rocky slopes in oak-pine woods. Flowering in
November and December.

The specimens examined show a wide range of variation in leaf
shape and pubescence between the two extreme forms described
above. Somecorrelation between leaf shape and pubescence is apparent
since the plants with 5- to 7-lobed leaves tend to be more pubescent
and those plants with 3-lobed leaves tend to be glabrous and glaucous.
A specimen from eastern Jalisco (McVaugh & Koelz 435) has more

PIPPEN—MEXICAN “CACALIOID” GENERA 413

rounded lobes than the other specimens, Despite these variations all
of these specimens form only one taxon.

Comparison of the types of Pericalia michoacana and Cacalia tri-
gonophylla and examination of a large sample of these taxa indicate
that they are conspecific. They have the same number of phyllaries
and flowers in a head and the morphological structure of the flowers
is the same. The only observable difference is in the pubescence and
the number of lobes. Blake based C. trigonophylla on a collection in
which the leaves were 3-lobed and subglabrous. No geographical or
ecological differences are noticed between these two extremes in
vegetative form and therefore no nomenclatural status is given to

these variations.

Jatisco: Mezia 1656 (US, MICH, isotypes of C. trigonophylla Blake); Mc-
Vaugh 14290, 21355, 14136, 14093, 22071 (MICH); Pippen 26, 38, 63*, 65*
(MICH); McVaugh & Koelz 435 (MICH). México: Hinton et al. 2755 (US),
4993 (NY, US); Moore & Cetto 5469 (MICH). Micnuoacdn: Hinton et al 12718,
13469 (MICH, NY); Pringle 10117 (US, isotype).

2. Pericalia sessilifolia (Hook. & Arn.) Rydb. Bull. Torrey Bot. Club 51:376.

1924.

Cacalia cordifolia H.B.K. Nov. Gen. et Sp. 4:168. t. 360. 1820. Holotype:
México; Santa Rosa, Humboldt & Bonpland s.n. (P, not seen; Field Mus.
Neg. 37865!), not Cacalia cordifolia L.f.

Cacalia sessilifolia Hook. & Arn. Bot. Beech. Voy. 436. 1841. Holotype:
Nayarit; between San Blas and Tepic, in 1837, Sinclair s.n. in Hooker
Herb. (K, not seen; MICH Neg. 621)).

Senecio ovatifolius Schultz Bip. Flora 28:498. 1845. Based on Cacalia cordi-

folia H.B.K. not S. cordifolius L.f.
Senecio beecheyanus Schultz Bip. Flora 28:499. 1845. Based on Cacalia

sessilifolia Hook. & Arn. Name illegitimate.

Senecio cardiophyllus Hemsl. Biol. Centr. Amer. Bot, 2:237. 1881. Based
on Cacalia cordifolia H.B.K. Name illegitimate.

Senecio sessilifolius (Hook. & Arn.) Hemsl. Biol. Centr. Amer. Bot. 2:247.
1881. Based on Cacalia sessilifolia Hook. & Arn.

Cacalia nutans Sessé & Moc. Pl. N. Hisp. ed. 1. 182. (Fl. Mex. ic. ined.
145; Field Mus. Neg. 30705!) 1889. Lectotype: Without locality (pre-
sumably “in humidis unbrosisque Tepalpae circuitibus prope Mexicanum”’),
Sessé & Moc. 1488 or 2822! (both numbers appear on the specimen),
Field Mus. Neg. 42285.

Pericalia ovatifolia (Schultz Bip.) Rydb. Bull. Torrey Bot. Club 51:377. 1924,

Plants 0.5-1.5 (-1.8 m., Balls 5575) m. tall; tubercles 2-2.5 cm.
thick, to 5 cm. long; entire plant glabrous and glaucous, often sparsely
pubescent when young; stems 3-9 mm. thick at the base; leaves 10-15,
petiolate or sessile, subpalmately veined with often 3 principal veins,
the central one usually dominant; blades cordate-based, broadly ovate
or broadly elliptic, the larger ones 4-13 cm. long from the basal sinus
to the apex, 5-12 (-14) cm. wide, usually 1-3 em. wider than long
(often as long as wide to 1% times longer than wide), the margin

414 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

variable from subentire to dentate to 9- to 14-lobulate, the lobules,
when present, usually broader than long; petioles (when present) sub-
terete, purplish green, 3-9 cm. long; inflorescence corymbose or pa-
niculate, 10-30 em. long, 7-18 cm. broad, the proximal branches often
in the axils of the distal leaves, the other branches subtended by sub-
ulate or narrowly ovate bracts 7-10 mm. long, 1 mm. wide; pedicels
(2-) 3-6 (-9) cm. long; calyculate bracts 10-16, linear or narrowly
triangular, 6-11 mm. long; heads 6-15 (—20), 15-20 mm. long; phyl-
laries 12-14 (-15), narrowly ovate or narrowly triangular, (9-) 11-14
mm. long, 2-3 mm. wide, greenish or purplish; receptacle ca. 7 mm.
across; flowers 40-50 or more; corolla (7—) 8-12 (-13) mm. long, the
tube (3.5) 4-5 (-7) mm. long, the throat 2-3 mm. long, 1-1.5 mm.
across at the top, the lobes about as long as the throat, 1.5-3 mm.
long, strongly recurved; anthers 3-3.5 (-4) mm. long; style branches
1-1.5 (-2) mm. long, recurved; mature achenes oblong or ellipsoid,
subterete in cross section, 3.5 mm. long, 1-1.5 mm. wide, ca. 0.5—1
mm. thick, glabrous, purplish brown, 10-ribbed; pappus white (7-)
8-10 mm. long, distally clavate; chromosome number, n=30 (Pippen).

DistrispuTtion: Durango, Nayarit, and Jalisco, east to Michoacin
and México. Common on rocky slopes in the oak-pine woodlands.
Flowering in September and October.

Comparison of Cacalia cordifolia H.B.KK. and P. sessilifolia (Hook.
& Arn.) Rydb. indicate that they are conspecific. They differ only in
the presence or absence of the petiole. Several specimens from Nayarit
have both sessile and petiolate leaves on the same plant. The heads,
flowers, and other morphological characters are similar. The sessile-
leaved plants are apparently vegetative variations that occur several
times within the range of the petiolate forms. This difference is not
significant enough for nomenclatural status.

This species is a victim of nomenclatural rules. Since Cacalia cordifolia
H.B.K. was superfluous when published because of the earlier C.
cordifoha L. f., a different taxon, the next oldest valid name for this
taxon is Cacalia sessilifolia [Pericalia sessilifolia (Hook. & Arn.)
Rydb.]. This is unfortunate since the petiolate-leaved plants are by
far more common than the sessile-leaved forms.

AUGUASCALIENTES: McVaugh & Koelz 147 (MICH); Pippen 5 (MICH).
Distrito FEDERAL: Balls 5575 (US); Bourgeau 715 (GH); Pringle 7983 (GH);
Lyonnet 194 (GH, NY, US); Purpus 5625 (GH, NY, US). Duranao: Gentry 6968
(GH, MICH, US); McVaugh 21658 (MICH); Maysilles 7579 (MICH); Pennell
18345 (NY, all of the specimens from Durango are of slightly smaller stature with
more, smaller leaves). GuaNasuatTo: Duges 481 (GH, US). Jaisco: McVaugh
13559, 13625, 17118, 17183, 18121 (MICH); McVaugh & Koelz 920 (MICH);
Mexia 1450 (US); Palmer 576 (GH, NY, US); Pippen 17, 18, 31, 33, 34, 61*, 64
(MICH); Pringle 1736 (GH, MICH, NY, US), 9869 (GH, NY, US), 11497 (GH,
MICH, US); Safford 1451 (US); Rose 2828 (GH, US). M&xtco: Matuda 21825

PIPPEN—MEXICAN “CACALIOID” GENERA 415

(NY). Micnoacin: Arsene s.n., Sept. 11, 1909, Sept. 14, 1911, 5729, 7347 (US);
Feddema 161 (MICH); Pippen 51 (MICH). Nayarit: Feddema 381, 1345 (MICH):
McVaugh & Koelz 479, 792, 656 (MICH); Mezia 635 (US); Ortega 5727 (US);
Palmer 1832 (GH, NY, US); Pippen 13, 15 (MICH). QuerETARO: Matuda 29787
(NY). San Luts Potosf: Palmer 47 (GH, NY, US); Parry & Palmer 541 (GH,
US); Schaffner 255, 33 or 37? (GH).

Three specimens in the herbarium at the University of Michigan
do not fit precisely into either of the above taxa, P. sessilifolia or P.
michoacana. Two of these, McVaugh 13469 and 13825 (from northeast
slopes of the Nevado de Colima, below Canoa de Leoncito, 2,250-
2,550 m.), have leaves similar to P. michoacana in that they are sub-
circular, 7-lobed, and pubescent, but they are about 0.5 times larger
with much longer petioles and the inflorescence has larger heads with
more flowers. The phyllaries are finely pubescent with fuscous hairs
and the heads are subtended by about 15 calyculate bracts as wide
as but longer than the phyllaries.

These specimens may represent a different species but, because of
the paucity of material available and its rather close similarity to P.
michoacana, I will only mention their existence for the present.

The third nonconforming specimen, McVaugh 13862 (Sierra de
Manantlin, 15-20 miles southeast of Autlin, near Asseradero El
Cuartén, 2,500 m.), differs from the above in that the leaves are less
deeply lobed, less pubescent, and the trichomes are mostly restricted
to the leaf margins; the phyllaries are glabrous, the subinvolucral
bracts no longer than the phyllaries and somewhat narrower. This
specimen approaches P. sessilifolia but differs in the longer petioles,
the more subcircular than ovate, 7-lobed leaves, and the presence of
pubescence. Future collections and field observations in this region
will be necessary to properly interpret this specimen.

4. Psacalium
Psacalium Cass. Dist. Sci. Nat. 43:461. 1826.

Perennial, subscapose herbs, 0.1-3 m. tall; roots fleshy, fibrous;
caudex tough, to 5 em. long, 3 cm. thick, the nodes and terminal
buds densely covered with masses of tawny, multicellular hairs, 1-2
cm. long; stems usually solitary, annual, erect, slender or stout,
usually terete, glabrous or pubescent; radical leaves forming a basal
rosette of two to several, centrally peltate with several principal
radiating veins, each pinnately subdivided, the blades variously lobed,
rarely subentire, mostly pubescent; cauline leaves alternate, pro-
gressively smaller upward, the uppermost bracteiform, usually non-
peltate with a broad leafy-margined petiole and palmately 3- to
7-lobed blade; inflorescence paniculate or corymbose, the branches
often stipitate glandular, with minute, multicellular glands, 0.5 mm.

416 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

long, subtended by small, linear-subulate or large, leafy, often vagi-
nate bracts; heads few to many, calyculate, the bracts 3-5, as long
as the phyllaries or shorter (longer in P. peltatum); involucre cam-

panulate, turbinate or less frequently cylindrical, the phyllaries
5-17, subequal, biseriate appearing uniseriate, the overlapped mar-
gins subscarious, the exposed surfaces pubescent or glabrous, the
tips fringed with minute trichomes up to 0.3 mm. long; receptacle
flat or nearly so, alveolate, not paleaceous, occasionally sparsely
pubescent; flowers all discoid, fertile, 5-80 per head; corollas glabrous,
creamy or nearly white, occasionally greenish or purplish tinted but
never yellow, the tube narrowly cylindrical, 0.5 mm. in diameter,
the limb divided into 5, equal, narrowly-triangular, 3-nerved, ascend-
ing or spreading, not recurved lobes, connate into a short, ampliate
or campanulate throat, about 1 mm. long, 1-1.5 mm. wide (or occa-
sionally lacking), the lobes longer than the throat and as long as to
one-half as long as the tube; anthers 2-4 mm. long, exserted beyond
the corolla throat, rounded or slightly sagittate at the base, the
terminal appendages trullate or ovate 0.5—-0.7 mm. long; style branches
0.7-2 mm. long, slightly flattened, recurved or spreading, the stig-
matic surface covering most of the adaxial surface, the abaxial surface
glabrous or hispidulous below, the unappendaged tips truncate to
obtuse with a fringe of nonglandular “pollen producing” trichomes;
achenes obovoid or ellipsoid, somewhat compressed and elliptic or
broadly elliptic in cross section, glabrous or pubescent, striate or
prominantly 10- to 12-ribbed; pappus a single row of white or creamy,
stiff, barbellate, capillary bristles (lacking in P. nanum and P. calvum);
chromosome number, n=30.

TyprE specizs: Psacalium peltatum (H.B.K.) Rydb.

This genus is easily recognized by the peltate basal leaves and is
the only genus discussed here with stipitate glandular pubescence in
addition to the usual nonglandular trichomes.

Key to Species of Psacalium

1. Inflorescence and upper portion of stem minutely stipitate glandular; plants
also pubescent with nonglandular hairs.
2. Achenes pubescent.

3. Corolla tube about twice as long as the lobes, 6-7 mm. long; heads nodding,

about 40 or fewer; pappus 9-10 mm. long ... . 3. P. holwayanum
3. Corolla tube about the same length as the lobes, 3.5-4 (-5) mm. long;
heads erect, 50 or more; pappus 6-7 mm. long . . . 2. P. eriocarpum

2. Achenes glabrous.
4. Flowers 5-6 per head; Piylaries & 5, mostly glabrous or very sparsely
pubescent. . .... . .. . I1. P. tabulare

4. Flowers 8 or more per head; phyll: wries ‘at least as numerous as the flowers,
densely pubescent.

PIPPEN—MEXICAN “CACALIOID” GENERA 417

5. Basal leaves lobed one-half or less the distance to the center; phyllaries
(7-) 8; flowers 8-10; bracts subtending the phyllaries either 3-5 and
one-half as long as the phyllaries, or 1 or 2 and as long as the phyllaries.

6. Lobes of the leaves 4 or 5; leaves 5-6 cm. in diameter, pubescent on the
adaxial surface; phylaries subtended by 1 or 2 bracts as long as or
longer than thep hyllaries. . ......, . . . 4. P. laxiflorum.

6. Lobes of the leaves 6 or 7; leaves 20-30 cm. in diameter, glabrous on
the adaxial surface; phyllaries subtended by 3-5 bracts about one-
half as long as the phyllaries. ... . . . 12. P. species

5. Basal leaves lobed about two-thirds the distance to the center; phyllaries
(8-) 12-15; flowers 20 or more; bracts subtending the phyllaries 3 or 4,
longer than the phyllaries . . 2... 1. we ee . 8 P. peltatum

1. Inflorescence and upper portion of the stem. lacking stipitate glands; plants
pubescent with nonglandular trichomes or glabrous.
7. Plants with fleshy underground tubers attached to the caudex.
8. Pappus present, 4-6 mm. long; flowers 5-8; phyllaries 5; plants 0.5 m. tall

ortaller . 2... 6 ew ee ee et ee ee ee 9. P. peltigerum
8. Pappus lacking; flowers 10 or ll: | phyla 7; plants up to about 15 cm,
tal. . 2... rr 6. P. nanum

7. Plants lacking underground tubers.

9. Pappus lacking; achenes obovoid, 10-12 mm. long, 5-6 mm. wide;
leaves deeply and finely dissected almost to the center, the major
segments again divided, the ultimate segments linear . 1. P. calyum

9. Pappus present; achenes ellipsoid, up to 7 mm. long, 2 mm. wide; leaves
lobed as deep as three-fourths the distance to the center, not deeply
and finely dissected.

10. Phyllaries densely pubescent.
11, Bracts subtending the phyllaries 3 or 4, longer than the phyllaries;
phyllaries 8-10; flowers 12 or 13; receptacle naked, not pubescent.
8. P. peltatum
11. Bracts subtending the phyllaries 3-5, less than half as long as the
phyllaries; phyllaries 5; flowers 5 or 6; receptacle sparsely pubescent.
5. P. megaphyllum
10. Phyllaries glabrous or very sparsely pubescent.
12. Phyllaries 9-10 mm. long; achenes glabrous; leaves pubescent,
densely so on the abaxial surface.
13. Phyllaries 5; flowers 5 or 6; ranging from Veracruz west to
Michoacdn. .... ..... Il. P. tabulare
13. Phyllaries 8 or 9; flowers 10-12; known only from central Oaxaca.
7. P. nelsonii
12. Phyllaries 5-7 mm. long; achenes pubescent; leaves glabrous or
sparsely pubescent along the veins on the abaxial surface.
10. P. poculiferum
1. Psacalium calvum (Brand.) Pippen, comb. nov.

Cacalia calva Brand. Univ. Calif. Publ. Bot. 4:193. 1911. Holotype: Puebla;
Cerro de Gavilan, August 1900, C. A. Purpus 4113 (UC, not seen; isotypes
GH!, US!).

Odontotrichum caluum (Brand.) Rydb. Bull. Torrey Bot. Club 51:420. 1924.

Lower portion of the plant not seen; plants ca. 0.5 m. tall; stem
and leaves densely pubescent to tomentose; flowering stem terete;

418 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

basal (?) leaves 2 or 3, long-petioled, peltate, subcircular in outline,
12-14 em. in diameter; blades deeply dissected into 6-8 divisions
extending almost to the center, the major divisions several times
subdivided, the ultimate divisions narrow, linear; petioles slender,
20-25 cm. long; cauline leaves reduced and bracteiform; inflorescence
corymbose, the branches subtended by linear-subulate bracts to 1
cm. long; heads about 20; phyllaries 12, narrowly ovate, 8 mm. long,
1 mm. wide; flowers 12-14; corolla about 7-8 mm. long (flower meas-
urements from the original description), the tube 3-4 mm. long, the
throat ca. 0.5 mm. long, 0.5-1 mm. across at the apex, the lobes 4
mm, long; anthers about 3 mm. long; style branches recurved, hirtel-
lous; mature achenes obvoid, 10-12 mm. long, 5-6 mm. wide, glabrous,
pale brown; pappus lacking entirely.

DistrisutTion: Known only from the type specimens.

The specimens examined lacked adequate flowering material, and
the poorly preserved leaves made it difficult to determine whether or
not they were peltate.

The most unusual features of this species are the extremely large,
obovoid achenes and lack of a pappus. Field examinations and study
of material collected in the future may indicate a change in the
taxonomic position of this taxon.

2. Psacalium eriocarpum (8. F. Blake) Blake Journ. Wash. Acad. Sci. 28:492. 1938.
Cacalia eriocarpa 8. F. Blake Journ. Wash. Acad. Sci. 129. 1929. Holotype:
Jalisco; Arroyo de Santa Gertrudis, San Sebastian, steep slopes in open

pine and oak woods, 1,500 m., Jan. 21, 1927, Ynez Mexia 1539 (US)).

Slender plants up to 1.3 m. tall, stems 5-10 mm. thick at the base,
purplish green, sparsely or densely pubescent with long, multicellular,
stiff, attenuate-tipped hairs, 1-1.5 mm. long, and with multicellular,
stipitate glands, 0.5 mm. long; leaves hirsute with nonglandular hairs
like those on the stem, the pubescence denser on the adaxial surface;
basal leaves 2 or 3, long-petioled, centrally peltate, subcircular in
outline, 15-25 (-30) em. in diameter, the venation radiate with 10 or
11 principal veins; blades shallowly lobed one-third the distance to the
center of the blade or less, the lobes 8-10, 1.5-2 times broader than
long, each 3-angulate or subdivided into 2 or 3 smaller lobules, the
margins spinulose-toothed or entire, the center of the blade flat, not
depressed; petioles 15-30 em. long, slender; cauline leaves 1 or 2,
greatly reduced, bracteiform, inflorescence paniculate, usually 30 cm.
long or longer, 10-15 cm. broad, the branches subtended by narrowly
triangular bracts 3-7 mm. long, 1 mm. wide; pedicels 3.5-5 (-9) mm.
long, calyculate bracts 3-5, subulate, 2-5 mm. long; heads 50-70
(-90), 1.2-1.5 em. long; phyllaries 5 or 8, narrowly ovate or elliptic,
(6-) 7-8 (-9) mm. long, 1-3 mm. wide, purplish tipped, pubescent,
occasionally sparsely stipitate glandular; flowers 6 (when 5 phyllaries)

PIPPEN—MEXICAN “CACALIOID” GENERA 419

or 9-11 (when 8 phyllaries); corolla about 8-9 mm. long, the tube
(3.5-) 4 (-5) mm. long, the throat short, campanulate, about 1 mm.
long and broad, the lobes about as long as the tube, 3-4 (-5) mm.,
narrowly triangular; anthers 3-4 mm. long; style branches 1.5-2 mm.
long, the abaxial surface hispidulous, especially distally, the tips
rounded or obtuse with a sparse subterminal fringe of hairs; mature
achenes not seen, the immature achenes pale green, densely villous;
pappus (5.5-) 6-7 (-8) mm. long, white; chromosome number, n=30
(Pippen).

DistripuTion: Western Jalisco. Mostly in pine and, less frequently,
oak woodlands. Flowering mid-November to December.

This species has two forms: the more common form has small heads
with 5 phyllaries and 6 flowers, the other has larger heads with 8
phyllaries and 9-11 flowers. These two forms are similar in all other
aspects, including range. The larger headed form is known only from
an area in between the known populations of the smaller headed form,
These forms are not given nomenclatural status. This species is
closely related to P. holwayanum but differs from it in the smaller
heads, erect inflorescence, and less deeply lobed leaves.

Jauisco: McVaugh 13954, 14287, 13684, 14165 (MICH); McVaugh & Koelz
919 (MICH); Pippen 36, 62*, 66, 67 (MICH).

3. Psacalium holwayanum (B. L. Rob.) Rydb. Bull. Torrey Bot. Club 51:3872.
1924.
Cacalia holwayana B. L. Rob. Proc. Amer. Acad. 43:45. 1907. Holotype:
Michoacin; Uruapan, Oct. 11, 1899, #. W. D. Holway 2617 (GH).
Psacalium langlasset Rydb. Bull. Torrey Bot. Club: 51:376. 1924. Holotype:
Guerrero; Sierra Madre, in 1898, #. Langlassé 576 (USI).

Plants up to 1 m. tall; stems 3-5 mm. thick at the base, densely
pubescent with stiff, white hairs up to 2 mm. long, 12- to 20-celled,
and also with 6-celled stipitate glands, 0.1-0.2 mm. long, abundant
distally; leaves sparsely or densely pubescent with stiff, nonglandular
hairs only; basal leaves 2 or 3, long-petioled, peltate, subcircular in
outline, (9—-) 12-14 (-26) cm. in diameter, the venation radiate with
8 principal veins; blades lobed about one-third the distance to the
center, the lobes 7 or 8, 2.5-4 cm. long, 3-6 cm. wide, each one 3-
angulate or divided into 2 or 3 smaller secondary lobes, the margins
irregularly and sparsely toothed; petioles (9-) 10-18 (—32) cm. long,
terete; cauline leaves 2 or 3, similar to the basal ones but smaller, 1-5
cm. in diameter, the uppermost subpeltate with petioles 1-5 (—10)
cm. long; inflorescence thyrsiform, 16-30 cm. long, 14-16 cm. broad,
the branches pubescent and stipitate glandular lke the stem, sub-
tended by either linear bracts 1—2 em. long, 1 mm. wide or occasionally
spathulate bracts, 3-toothed at the tip; pedicels 5-15 mm. long;

420 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

calyculate bracts 3-5, subulate, 4-7 mm. long; heads about 40 or
fewer, nodding, 1.5-3 cm. long; phyllaries 8, narrowly ovate, 9-13
mm. long, 1.5-3 mm. wide, purplish, pubescent with glands and sparse-
ly hirsute with nonglandular, long, stiff hairs; receptacle 3-3.5 mm.
long; flowers 10 or 11; corolla 10-11 mm. long, the tube 6-7 mm. long,
the throat short, campanulate, ca. 1 mm. long, 1.5-2 mm. across,
the lobes 3-3.5 (-5, Pringle 13294) mm. long, linear, 0.5 mm. wide,
anthers 3.5-4 mm. long; style branches 1.5-2 (-3) mm. long, hispid
on the abaxial surface, the tips truncate with a terminal fringe of
trichomes; mature achenes 5-6 mm. long, 1.5 mm. wide, 0.5-0.7 mm.
thick, oblong, or ellipsoid, pale, densely pubescent with unicellular,
adpressed, ascending hairs; pappus 9-10 mm. long, white; chromosome
number, n=30 (Turner, ined.).

DisrriputTion: Known from west central Michoacdén eastward to
México and from onecollection in Guerrero. Mostly in pinewoodlands.
Flowering mid-November to December.

An isoparatype, Langlassé 576 (US), was described by Rydberg as
Psacalium langlassei, because the leaves were less deeply lobed. Com-
parison of this specimen with the type and other specimens of P.
holwayanum indicates that they are conspecific because they are
similar in all characters except leaf lobes. The specimen Rydberg
examined was merely less deeply lobed.

GueErrRERO: Langlassé 576 (GH, paratype of C. holwayana, isotype of P.
langlassei). M&xico: Gilly 5 (MICH); Matuda 30015 (NY). Mucnoackn:
King & Soderstrom 4890 (MICH); Pringle 13297 (GH, US), 13672 (GH, para-
type; US), 138979 (GH, MICH, US); McVaugh 21953* (MICH).

4. Pascalium laxiflorum Benth. Pl. Hartw. 41. 1841. Holotype: Michoacan;
Morelia, pine woods, in 1836, Hartweg 318 (K, not seen; Field Mus. Neg.
37949!).

Senecio moreliae Hemsl. Biol. Centr. Amer. Bot. 2:243. 1881. Based on
Psacalium laxiflorum Benth. not S. laxiflorum Viv.

Slender plants 6-7 cm. tall; stems 3-4 mm. thick at the base,
pubescent with multicellular stiff hairs, 1-1.5 mm. long, and minute,
2- to 3-celled stipitate glands ca. 0.1 mm. long, most abundant dis-
tally; leaves pubescent on the adaxial surface with nonglandular
hairs, subglabrous abaxially; basal leaves 2 or 3, petiolate, peltate,
subcircular in outline, 5-6 cm. in diameter, the venation radiate with
5 principal veins; blades lobed about one-half the distance to the
center, the lobes 4 or 5, 1-2 em. long and broad or occasionally 1.5-2
times broader than long, each slightly 3-angulate, the tips rounded
or obtuse, the margins mucronulate-toothed, the sinuses broadly
rounded; petioles 6-8 cm. long, terete; cauline leaves usually 2, the
lowermost similar to the basal ones, the upper one nonpeltate, 2.5—-4
em. long, 1-2 cm. wide with the petiolar portion leafy-margined,

PIPPEN—MEXICAN “CACALIOID” GENERA 421

ovate, the blade portion 3-lobed; inflorescence paniculate, 16-26 cm.
long, 11-12 cm. broad, the branches pubescent and stipitate glandular
like the stem, subtended by narrowly ovate or ovate bracts 2-2.5 cm.
long, 0.5-1 cm. wide progressively smaller upward, the bracts glan-
dular on the adaxial surface, glabrous abaxially; pedicels 5-10 mm.
long; calyculate bracts 1-2, subulate, 8-10 mm. long; heads 30 or
fewer, 14-15 mm. long; phyllaries 8, narrowly ovate, (6-) 7-8 mm.
long, 1.5-3 mm. wide, purplish, densely stipitate glandular; flowers 8;
corolla 8-9 mm. long, the tube 5-5.5 mm. long, the throat ampliate,
0.5 mm. long, 0.7-1 mm. wide, the lobes 3-3.5 mm. long, linear;
anthers 3 mm. long; style branches 1.5 mm. long, hispid on the
abaxial surface, the tips truncate or rounded with a terminal or
subterminal fringe of trichomes; mature achenes not seen, immature
achenes glabrous, striate; pappus 6-7 mm. long, white.

DistrisuTion: Known only from the type collection and one col-
lection from southern Durango (Rose 2337, GH, US). In pine-wood-
lands. Flowering in August.

This species has smaller leaves than the other members in the
section, and they tend to be 4-lobed and somewhat cross shaped. It
is related to P. holwayanum and P. eriocarpum.

5. Psacalium megaphyllum (Rob. & Greenm.) Rydb. Bull. Torrey Bot. Club 51:
374. 1924. PuLaTE 6
Cacalia megaphylla Rob. & Greenm. Amer. Journ. Sci. III. 50:157. 1895.
Holotype: Jalisco; on hillside near Guadalajara, Oct. 10, 1889, C. G.
Pringle 2490 (GH).
Cacalia obtusiloba Rob. & Greenm. Amer. Journ. Sci. III. 50:158. 1895.
Holotype: Oaxaca; Sierra de San Felipe, 6,000 ft., Nov. 17, 1894, C. G.

Pringle 5840 (GH).
Psacalium obtusilobum (Rob. & Greenm.) Rydb. Bull. Torrey Bot. Club

51:374. 1924.

Stout plants to 3 m. tall; entire plant pubescent, the pubescence
denser in the inflorescence and on the abaxial surface of the leaves,
the hairs 0.5-0.7 mm. long, 5- to 15-celled, white; stem to 2 cm.
thick at the base; basal leaves 3 or 4, long-petioled, peltate, subcircular
in outline, 30-60 cm. in diameter, the venation radiate with 5-7
principal veins; blades lobed one-half to two-thirds the distance to
the center, the primary lobes 5-7, 7-15 cm. long from the base of
the sinus to the distal edge, 5-7 cm. wide at the base, each usually
divided into 3 lobes, the secondary lobes (5—-) 7-10 cm. long, 3-5 cm.
wide at the base, each divided into several smaller, triangular lobules
1-3 cm. long, the margins coarsely toothed, the center of the blade
depressed; petioles 25-80 cm. long, terete, purplish green; cauline
leaves 5-7 or more, the lowermost usually smaller than the basal
ones but similar to them, 30-35 cm. in diameter, the upper cauline
leaves progressively smaller upward, subpeltate or nonpeltate, the

492 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

uppermost 8-15 cm. long, 10-20 cm. wide, palmately 3- to 5-lobed,
the petioles 3-7 cm. long; inflorescence broadly paniculate-corymbose,
to 1.5 m. high, ca. 1 m. broad, the proximal branches in the axils of
the uppermost leaves, the distal branches subtended by ovate bracts
2-5 cm. long, 3-7 mm. wide; pedicels about as long as the phyllaries;
calyculate bracts 3-5, subulate, 1.5-2 mm. long; heads at least 150,
10-12 mm. long; phyllaries 5, narrowly elliptic or narrowly ovate,
(5.5-) 7-8 (-10) mm, long, (1.5-) 2-3 mm. wide, densely white
puberulent; receptacle sparsely pubescent with filiform, multicellular
hairs; flowers 5 (-6); corolla 6-8 mm. long, the tube 3-4 mm. long,
the throat 0.5 mm. long and broad (or occasionally lacking), the
lobes narrowly triangular, 2.5-4 (-5.5) mm. long, reflexed; anthers
2-3.5 mm. long; style branches 1—-1.5 mm. long, the abaxial surface
hispid, denser distally, the tips obtuse with a sparse terminal fringe
of trichomes; mature achenes oblong or ellipsoid, 5-5.5 mm. long,
1.5-2 mm. wide, 1 mm. thick, pale green, glabrous, 15- to 20-ribbed;
pappus 4-5 (-7) mm. long, white; chromosome number, n=30
(Turner, ined.).

Distripution: From Zacatecas southeast to Oaxaca. Common, at
least in western Mexico, in rocky grasslands, roadsides, and grazed
areas. Flowering in September and October.

This species is striking because of its very large size and especially
its extremely large basal leaves, depressed in the center. Most existing
herbarium specimens do not adequately represent the species because
even one leaf is larger than the average herbarium sheet.

Robinson and Greenman in describing Cacalia megaphylla and C.
obtusiloba stated that the latter differed from the former in having
smaller heads and more rounded basal leaf lobes. Comparison of the
types of these two species, however, indicates that they are conspecific
because of several common characters: Number of phyllaries and
flowers, pubescence of phyllaries, leaves and stem, general habit, and
leaf shape (the latter somewhat variable).

AGUASCALIENTES: McVaugh & Koelz 39 (MICH); Pippen 8* (MICH). Guer-
RERO: McVaugh 21897, 21910 (MICH); Moore 5119 (MICH). Jauisco: McVaugh
13321 (MICH); Pippen 16, 24, 39 (MICH). México: Hinton 2200 (US), 2355
(NY); Matuda 26572 (NY). Micnoacdn: Arséne 2386 (US); Hinton et al. 13480,
15629 (MICH, NY); Pippen 47 (MICH); McVaugh 21928 (MICH). More ros:
Pringle 6165 (GH, NY, US). Nayarit: Meria 877 (GH, MICH, Us). PuEBta:
Arsene & Nicholas 334 (US); Arséne, s.n., November 1908 (US). ZacaTECAs:
McVaugh 17650 (MICH).

6. Psacalium nanum Pippen, sp. nov. PLATES 7, 8
Plantae nanae usque ad 12 cm. altae, tuberibus pluribus, carnosis,
usiformibus, subterraneis, caulibus hirsutis scaposis, foliis radicalibus
fusif bus, subte , libus | t I , fol dicalib
peltatis, suborbicularibus, usque ad 5 cm. diametro, profunde 7-

PIPPEN—MEXICAN “CACALIOID” GENERA 423

lobatis, supra hirsutis, subtus glabris, petiolis tenuibus 5-10 cm.

longis hirsutis; inflorescentia corymbosa, 2-3 cm. longa latasque,

bracteis foliosis, trullatus, usque ad 2 cm. longis; capitula usque ad 25,

ca. 7 mm. longa; phyllaria 8, elliptica 3.5-4.5 mm. longa, glabra:

flores 10-11, corollis 3.5-4 mm. longis, tubo 1.5-2 mm. longo, lobis

2-3 mm. longis; achaenia matura non visi; pappus nullus.

Plants scapose, up to 12 cm. tall with several fleshy, elongate,
fusiform tubers attached to the caudex beneath the soil; stems,
petioles, and adaxial surface of the leaves hirsute with tawny multi-
cellular hairs to 1.5 mm. long, the abaxial surface of the leaves gla-
brous; stems slender, 2 mm. thick at the base; basal leaves 7-15, long-
petioled, centrally or subcentrally peltate, subcircular in outline,
to 5 cm. across, the venation radiate with 7 principal veins; blades
deeply lobed one-half to two-thirds the distance to the center of
the blade, the primary lobes 7, each divided into 3 major lobes and
2 or 3 lesser lobules, the secondary lobes often again lobed, the ulti-
mate segments triangular, mucronate; petioles slender, 5-10 cm.
long; inflorescence corymbose, 2-3 cm. long and broad, tightly com-
pact, the primary branches subtended by petiolate, leafy bracts up
to 2 cm. long, the blade portion obtrullate; pedicels mostly glabrous;
calyculate bracts usually 3, subulate, up to 1.5 mm. long; heads 25
or fewer, about 7 mm. long; phyllaries 7, elliptic, 3.5-4.5 mm. long,
0.7-1 mm. wide, glabrous; flowers 10 or 11; corolla, 0.7-1 mm. wide,
elliptic, strongly 3-nerved, the throat none; anthers 2 mm. long; style
branches 0.7 mm. long, hispid on the abaxial surface, the tips truncate
or rounded with a terminal or subterminal fringe of trichomes;
mature achenes not seen, the immature achenes brown, glabrous,
10-ribbed; pappus entirely lacking.

Houotype: Guerrero; Teitepec (Mina Distr.). Open stunted pine
forest, on mossy rocks, 3,600 m. (acc. to the label), July 17, 1939,
G. B. Hinton et al. 14464 (US!, isotypes MICH!, NY!).

DistrrisuTion: Known only from the type collection. Flowering in
July.

This species has affinities with both P. peltigerum (through the
underground tubers) and P. calvum (through the lack of a pappus).
The small size makes this species unique in this genus.

7. Psacalium nelsonii Rydb. Bull. Torrey Bot. Club 51:374. 1924. Holotype:
Oaxaca; vicinity of Cerro de San Felipe, 9,500-11,000 ft., in 1894, EH. W.
Nelson 1111 (US!).

Roots, base of plant and stem below the inflorescence not seen;
leaves, presumably basal, petioled, peltate, subcircular in outline, 30-
35 cm. in diameter, the venation radiate with 7 or 8 principal veins;
blades lobed about one-half the distance to the center of the blade,
slightly longer than wide, each divided into 2 or 3 secondary lobes

424. CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

which are obtusely deltoid, mucronate-tipped, the adaxial surface
pubescent with multicellular hairs up to 1.5 mm. long, the abaxial
surface gray-tomentose; cauline leaf (only one seen) similar to the basal
leaves but subpeltate, the petiole attached 2-3 mm. above the base of
the blade, the blade 8-lobed; inflorescence paniculate, at least 35 cm.
long, 15 cm. broad, the branches purple-tomentose, the tomentum
denser toward the heads, the branches subtended by leafy bracts, 4-10
em. long, obovate, palmately 3- to 7-lobed or subentire; calyculate
bracts about 5, linear-subulate, 4-5 mm. long; heads many, about 12
mm. long; phyllaries 8 or 9, elliptic, narrowly ovate or oblong, 10 mm.
long, 2-3 mm. wide, glabrous, purplish tinted; flowers mostly immature,
10-12; corolla about 7 mm. long, purplish tinted when dry, the tube
3.5-4 mm. long, the throat 0.5-1 mm. long, 1-1.5 mm. broad, the lobes
3 mm. long; anthers 2-3 mm. long; style branches 1.5 mm. long;
mature achenes not seen, the immature achenes glabrous; pappus
white, about 5 mm. long.

Distripution: Known only from the type specimen. Flowering
time unknown.

The type collection consists only of a basal leaf and a portion of the
inflorescence. It is apparently related to P. peltatum because of the
leafy inflorescence bracts, the number of phyllaries and flowers, and
the glabrous achenes. It differs from P. peltatum because of the gla-
brous phyllaries, short subinvolucral bracts, tomentose condition of
the leaves, and the outline of the basal leaves. This specimen appears
to be significantly different from any of the known species of Psacalium
and therefore I have followed Rydberg in recognizing it as a distinct
species, despite the incomplete condition of the type specimen.

8. Psacalium peltatum (1.B.K.) Cass. Dict. Sci. Nat. 43:461. 1826.
Cacalia peltata H.B.K. Nov. Gen. et Sp. 4:170. t. 861. 1820. (Folio ed. 4:133.
1820.
Senecio. peltiferus Hemsl. Biol. Centr. Amer. Bot. 2:245. 1881. Based on
Cacalia peltata H.B.K., not 8S. peltatus DC.

Plants up to 2.5 m. tall; stems and leaves sparsely to densely pilose
or hirsute with multicellular, conical-attenuate nonglandular hairs,
0.5-2 mm. long, the crosswalls often purple, the stems also occasionally
densely covered with minute, multicellular stipitate glands, 0.3-0.5
mm. long, (lacking in var. peltatum); stems stout, terete; basal leaves
3 or 4, long-petioled, peltate, subcircular in outline, 10-30 (-65) cm.
in diameter, the venation radiate with 6-8 principal veins; blades
deeply lobed about two-thirds the distance to the center, the primary
lobes 7-9, each sinuately 5- to 7-lobed, the distal secondary lobes often
larger than the others, again lobed or 3-angulate; lowermost cauline
leaves similar to the basal ones and about as large; upper cauline

PIPPEN—MEXICAN “CACALIOID” GENERA 425

leaves progressively smaller upward, subpeltate, the uppermost non-
peltate with leafy-margined petioles, the blade palmately 3- to 5-
lobed; inflorescence racemose, paniculate or corymbose, the branches
densely stipitate-glandular, subtended by sessile, leafy bracts, ovate
or narrowly ovate, less often linear; heads 10-100, 1.5-2.5 cm. long,
phyllaries 8-10 or 12-14, pubescent; the involucre subtended by 3 or
4 narrowly ovate or linear bracts longer than the phyllaries; flowers
12-20 (-40) ; corolla about 9-14 mm. long, the tube 6-10 mm. long, the
throat ampliate, 0.5 mm. long, 1-1.5 mm. broad, the lobes narrowly
triangular, 3-4 mm. long; anthers 3-3.5 (-4) mm. long; style branches
1.5-2 mm. long, recurved, hispid on the abaxial surface, the tips
truncate or obtuse with a sparse subterminal fringe of trichomes;
mature achenes oblong or ellipsoid, 5.5-7 mm. long, 1-1.5 mm. wide,
0.7-1 mm. thick, glabrous, creamy, multistriate; pappus 6-8 (-9) mm.
long, white.

This is a variable and complex species usually distinguished by the
sessile, elliptic or lobed inflorescence bracts, the 3 or 4 long, calyculate
bracts subtending the phyllaries, the creamy, glabrous achenes, and
the usually deeply lobed basal leaves.

Within this complex are three smaller groups, recognized as varieties,
which seem to maintain themselves both morphologically and geo-
graphically.

Of several other specimens that are apparently a part of this com-
plex, two are quite similar to var. conzattii and are discussed with
that taxon. Another specimen is unique (Guerrero, between Petlacala
and Buenavista, G. B. Hinton 14877, MICH, NY) and is from a
poorly explored region where no specimens of P. peltatum have been
collected. This specimen has much less deeply lobed leaves with
much broader lobes than P. peltatum but with an inflorescence like
var. peltatum, except it is glandular-pubescent. More material from
this region as well as field observations will be necessary to determine
the best taxonomic position of this plant.

Key to Varieties of Psacalium peltatum

1. Stem and inflorescence glandular pubescent, the glands stipitate, 0.5 mm.
long; plants also pubescent with long, stiff, nonglandular hairs.
2. Heads few, ca. 10, 2-2.5 em. long; calyculate bracts 15-17 (-25) mm. long;
pappus bristles 7-9 mm. long; flowering in September and October.
8c. P. peltatum var. conzattii
2. Heads many, at least 50, ca. 1.5 cm. long; calyculate bracts 20-35 mm.
long, pappus bristles 6-7 mm. long, flowering in July and August.
8b. P. peltatum var. adenophorum
1. Stem and inflorescence lacking stipitate glands but sparsely to densely hirsute
or pilose with nonglandular hairs . . . 8a. P. peltatum var. peltatum

426 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

8a. Psacalium peltatum var. peltatum

Cacalia peltata H.B.K. as to holotype: Michoscan; in woods around Pdzcuaro
[Pdtzcuaro], 1130 hex., Humboldt & Bonpland 4346 (P, not seen; Field
Mus. Neg. 37873!).

Cacalia peltata var. coultert Greenm. Proc. Amer. Acad. 40:51. 1904. Lectotype:
Hidalgo (Veracruz on the label); Real del Monte, Th. Coulter 420 (GII!).

Psacalium coultert (Greenm.) Rydb. Bull. Torrey Bot. Club 51:373. 1924.

Psacalium argutum Rydb. Bull. Torrey Bot. Club 51:373. 1924, Holotype:
San Luis Potosf, in 1879, J. G. Schaffner 725 (NY1).

Stems and leaves sparsely or densely pilose or hirsute with non-
glandular, multicellular trichomes, 0.5-2 mm. long, the stem and
inflorescence not stipitate glandular; basal leaf lobes (7-) 10-12
(-20) em. long, 3-5 cm. wide at the base, the sinuses obtuse or broadly
rounded; petioles (10-) 20-40 em. long, the petiole of the lowermost
cauline leaf more dilated than those of the basal leaves, and leafy-
margined; inflorescence paniculate or corymbose, about 1 m. long
and broad, the leafy bracts ovate, 3-4 (-7) cm. long, 2-3 em. broad,
occasionally 3-lobed near the distal end; calyculate bracts narrowly
ovate or ovate, (11-) 12-17 (-25) mm. long, one of these usually
broader than the others surrounding the same head; heads many,
ca. 15 mm. long; phyllaries 8-10, narrowly ovate, 10-12 (-15) mm.
long, 2-3 mm. wide, pubescent but not stipitate glandular; flowers
12-17 (-20); corolla 8.5-10.5 mm. long, the tube (5.5-) 6-7 mm.
long, the lobes 3-3.5 mm. long, 1 mm. wide; anthers 3.3-5 mm. long,
pappus 7-8 mm. long; chromosome number, n=30 (Pippen).

Distrisution: From southern Chihuahua south through Durango
to Jalisco, eastward across the Trans-Mexican volcanic belt in
Michoacén and México to Puebla. Grassland openings and borders
of the pine woodlands. Flowering September and October.

The pubescence varies from sparsely short puberulent to densely
pilose or villous. Greenman distinguished the villous specimens as
var. coulteri. Apparently this was the only character used and it
does not appear to be consistent or correlated with any other charac-
ters, therefore, I have not recognized the var. coullert.

Rydberg distinguished Psacalium argutum, a group of specimens
from San Luis Potosi, as differing from P. peltatum by the presence
of longer calyculate bracts and more sharply pointed basal leaf lobes.
Although such a tendency is noted in these specimens, it is neither
consistent nor restricted to this region. There are no other significant
differences in these specimens and I have not supported this separa-
tion.

Cuinuanua: Gentry, Correll, & Arguelles 18016 (US). Duranao: Nelson 4954
(NY, US); Pennell 18343 (NY); Iberra Garcia 414 (US). Disrriro FEDERAL:
Matuda 26175, 26632 (NY). Hipauao: Pringle 9870 (GH). Jauisco: McVaugh
13821 (MICH). México: Hinton 1908 (US); Matuda 28343, 29483, 29591 (NY);
Rose & Painter 7939 (GH, US, paratypes of v. cowltert Greenm.); Lyonnet 3122

PIPPEN—MEXICAN “CACALIOID” GENERA 427

(US). MicHoactn: Nelson 6582 (GH, NY, US); Pippen 44, 49*, 53 (MICH);
Pringle 3340 (GH, NY, US); Seler 1259 (GH, NY, US). Moretos: Lyonnet 794
(US); Pringle 9871 (GH, MICH, NY, US). PuEBia: Beaman 3619 (TEX);
Purpus 3038 (NY, US). San Luis Porosf: Schaffner 294 (GH); Parry & Palmer
543 (GH, NY, US). Tuaxcata: Sharp & Hernandez X, 44508 (NY). VeERAcRUz:
Seaton s.n., August 1891 (GH, paratype of v. coulteri Greenm.).

8b. Psacalium peltatum var. adenophorum §. F. Blake, Journ. Wash. Acad.
Sei. 32:150. 1942. Holotype: Nuevo Leon: Municipio de Rayones, upper
west slopes of Sicrra de Cebolla, above 2,750 m., Aug. 21, 1939, C. H.
Muller 2911 (US!).

Plants ca. 1.5 m. tall; stems and leaves sparsely or densely pilose
or hirsute with multicellular, nonglandular hairs, the stem and inflores-
cense also densely covered with minute stipitate glands; stem 5-9
mm. thick at the base; leaves similar to those of the typical variety;
inflorescence paniculate or corymbose, ca. 30 cm. long, 11-17 cm.
broad, the bracts up to 6 cm. long, 1.5 em. broad; calyculate bracts
2.0-3.5 cm. long, 3-10 mm. wide; heads ca. 40, about 2 cm. long;
phyllaries (8—) 12 or 13, narrowly ovate, (10—-) 15-16 mm. long, 2-3
mm. wide; flowers ca. 20; corolla 13-14 mm. long, the tube (7-)
9-10 mm. long, the lobes ca. 4 mm. long; anthers 3-3.5 mm. long;
mature achenes not seen; pappus 6-7 mm. long.

DistripuTion: Known only from the southern region of Nuevo
Leon and Tamaulipas. Flowering in July and August.

This taxon is known from only one collection in addition to the
type (Tamaulipas: Stanford, Taylor, & Lauber 2535, NY, TEX, US).
The longer subinvolucral bracts and presence of stipitate glands
separate it most easily from the typical variety. Time of flowering
and geographical range separate it from var. conzatti.
8c. Psacalium peltatum var. conzattii (Rob. & Greenm.) Pippen, comb. nov.

Cacalia peltata var. conzattii Rob. & Greenm. Proc. Amer. Acad. 32:49.
1896. Lectotype: Oaxaca; Sierra de San Felipe, 10,000 ft., Dec. 13, 1895,
C. G. Pringle 6238 (GH!).

Psacalium conzattit Rydb. Bull. Torrey Bot. Club 51:372. 1924.

?Psacalium mollifolium S. F. Blake, Journ. Wash. Acad. Sci. 28:491. 1938.
Holotype: Guerrero; Taxco, Aug. 12, 1937, Ruth Q. Abbott 353 (GH)).

Plants usually 1 m. tall or less; stems and leaves sparsely or densely
pilose or hirsute with nonglandular trichomes, these especially dense
on the abaxial surface of the leaves, the stem and inflorescence also
densely stipitate-glandular; stem 4-10 mm. thick at the base; basal
leaves subcircular, 14-25 cm. in diameter; blades deeply lobed usually
more than three-fourths the distance to the center, the primary lobes
4-10 cm. long, 2-4 cm. wide at the base; inflorescence sub paniculate
10-20 (-30) em. long, 11-16 cm. broad, the branches subtended by
bracts about 3 cm. long, 0.7-1.5 cm. wide; calyculate bracts narrowly
elliptic, 15-17 (-25) mm. long, 1-2.5 mm. wide; heads usually 10

266-—836—68——-5

428 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

(-30), 2-2.5 em. long; phyllaries 12-15, narrowly ovate, 12-15 (-18)
mm. long, 2-3 (-4) mm. wide, pilose with nonglandular hairs, stipitate-
glandular; flowers (20-) 25-40; corolla 10-12 mm. long, the tube
(6-) 7-8 (-9) mm. long, the lobes 3.5-4 mm. long, 1 mm. wide; an-
thers 3-3.5 (-4) mm. long; mature achenes 6-7 mm. long, 1-1.5 mm.
wide, ca. 10-ribbed; pappus 7-8 (-9) mm. long.

DisrriputTion: Known mostly from central Oaxaca and question-
ably from Guerrero and Durango. Openings in pine woodlands.
Flowering in September and October.

Two specimens were cited in the original description as syntypes:
Oaxaca; Cerro de San Felipe, 9,000 ft., Nov. 29, 1895, C. Conzatti
27 (GH!), Dec. 13, 1895, C. G. Pringle 6238 (GH!). Despite the fact
the taxon was named for Conzatti, possibly indicating his specimen
as the lectotype, Pringle 6238 is a more complete and more repre-
sentative specimen and is thus a better choice for the lectotype.

Two specimens have been studied that are obviously closely
related to this taxon but differ in certain respects. One specimen
(15 mi. W. of El Salto, U. T. Waterfall 12688, MICH) has heads
and stipitate glands like var. conzattii but the leaves are smaller
with obovate lobes; most significant is its occurrence in Durango
which is quite far from Oaxaca. Such a separation in range is difficult
to explain if this specimen is a part of this taxon.

The other specimen (Guerrero: Taxco, Ruth Q. Abbott 353, GH)
was the basis of Psacalium mollifolium S. F. Blake. It differs from
var. conzattii in that the basal leaves are lobed less than half the
distance to the center and the abaxial surface is densely tomentose.
The inflorescence is similar to var. conzattit, including the long calycu-
late bracts. These differences do not appear significant enough to
give this specimen nomenclatural status; at least not until field
observations and study of future material can be made.

Oaxaca: Pringle 6238 (US, isotype); Camp 2401 (NY); McVaugh 21826
(MICH).

9. Psacalium peltigerum (Rob. & Seat.) Rydb. Bull. Torrey Bot. Club 51:374.
1924,

Cacalia peltigera Rob. & Seat. Proc. Amer. Acad. 28:111. May 13, 1893.
Plants up to 1.5 m. tall with several fleshy underground tubers up
to 5 cm. long, 3 em. thick attached to the caudex; stems 5—12 mm.
thick at the base, mostly glabrous or sparsely hirsute; leaves sparsely
or densely pubescent with stiff, white hairs up to 1 mm. long; basal
leaves 2 or 3, long-petioled, peltate, subcircular in outline, (10—-) 20-30
cm. in diameter, the venation radiate with 7-9 (-12) principal veins;
blades angulate or lobed one-third to four-fifths the distance to the
center, the primary lobes 9-11, each divided into 2 or 3 major lobes
and several lesser lobules, the secondary lobes usually lobed or coarsely

PIPPEN—-MEXICAN “CACALIOID” GENERA 429

toothed, or if the blade angulate the margins mucronate-tipped;
petioles 29-30 cm. long, pubescent proximally, becoming glabrous to-
ward the blade; cauline leaves 3 or 4, the lowermost similar to the
basal ones but smaller; upper cauline leaves progressively smaller up-
ward, either nonpeltate with the blades 3- to 7-palmately lobed or
remaining peltate and angulate, not lobed; inflorescence mostly corym-
bose, 9-30 em. long and broad, the branches subtended by linear-
subulate bracts, 1.5—-2 cm. long, 1-2 mm. wide; pedicels 1-2 em. long;
calyculate bracts 3-5, subulate, 2 (-5) mm. long; heads many, 12-15
mm. long; phyllaries 5, narrowly oblong, narrowly elliptic, or occa-
sionally spathulate, (6—) 7-8 (-10) mm. long, 1-2 mm. wide, glabrous;
receptacle 1-1.5 mm. across; flowers 5 or 6; corolla 9-10 mm. long, the
tube (3.5-) 4-5 mm. long, the lobes as long as the tube or slightly
longer, narrowly elliptic, the throat none; anthers 3-3.5 mm. long,
style branches 1-1.5 mm. long, distally hispid on the abaxial surface,
the tips truncate or rounded with a subterminal or terminal fringe of
trichomes; mature achenes ellipsoid, 4-5.5 mm. long, 1.5-2 mm. thick,
glabrous, tan, prominently 10-ribbed; pappus (4—) 5-5.5 (—6) mm.
long, creamy or tawny.

This species is easily distinguished by the presence of fleshy sub-
terranean tubers. Microscopic examination of freehand sections reveals
that these tubers are modified stems with discrete vascular bundles.
Druse-shaped crystals were found in the parenchyma cells. In green-
house-grown plants these tubers formed during the first season of
growth and persisted through the dormant period. There isnoevidence
that they function in vegetative propagation as do the tubercles in the
genus Pericalia.

Three varieties are recognized and may be separated primarily by
the outline of the basal leaves and geographical distribution, These
three varieties exhibit an interesting range in leaf shapes from almost
entire (var. hintonii), to broadly lobed (var. larilobum), to deeply
linear-lobed, the lobes five times longer than broad (var. peltigcrum).

Key to Varieties of Psacalium peltigerum

1. Basal leaves angulate, not lobed, the margin with mucronate tips; stem hir-

sute ..... eee ee ee ew ww es 6 OD. P. peltigerum var. hintonii

1, Basal leaves lobed at least half the distance to the center, the margins toothed,

the teeth triangular, not mucronate; stems mostly glabrous or very sparsely
hirsute.

2. Lobes of the basal leaves extending two-thirds to four-fifths the distance

to the center of the blade, 1-2 em. wide at the base, much longer than

broad, the ultimate segments of the lobes narrowly triangular or linear-

attenuate, about 5 times longer than wide; stem glabrous.
9a. P. peltigerum var. peltigerum

430 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

2. Lobes of the basal leaves extending less than one-half the distance to the
center of the blade, 5-7 cm. wide at the base, about as long as wide, the
ultimate segments of the lobes obtusely deltoid or triangular, about as
long as broad; stem hirsute at least on the proximal half.

9c. P. peltigerum var. latilobum

9a. Psacalium peltigerum var. peltigerum
Cacalia peltigera Rob. & Seat. as to Holotype: Jalisco; bluff of the barranca
near Guadalajara, Sept. 14, 1891, C. G. Pringle 5154 (GH!).

Blades of the leaves deepiy lobed two-thirds to four-fifths the
distance to the center, the primary lobes 9-11, 1-2 em. wide at the
base, much longer than wide, each subdivided into 2 or 3 major lobes
and several lesser lobules, the secondary lobes usually lobed or coarsely
toothed, the ultimate segments narrowly triangular or linear-atten-
uate, ca. 5 times longer than wide.

DisrriBution: Central and western Jalisco. In the dry grasslands
around Guadalajara and the pine woodlands to the west. Flowering
in July and August.

In the original description two specimens were listed as syntypes:
Jalisco; Rio Blanco, July 1886, #. Palmer 171 (GH!); bluff of the
barranca near Guadalajara, Sept. 14, 1891, C. G. Pringle 5154 (GH!).
Pringle 5154 is chosen as the lectotype because it is more complete.
The name was published in a paper concerning plants discovered by
Pringle, and this specimen is annotated in what appears to be Robin-
son’s writing.

The leaves of this variety exhibit considerable latitude in shape and
lobing, but are always deeply lobed with the ultimate segments very
narrow. ‘The inflorescence is more elongate and broader than in the
other varieties.

Jauisco: Pringle 4627, 9872 (GH, NY, US); King 3656 (MICH, NY, TEX,
US); McVaugh 18603, 15094, 20280 (MICH); Wilbur & Wilbur 1996 (US).
9b. Pscalium peltigerum var. hintonii Pippen, var. nov. PLATE 9

Varietas peltigera foliis profunde lobatis, caule glabro, inflorescentia
majore differt.

Plants to ca. 0.5 m. tall; stems hirsute with multicellular white
hairs to 2 mm. long; blades of the basal leaves shallowly 9- to 11-
angled but not lobed, the margin mucronate-toothed; upper cauline
leaves similar to but smaller than the lower ones; inflorescence tightly
corymbose, 10 cm. long and broad (those examined not mature).

Hoxotyps: México; Carboneras, Toma, July 21, 1935, G. B. Hinton
et al. 7966 (MICH!, isotype NY}).

DistriBuTiIon: Known only from the type collection.

The floral characters are similar to var. peltigerum and the plants
have similar fleshy subterranean tubers. They differ in that the leaves
are angulate, not deeply lobed, and the stem is hirsute. Field obser-

PIPPEN—MEXICAN “CACALIOID” GENERA 431

vations and study of future material may indicate that var. hintonii
deserves specific rank, but for the present it is considered only as a
variety.

9c. Psacalium peltigerum var. latilobum Pippen var. nov. PuaTE 10

Varietas peltigera foliorum lobis longioribus quam latioribus, caule
glabro differt.

Plants to ca. 0.5 m. tall; stems sparsely pubescent; blades of the basal
leaves lobed less than one-half the distance to the center, the primary
lobes 10 or 11, 5-7 cm. wide at the base and equally long, the lobes
subdivided into 2 or 3 secondary lobes, obtusely triangular or deltoid,
about as broad as long; margins of the lobes coarsely, triangular-
toothed; inflorescence corymbose, tightly compact, ca. 9 cm. long
and broad (inflorescence not fully matured).

Hoxotyre: Guerrero; 1}, miles W. of Omiltemi, 7,800 ft., June 11,
1953, T. W. McCorele & C. M. Rowell, Jr. 3453 (MICH, sheet 1!;
isotype, sheet 2!).

Distripution: Known only from the type collection.

10. Psacalium poculiferum (S. Wats.) Rydb. Bull. Torrey Bot. Club 51:375,
1924,

Slender herbs ca. 1 m. tall; roots exuding a reddish-orange sap when
cut; stems 4-7 mm. thick at the base, densely puberulent proximally
becoming subglabrous upward and again puberulent in the inflo-
rescence; leaves glabrous or sparsely puberulent mostly along the veins
with 6- to 10-celled hairs ca. 0.5 mm. long; basal leaves 2 or 3, long-
petioled, centrally or subcentrally peltate, subcircular in outline, 15—
25 (-60) cm. in diameter; blades lobed one-third to two-thirds the
distance to the depressed center, the primary lobes 7 or 8 (—10), 4-8
cm. wide at the base, usually 2-3 cm. longer than broad, each divided
into 2 or 3 secondary lobes or merely triangulate, the margins irregu-
larly toothed; petioles 15-30 (—40) cm. long, terete, slender, usually
sparsely pubescent; cauline leaves 3 or 4, the lowermost similar in
size and shape to the basal ones except the petiole leafy-margined
and dilated at the base; cauline leaves progressively smaller upward;
the uppermost bracteiform, nonpeltate, the petiolar portion leafy-
margined, auriculate-based, the blade portion 5- to 7-lobed; inflo-
rescence paniculate or corymbose, 12-30 cm. long and broad, the lower
branches in the axils of the upper leaves; pedicels tomentose; calycu-
late bracts 3, subulate, about 1 mm. long (or lacking); heads about
100, 9-10 mm. long; phyllaries 3-5 (usually 4), varying on the same
plant, elliptic or obovate, (4-) 5-6 (-7) mm. long, 1-3 mm. wide,
glabrous, the tips purplish; flowers 3-5, opposite the phyllaries;
corolla 6-8 mm. long, the tube as long as the lobes, 3-4 mm., the
lobes linear, 0.5 mm. wide, the throat none; anthers 2-3 mm. long;

432 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

style branches ca. 1 mm. long, the tips truncate or rounded with a
terminal or subterminal fringe of trichomes; mature achenes oblong
or narrowly ellipsoid, 5 mm. long, 1--1.6 mm. wide, pubescent with
fuscous, unicellular hairs; pappus (3.5-) 5-6 (-8) mm. long, creamy.

DistriputTion: Western Jalisco to Nayarit, north to Sinaloa.
Rocky, grass-oak dominated slopes and pastured areas. Flowering in
mid-July and August.

This species is unusual because of the very small heads with mostly
4 phyllaries and flowers. It is closely related to P. tabulare but differs
in its sparser pubescence and smaller heads. Both, however, have
similar leafy, auriculate-based, cauline leaves.

Jatisco: Pippen 19, 21, 27, 60 (MICH); Pr*ngle 9874, 4414 (GH, NY, US);
Wilbur & Wilbur 2017, 2166 (MICH, US). Nayarit: Rose 1935 (GH, US);
King 3677 (MICH, NY, TEX, US); Pippen 11 (MICH); McVaugh 16406, 18838
(MICH). Sinatoa: Ortega 7146 (US, the leaf of this specimen is somewhat
atypical but this inflorescence is very similar to P. poculiferum).

11. Psacalium tabulare (Hemsl.) Rydb. Bull. Torrey Bot. Club 51:375. 1924.
Senecio tabularis Hemsl. Biol. Centr. Amer. Bot. 2:248. 1881. Holotype:
Veracruz; valley of Orizaba, Mscamella, Aug. 16, 1866, Bourgeaw 2926
(KX, not seen; isotypes GH! and C, Field Mus. Neg. 22631!).
Cacalia tabularis (Hemsl.) A. Gray, Proc. Amer. Acad. 19:52. 1883.

Plants up to 2 m. tall (Balls 5350); stems at least 7 mm. thick at
the base, purplish, densely pubescent; leaves sparsely pubescent on
the adaxial surface, densely pubescent on the abaxial surface, the
hairs multicellular, 1-1.5 mm. long, 15- to 20-celled; basal leaves 2-4,
long-petioled, peltate, subcircular in outline, 16-27 (-60, Balls 5350)
cm. in diameter, the venation radiate with 8 principal veins; blades
lobed about half way to the center, the primary lobes 7 or 8, 5-8 em.
long and broad, each usually divided into 3 triangular secondary lobes,
1-2.5 cm. long, 2-3 cm. broad, the margins coarsely deltoid-toothed ;
petioles up to 35 cm. long, terete, pubescent; cauline leaves 2—4; the
lowermost subcentrally peltate, the blades broadly ovate or subcir-
cular, 14-16 cm. long, 15-17 cm. wide, palmately 6- to 7-lobed, the
petioles attached to the blade about 3 cm. above the base, leafy-mar-
gined, auriculate-based; uppermost cauline leaves sessile, auriculate-
based, 3-lobed distally; inflorescence paniculate, 30-40 em. long, 8-30
em. broad, the branches often sparsely stipitate glandular, subtended
by narrowly triangular bracts 1-5 em. long, 0.3-1 em. wide; pedicels
5-10 mm. long, with 2-5 subulate bracts, 2-6 mm. long; heads 75 to
over 100, about 12 mm. long; phyllaries 5, narrowly ovate, 9-10 mm.
long, 2-3 mm. broad, mostly glabrous or occasionally sparsely pu-
bescent; receptacle 1.5 mm. across; flowers 5, opposite the phyllaries;
corolla 7-9 mm. long, the tube (3.5—-) 4-5 mm. long, the throat cam-
panulate 0.5-1 mm. long and wide, the lobes 3 mm. long, linear; an-
thers 3 mm. long; style branches 1-1.5 mm. long, hispid on the abaxial

PIPPEN—MEXICAN “CACALIOID” GENERA 433

surface, the tips truncate or obtuse with a terminal or subterminal
fringe of trichomes; mature achenes narrowly ellipsoid, 5-5.5 mm.
long, 1.5 mm. wide, 1 mm. or less thick, glabrous, pale, about 18-
ribbed; pappus (5.5-) 6-7 mm. long, white.

DistriBuTION: Known mostly from western Veracruz and eastern
Puebla and one specimen collected in Michoacén. Openings in oak-
pine woodlands. Flowering from August to October.

This species is very closely related to P. poculiferum, but differs
in that the heads always have 5 phyllaries and flowers, and glabrous
achenes,

Some of the specimens examined are sparsely stipitate-glandular
on the inflorescence branches but this is not constant in this species.

Puresia: Purpus 5630 (GH, NY, US). Micnoackn: Pippen 55 (MICH).
Veracruz: Balls 5350 (US); Muller s.n., August 1843 (NY); Sharp 44823 (NY).
12. Psacalium species

Plants to about 1 m. tall; stems slender, to 5 mm. thick at the
base, subscapose, densely hirsute below with white, nonglandular
multicellular hairs up to 1 mm. long, distally sparsely hirsute with
nonglandular hairs as well as minutely stipitate glandular; leaves
glabrous on the adaxial surface, sparsely pubescent along the veins
on the abaxial surface; basal leaves 2 or 3, long-petioled, peltate,
subcircular in outline, 20-30 em. across, venation radiate with about
8 principal veins; blades lobed one-third to one-fourth (rarely one-
half) the distance to the center of the blade, the primary lobes 6 or
7, 5-8 cm. wide at the base about as long as wide, each usually divided
into 3, triangular attenuate-tipped secondary lobes; petioles slender,
to 30 cm. long, pubescent; cauline leaves 1 (or 2), similar to the
basal leaves but much smaller; inflorescence paniculate, ca. 35 cm.
long, 25 em. broad, the branches densely stipitate-glandular, sub-
tended by petiolate, leafy bracts, the blade to 6 cm. long, elliptic,
entire or palmately 3- to 5-lobed; pedicels purple, to 3 cm. long;
calyculate bracts 3-7, subulate, 4-5 mm. long; heads few, immature;
phyllaries (7 or) 8, oblong or elliptic, ca. 10 mm. long, 2-3 mm. wide,
hirsute, occasionally sparsely glandular; flowers ca. 9, achenes gla-
brous; pappus white, chromosome number, n=30 (Turner, ined.).

DistrisuTion: Known only from Guerrero. Flowering probably in
November.

These specimens appear to be somewhat intermediate between
Psacalium eriocarpum and P. holwayanum. They differ from both of
these species by the glabrous achenes, the leafy inflorescence bracts,
the mostly glabrous basal leaves, as well as the distinct geographical
range.

These specimens probably represent either a distinct species or a
variety of one of the above. The material examined was immature

434 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

and could not be studied thoroughly. Perhaps after future collections
have been studied and field observations made it will be possible to
assign a name,

GuERRERO: McVaugh 21913* (MICH); Sharp 441496 (NY).
Doubtful Names and Excluded Taxa

Cacalia albicans Sessé & Moc. Fl. Mex. ed. 2. 180. 1894. Known only from the
type description. This cannot be associated with any species of the genera
discussed in this paper. According to the description the leaves are lanceolate
and sinuately pinnatifid and therefore this probably should be referred to
Senecio.

Cacalia amplexicaulis Schultz Bip. ex Klatt, Leopoldina 24:125, in synonymy,
1888. Listed as a synonym of Senecio bracteatus Klatt.

Cacalia berlandiert DC. Prod. 6:328. 1838. =—Senecio desortum Hemsl. (Biol.
Centr. Amer. Bot. 2:239. 1881).

Cacalia calotricha S. F. Blake, Contr. Gray Herb. II. 52:58. 1917. This should
be referred to Senecio. The structure of the corolla is similar to that of S.
angulifolius in that the throat is longer than the recurved lobes. The plant
is apparently a shrub. It is known only from the type.

Cacalia eriopoda Schultz Bip. ex Klatt, Leopoldina 24:125, in synonymy, 1888.
Listed as synonym of Senecio eriopodus Klatt.

Cacalia guatemalensis Standl. & Steyermark, Publ. Field Mus. Bot. 23:99. 1944.
This should probably be referred to Senecio sect. Mulgedifolii since the
description states that the flowers are purple and the inflorescence is glan-
dular pubescent. This species is not a Psacaliuwm, as suggested by Cuatrecasas
(Brittonia 8:157. 1955) since, according to the description, the leaves are
not peltate.

Cacalia incana Sessé & Moc. Pl. N. Wisp. ed. 1. 132. 1889. A later homonym of
C. incana L., 1753.

Cacalia liebmannii Schultz Bip. ex Klatt, Leopoldina 24:125, in synonymy, 1888.
Listed as a synonym of Senecio liebmannii Buchinger.

Cacalia lyrata Sessé & Moc. Pl. N. Hisp. ed. 1. 131. 1889. Known only from the
type description. This is probably referable to Senecio sect. Mulgedifolti
because, according to the description, the flowers are purple and the leaves
are lyrate-pinnatifid.

Cacalia macrophylla Sessé & Moc. Pl. N. Hisp. ed. 1. 132.1889. Later homonym
of Cacalia macrophylla Bieb. 1808.

Cacalia pallescens Schultz Bip. ex Klatt, Leopoldina 24:126, in synonymy, 1888.
Listed as a synonym of Senecio pallescens Klatt. Rydberg transferred this
species to Odontotrichum based on a tracing in Klatt’s herbarium.

Cacalia parasitica Schultz Bip. Bot. Zeit. 15:759. 1857. According to Greenman
(Contr. U.S. Nat. Herb. 23:1635. 1924) this is Senecio parasiticus Greenm.
It is a shrub. No specimens have been seen by me.

Cacalia peltata Sessé & Moc. Pl. N. Hisp. ed. 1. 132. 1889. A later homonym of
Cacalia peltata H.B.K. [Psacalium peltatum (H.B.K.) Cass.]

PIPPEN:

MEXICAN “CACALIOID” GENERA 435

Cacalia pinnatifida Sessé & Moc. Pl. N. Hisp. ed. 1. 132. 1889. A later homonym
of C. pinnatifida L., 1767. The description and photograph of the type (an
illustration) indicate that this is probably Odontotrichum amplifolium (DC.)
Rydb,

Cacalia prenanthoides T1.B.K. Nov. Gen. et Sp. 4:167. 1820. =Erechtites valeri-
anaefolia f. prenanthoides, Belcher (Ann. Mo. Bot. Gard. 43:30. 1956).

Cacalia purpurascens Schultz Bip. ex Klatt, Leopoldina 24:126, in synonymy,
1888. Listed as a synonym of Senecio purpurascens Klatt.

Cacalia pudica Standl. & Steyermark, Publ. Field Mus. Bot. 23:255. 1947. This
is not in any of the genera herein considered, because the flowers are purple
and the throat of the corolla is much longer than the lobes. It is more closely
related to Senecio Sect. Mulgedzfoliz.

Cacalia runcinata H.B.K. Nov. Gen. ct Sp. 4:168. 1820. =Senecio roseus Schultz
Bip. (Flora 28:498. 1845).

Cacalia runcinata Less. Linnaea 5:162. 1830. =Senecio coulteri Greenm. (Ann.
Mo. Bot. Gard. 1:272. 1914).

Cacalia sarracenica Sessé & Moc. Fl. Mex. ed. 2. 180. 1894. A later homonym
of Cacalia sarracenica L.., 1753.

Cacalia sinuata Sessé & Moc. Pl. N. Hisp. ed. 1. 1889. A later homonym of
Cacalia sinuata Ceryv. 1824 [Odontotrichum sinuatum (Cerv.) Rydb.].

Cacalia spathulata Sessé & Moc. ex D. Don, Trans. Linn. Soc. 16:250. 1833.
=Chaptalia nutans (according to Blake).

Cacalia subdecurrens Schulz Bip. ex Klatt, Leopoldina 24:125, in synonymy,
1888. Listed as a synonym of Senecio deformis Klatt.

Cacalia suffulta Greenm. Proc. Amer. Acad. 32:310. 1897. This probably should
be referred to Senecio. It is not a member of the genera herein discussed,
because the flowers are bright orange, the throat of the corolla is longer than
the lobes, and the achenes have hygroscopic trichomes.

Cacalia tepicana M. E. Jones, Contr. West. Bot. 15:156. 1929. =Senecio hartwegii
Benth. according to Blake (Contr. U.S. Nat. Herb. 29:136. 1945).

Cacalia toluccana DC. Prod. 6:328. 1838. =Senecio callosus Schultz Bip. according
to Greenman (Ann. Mo. Bot. Gard. 1:271. 1914).

Cacalia tuberosa Sessé & Moc. Pl. N. Hisp. ed. 1. 182. 1889. A later homonym of
Cacalia tuberosa Nutt. 1818.

Cacalia viscosa Sessé & Moc. Pl. N. Hisp. ed. 1. 1382. 1889. Known only from the
type description which states that it is a shrub; therefore, it is most probably
not a species in the genera discussed in this paper.

Odontotrichum delphinifolium Rydb. Bull. Torrey Bot. Club 51:419. 1924.
Rydberg saw only the type specimen which was in fruit. This species has
since been collected in flower. It is not an Odontotrichum, because the inflo-
rescence has only one large head, yellow radiate flowers, and no pappus.
This taxon should probably be referred to the Heleneae.

436 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Odontotrichum liebmannii (Buchinger) Rydb. Bull. Torrey Bot. Club. 51:419.
1924. Rydberg transferred this species from Senecio on the strength of a
tracing of a leaf in the Klatt Herbarium at the Gray Herbarium. I have
seen no specimens and cannot associate the description or the tracing of
Klatt’s with any species of the genera of this paper (see C. liebmannii Schultz
Bip. above).

Psacalium thyrsoideum DC. Prod. 6:335. 1838. Known only from the type de-
scription. It was questionably published by DeCandolle as a Psacalium.
Thus far the description has not been matched with any other cacalioid
species. The type specimen is probably in Prague and must be examined
before the position of this taxon can be determined.

Collections Cited Excluding Types

(Abbreviations: D.=Digitacalia; O.=Odontotrichum; Pe.=Pericalia; Ps.=
Psacalium.)

Arsine, G, ConzattI, C.
s.n. Nov. 1908, Ps. megaphyllum 715. D. heteroidea
s.n. Sept. 9, 1909, D. jatrophoides 4015. O. purpusii
s.n. Nov. 4, 1909, Pe. sessilifolia
s.n, Sept. 14, 1911, Pe. sessilifolia Conzartt, C., & Gonzauez, V.
93. O. sinuatum 397. O. amplifolium
93a. O. sinuatum 1237. O. purpusii

334. O. sinuatum
1181. O. sinuatum
2386. Ps. megaphyllum s.n. July 4, 1937, O. decompositum
5132. O. palmeri
5729. Pe. sessilifolia
5740. O. sinuatum 1235. O. decompositum
7347. Pe. sessilifolia

Darrow, R.

Darrow, R., Puiures, & Putz

Dopps, Donatp, & Simpson, Rost. F.

Arskne, G., & Nicouas, Bro. 32. Pe. michoacana

334. Ps. hyll
3 Ss. megaphyllum DiaveEt, L.

Batis, Epwarp K. s.n. July 1912, O. palmeri

5299. Ps. peltatum v. peltatum

5333. O. amplifolium Duass, A.
5350. Ps. tabulare 451. O. sinuatum
5428. O. tussilaginoides 481. Pe. sessilifolia

5575. Pe. silifoli
€. sessuuola Eaaueston, W. W.
BARCLAY 10765. O. decompositum

s.n. QO. pringlei
pring FEDDEMA, CHARLES

BarkKLeEy, Frep A, 10. O. sinuatum

14A624. O. decompositum 161. Pe. sessilifolia
183. O. palmeri
BEAMAN, JOHN 308. O. palmeri

3619. Ps. peltatum v. peltatum 381. Pe. sessilifolia

1345. Pe. ilifoli
BLUMER, J. C. 345. Pe. sessilifolia

Ferris, Roxana 8.
5881. O. pringlei

1378. O. decompositum

BourRGEAU
715. Pe. sessilifolia

716. O. sinuatum 2491 bis. O. cirsiifolium
2926. Ps. tabulare

GALEotTTI, H.

Gentry, Howarp 8.
Camp, W. H. 536M. O. decompositum
2401. Ps. peltatum v. conzattii 1959. O. decompositum
437

438

2815. O. decompositum
6968. Pe. sessilifolia
6969. O. pachyphyllum
8562. O. sinuatum

GENTRY, Howarp 8., CorreEui, &
ARGUELLES

18016. Ps. peltatum v. peltatum

GHIESBREGHT

805. QO. cirsiifolium
GILuy, C. L., Sr.

5. Ps. holwayanum

GoupMAN, E. A.

132. O. sinuatum

Goppina, L. N.

s.n. Aug. 28, 1912, O. decompositum
784. O. decompositum

GouLp, Frank W.
8989. O. sinuatum

GRAHAM, SHIRLEY

94, O. sinuatum
99. O. globosum
100. O. radulifolia
124, O. palmeri

HARTWEG

125. O. sinuatum

Hewitt, W. P.

68. O. decompositum

Hinton, G. B.

1266. O. goldsmithii v. goldsmithii
1908. Ps. peltatum v. peltatum
2200. Ps. megaphyllum

2355. Ps. megaphyllum

2487. Ps. holwayanum

2755, Pe. michoacana

4993. Pe. michoacana

7984. O. goldsmithii v. goldsmithii
8066. O. goldsmithii v. goldsmithii
8211. O, cirsiifolium

8349. O. amplifolium

9202. O. goldsmithii v. goldsmithii
9436. O. cirsiifolium

12146. O. brachycomum

12572. D. napeifolia

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

12718. Pe. michoacana

15463. O. amplifolium

13469. Pe. michoacana

13480. Ps. megaphyllum

14464. Ps. nanum

14877. Ps. peltatum v. peltatum
15078. O. goldsmithii v. goldsmithii
15288. O. brachyecomum

15624. O. brachycomum

15629. Ps. megaphyllum

Iparra Gractra, P.

407, O. globosum
414. Ps. peltatum v. peltatum

JongEs, Marcus E.

25053. O. decompositum
27692. O. pringlei

Jony, P.
s.n. in 1892. O. platylepis

Kearny, T. H., & Harrison, G.

6045. O. decompositum
6194. O. decompositum

Kearny, T. H., & Presses
10092. O. decompositum

KeENoyeR, L. A.

2110. O. radulifolium
2330. O. radulifolium

Kina, Ropert M.

3615. QO. silphiifolium

3656. Ps. peltigerum v. peltigerum
3659. O. palmeri

3677. Ps. poculiferum

6737. O. sinuatum

Kine, Ropert M., & Soperstrom, T. R.

4904. O. brachycomum

Knostocn, I. W.

1273. O. globosum
1293. O. decompositum

Kruckebera, A. R.
4644. O. decompositum

LEAVENWORTH, Wm. C.
539. O. palmeri

PIPPEN—MEXICAN

LEesvEuR, HarpDE

145, O, decompositum

LyonneEt, E.

194. Pe. sessilifolia
794. Ps. peltatum v. peltatum
3122. Ps. peltatum v. peltatum

Mannine, W. E. & M. 8.
53560. O. radulifolium

Matupa, E.

19148, O. silphiifolium

19417. O. silphiifolium

19706. O. sinuatum

21808. O. sinuatum

21825. Pe. sessilifolia

26175. Ps. peltatum v. peltatum
26572. Ps. megaphyllum

26652. Ps. peltatum v. peltatum
26882. O. sinuatum

27730. O. amplifolium

28343. Ps. peltatum v. peltatum
28821. O. cirsiifolium

29313. O. amplifolium

29483. Ps. peltatum v. peltatum
29591. Ps. peltatum v. peltatum
29609. O. amplifolium

29787. Pe. sessilifolium

30015. Ps. holwayanum

MAYSILLEs, JAMES

7579.
7587.
7752.
7755.
7784,
7793.
7880.
8448.
8485.

Pe. segsilifolia

O. radulifolium
O. sinuatum

Pe. sessilifolia

. pachyphyllum
. sinuatum

. Sinuatum

. sinuatum

. Sinuatum

ooo0°0

McVauau, Rogers

12810. O. sinuatum

12929. O. palmeri

13321. Ps. megaphyllum

13559. Pe. sessilifolia

13616. O. pringlei

13625. Pe. sessilifolia

13684, Ps. eriocarpum

13821. Ps. peltatum v. peltatum
14093. Pe. michoacana

“CACALIOID”

GENERA 439

14136. Pe. michoacana

14165. Ps. eriocarpum

14287, Ps. eriocarpum

14290. Pe. michoacana

14390. Pe. michoacana

15094. Ps. peltigerum v. peltigerum
15313. O. cervinum

15460. O. goldsmithii v. goldsmithii
16132. O. goldsmithii v. goldsmithii
16406. Ps. poculiferum

17118. Pe. sessilifolium

17160. D. jatrophoides

17183. Pe. sessilifolia

17650. Ps. megaphyllum

17726. O. amplum

18121. Pe. sessilifolia

18603. Ps. peltigerum v. peltigerum
18838. Ps. poculiferum

20280. Ps. peltigerum v. peltigerum
20481. O. palmeri

21355, Pe. michoacana

21658. Pe. sessilifolia

21659. O. pachyphyllum

21729. O, pachyphyllum

21731. O. pachyphyllum

21732. O. globosum

21733. O. sinuatum

21734-39. O. globosum x O. sinuatum
21765. O. pringlei

21823. D. napeifolia

21824. O. amplifolium

21826. Ps. peltatum v. conzattii
21897. Ps. megaphyllum

21903. D. tridactylitis

21910. Ps. megaphyllum

21928. Ps. megaphyllum

21934. O. brachycomum

21953. Ps. holwayanum

21960. O. palmeri

21964. O. brachycomum

McVauau, R., & Koriz, W.

39. Ps. megaphyllum
48. O. platylepis

49. O. sinuatum

57. O. amplum

147. Pe. sessilifolia
276. O. sinuatum
435. Pe. michoacana
441. O. palmeri

479. Pe. sessilifolia
601. O. pringlei

656. Pe. sessilifolia

440)

792. Pe. sessilifolia
919. Ps. eriocarpum
920. Pe. sessilifolia
1104. O. pringlei
1253. O. pringlei
Mexia, YNEZ
635. Pe. sessilifolia
845. O. pringlei
877. Ps. megaphyllum
1450. Pe. sessilifolia
1656. Pe. michoacana
Mearns, Epaar A.
531. O. decompositum
2219. O. decompositum
Meyer, F. G., and Rogers, D. J.
2608. O. radulifolium
MickEL, Joun T.
501. O. radulifolium
561. O. radulifolium
Moors, H. E., Jr.

3421. O. silphiifolium
5119. Ps. megaphyllum

Moors, H. E., Jr. & Cerro, M.

5469. Pe, michoacana
Mutter, C. H.
3420. O. decompositum
Miuuer, F,

s.n. in 1853. Ps. tabulare
1619. QO. silphiifolium

Narvagz Monets, M., & SALAzar, A. E.

557. O. platylepis
Newson, E. W.

1061. O. amplifolium

1133. D. napeifolia

1420. O. cirsiifolium

2080. D. tridactylitis

2243. O. amplifolium

3117. O. cirsiifolium

3467. D. chiapensis

4758. O. decompositum

4759. O. globosum

4954, Ps. peltatum v. peltatum

6582. Ps. peltatum v. peltatum
OwWNBEY, GERALD B., & OWNBEY,

FINDLEY
1874. O. pachyphyllum

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Orcutt, C. R.
3746. O. silphiifolium

OrTEGA, JESUS G.
5727. Pe. sessilifolia
7146. Ps. poculiferum
PALMER, EDWARD
s.n. July 1890. O. decompositum
s.n. 1892. O. pringlei
47. Pe. sessilifolia
168. O. palmeri
576. Pe. sessilifolia
651. O. sinuatum
689. O. platylepis
1234. O. pringlei
1832. Pe. sessilifolia
Parry & PALMER
541. Pe. sessilifolia
543. O. radulifolium
543. Ps. peltatum v. peltatum

PENNELL, Francis W.
18332. O. pachyphyllum
18348. Ps. peltatum v. peltatum
18345. Pe. sessilifolia
18582. O. globosum

Puituips, Epwin A.
651. O. decompositum

PipPEN, Ricuarp W.

1. O. sinuatum
2. O. amplum
3. O. amplum
4. O. amplum

5. Pe. sessilifolium
6. Ps. peltatum v. peltatum
7. O. sinuatum

8. Ps. megaphyllum
9. O. sinuatum

10. O. palmeri

11. Ps. poculiferum
12. O. palmeri

13. Pe. sessilifolia
15. Pe. sessilifolia
16. Ps. megaphylum
17. Pe. sessilifolia
18. Pe. sessilifolia
19. Ps. poculiferum
20. O. palmeri

21. Ps. poculiferum
22. O. sinuatum

23. O. platylepis

PIPPEN—MEXICAN

24. Ps. megaphyllum

25. O. multilobum

26. Pe. michoacana

27a. O. palmeri

27. Ps. poculiferum

28. O. palmeri

29. O. ef. palmeri

30. O. multilobum

31. Pe. sessilifolia

32. O. palmeri

33. Pe. sessilifolia

34, Pe. sessilifolia

36. Ps. criocarpum

37. O. pringlei

38. Pe. michoacana

39. Ps. megaphyllum

40. O. multilobum

43. O. palmeri

44, Ps. peltatum v. peltatum
45. O. sinuatum

46. D. jatrophoides

47. Ps. megaphyllum

48. Pe. sessilifolia

49. Ps. peltatum v. peltatum
40. O. sinuatum

51. Pe. sessilifolia

52. O. cirsiifolium

53. Ps. peltatum v. peltatum

54. O. silphiifolium
56. O. sinuatum
57. O. sinuatum
58. O. platylepis
59. O. pringlei

60. Ps. poculiferum
61. Pe. sessilifolia
62. Ps. eriocarpum

63. Pe. michoacana
64. Pe. sessilifolia

65. Pe. michoacana
66. Ps. eriocarpum
67. Ps. eriocarpum

Powe 1, A. M., & EpmMonpson, J.

869. O. palmeri
929. O. radulifolium
948. O. sinuatum

PRINGLE, C. G.

767. O. decompositum
1736. Pe. sessilifolia
1811. O. pringlei
1816. O. platylepis
2304. O. palmeri

“CACALIOID” GENERA 441

3340. Ps. peltatum v. peltatum

3566

4095.
4262.
4274.
4414,
4672.
4778.
4785.
4984,
5272.
5741,
6018.
6164.

6165
6176
6238
6453
7983
9080
9868
9869
9870
9871

9872.

9873
9874
9875
9877
1002
1011

. O. radulifolium

O. radulifolium

D. jatrophoides

O. sinuatum

Ps. poculiferum

Ps. peltigerum v. peltigerum
D. napeifolia

O. amplifolium

O. cirsiifolium

O. cirsiifolium

D, jatrophoides

O. paucicapitatum

D. tridactylitis

. Ps. megaphyllum

. D. heteroidea

. Ps. peltatum v. conzattii
. O. silphiifolium

. Pe. sessilifolia

. O. silphiifolium

. Ps. tussilaginoides

. Pe. sessilifolia

. Ps. peltatum v. peltatum
. Ps. peltatum v. peltatum
Ps. peltigerum v. peltigerum
. O. platylepis

. Ps. poculiferum

. O. sinuatum

. D. tridactylitis

4, O. napellifolium

7. Pe. michoacana

10126. O. brachycomum

1149
1149
1150
1329
1367
1397

7. Pe. sessilifolia
9. O. pringlei

0. O. palmeri

7. Ps. holwayanum
2. Ps. holwayanum
9. Ps. holwayanum

Purpus, Caru A,

3038. Ps. peltatum v. peltatum
3139. O. amplifolium

3845.
4113.
5625.
5630.

1935.
1986.
2101.
2248.
2337.

O. purpusii
Ps. calvum
Pe. sessilifolia
Ps. tabulare

Roses, J. N.

Ps. poculiferum
O. cervinum

O. filicifolium
cf. O. palmeri
Ps. laxiflorum

4492 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

2828. Pe. sessilifolia SHarp, A. J.

3040, D. jatrophoides 44508. Ps. peltatum v. peltatum
3471. O. globosum x O. sinuatum 441431. O. goldsmithii v. rowellii
3575, O. amplum 441465. O. amplifolium

7808. O. sinuatum
SHREVE, FORREST
Ross, J. N., & Hay, Roperr 8031. O. globosum

5311. O. silphiifolium

oes Situ, Cuarues L.
5547, O. tussilaginoides

287. D. heteroidea
Ross, J. N., & Painter, J. H. 380. D. tridactylitis
6738. O. tussilaginoides 387. D. heteroidea

7939. Ps. peltatum v. peltatum 388. D. jatrophoides
Smitu, Lucius C.

123. D. jatrophoides
9187. O. silphiifolium 124. O. amplifolium

966. D. tridactylitis
Smytu, E. G.

Ross, J. N., BT AL.

RoweE.t, CuestER M., Jr.

3387. O. goldsmithii v. rowellii ;

3604. O. goldsmithii v. rowellii 172. QO. sinuatum

3779. O. napellifolium . Sranrorp, Rernerrorp, & NorTHCRAFT
3886. O. goldsmithii v. rowellii dulifoli

3896. O. napellifolium 818. Q. radulifolium

STANFORD, TayLor, & LAUBER

2421. O, radulifolium
15974. O. amplifolium 2535, Ps. peltatum v. adenophorum
16015. O. amplifolium
y HH. T.
16073. O. goldsmithii v. rowellii _ __ Townsnnp, C.
s.n. in 1909. O. decompositum

Sarrorp, WiuiiAM E. Townsenp, C. H. T., & Barser, C. M.
1432. O. platylepis 158. O. decompositum
1451. Pe. sessilifolia
Tucker, J. M.
SaLazaRr, F, 2456. O. decompositum

s.n. Aug. 1, 1913. O. napellifolium VISCHER

RzEpOwSKI, Jd.

14. Pe. sessilifolia

SCHAFFNER, J. G, :
58. O. sinuatum

s.n. Sept. 1955. Pe. sessilifolia
255. Pe. sessilifolia WarerraLt, U. T.

294. O. radulifolium 12668. O. pachyphyllum
294. Ps. peltatum v. peltatum Wuitr, STEPHEN 8,

2741. O. decompositum
3289. O. decompositum
148. O. radulifolium 4353. O. decompositum

SEATON, HENRY Witzur, R. L. & C. R.

s.n. in 1891. Ps. peltatum v. peltatum 2017. Ps. poculiferum
2166. Ps. poculiferum

SCHUMANN, W.

SEEMANN
Witcox, Timotuy E.

s.n. O. pachyphyllum s.n. in 1891. O. decompositum
SELER, Ep. WriGut, CHARLES

1259. Ps. peltatum v. peltatum 1286. O. decompositum

PLATES

CONTR. U.S. NAT. HERB. VOL. 34, PT. 6 PIPPEN—PLATE 1

© G. PRINGLE,
PLANTAL MEXICANA.
184.
ers ~STATE OF dakaca~
a . on Cy
= eaten Livciee erties
é feb, & Sh heen TCP,
Sierra de San Fitlpe, ait eee wee
et “et Per °

ore

Digitacalia tridactylitis (Rob. & Greenm.) Pippen (type, Pringle 5841, GH).

CONTR. U.S. NAT. HERB. VOL. 34, PT. 6 PIPPEN—PLATE 2

Chester M. Rowell fr.

Date hidy. hh UR oe

Odontotrichum goldsmithit var, roveellit Pippen, var. nov. (type, Rowell 2991, MICH).

CONTR. U.S. NAT. HERB. VOL. 34, PT. 6 PIPPEN—PLATE 3

PLANTS OF SOUTHWESTERN JALISCO, MEXICO
LAPEDITION OF THE BOTANICAL GARDENS, USIVERAITY GF aCnCMIGAN

Odontotrichum multilobum Pippen, sp. nov. (type, Wilbur 2094, MICID.

CONTR. U.S. NAT. HERB. VOL 34, PT. 6 PIPPEN—PLATE 4

fei, Watson. me sp.

Odontotrichun pringler (S. Wats.) Rydb. (lectotype, Pringle 1749, GH).

CONTR. U.S NAT. HERB. VOL. 34. PT. 6 PIPPEN- PLATE 5

r—

WOES Cacelia cifelwomesnna.
i

Pericalia michoacana (B. L. Rob.) Rydb. (type, Pringle IOLL7, GH).

CONTR. U.S. NAT. HERB. VOL 34, PT. 6 PIPPEN—PLATE 6

ho Cacalia pritata, TAR.

Psacalium megaphyllum (Rob. & Greenm.) Rydb. (type, Pringle 2090, GH).

CONTR. U.S. NAT. HERB. VOL. 34, PT PIPPEN—PLATE 7

Bsagqys i.
aS Ly
om,

UNITED STATES NATIONAL MUSEUM

HERBARIUM of GEO. B. HINTON No 24464

Pamily
Name

Determined by

Locality Teotepec 3600 m,

District Mina, Gro, _ México.
Collected byHs @t al. date 7-17=39
Vernac. Name

Habitat Open stunted pine forest - on
Description ( mossy rocks,
Uses Flower white,

Psacalium nanum Pippen, sp. nov. (type, Minton M4464, US).

CONTR. U.S. NAT. HERB. VOL. 34, PT. 6 PIPPEN—PLATE 8

enlargement of Psacaltum nanum Pippen, sp. nov. (type, Hinton 14464, US).

CONTR. U.S. NAT. HERB. VOL. 34, PT. 6 PIPPEN-— PLATE 9

i

4 ” EO B HINTON N Pb

i

, ute i

P : Cacalis

4 | .
aw : :
. ye r, are 4 fy &
« ‘ ‘s f= le

& LA
J a :

lower withe

Pracalium peltigerum var. hintonti Pippen, var. nov. (type, Minton 7966, MICH).

CONTR. U.S. NAT.

Psacalium peltigerum var. latilobum Pippen,

HERB. VOL. 34, PT. 6

MICH).

; << ah y¥@

(MOTEIG

var. nov. (type, MecCorcle and Rozwell 3453

remy urg 7

PIPPEN—PLATE 10

PLANTS OF GUERRERO. MEXICO

Nerbeceoos plank unk AM ck

TOoks. Maple & green \e aves nik
» steed inSloreScence. mart

De we ind cep OM. go. La mi wW
BoOokk

Thelton Ww. NECorcle ana

Chester M. Rowell. Jr is
Su « Fiet

JJ,

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1960. Compositae, pp. 829-971. In Kearney and Peebles, Arizona Flora,
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CassinI, H.
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1827. Cacalia in Dict. Sci. Nat. (Paris) 48:459.
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1959. Sedum of the Trans-Mexican volcanic belt. Ithaca.
Cuiusius, CaRoLus (CHARLES DE L’ ECLUSE)
1601. Rariorum plantarum historiae. (Lib. 1-3); (Lib 4-5) Antwerp.
ConTRERAS Arias, A.
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1963. Manual of vascular plants.
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1960. Studies on Andean Compositae—IV. Brittonia 12:182-195.
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1950. Gray’s manual of botany, ed. 8.
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1917. Illustrierte Flora von Mittel-Europa 6 (1) :403-407. Munchen.
HeEms.Ley, W. G.
1881. Biologia Centrali-Americana Bot. 2:235-247. London.

266-836-——67——_8 443

444 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Hircucock, A. 8., and Green, M. L.
1947. Supplement species lectotypicae generum Linnaei. Brittonia 6:114-
118.
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1890. Compositae im Engler and Prantl., Die natiirlichen Pflanzen-
familien 4 (5) :225-272. Leipzig.
LANGLET, O. F. J.
1936. Nagra Bidrag till Kannedomen om Kromosomtalen inom Nym-
phaeaceae, Ranunculaceae, Polemoniaceae och Compositae. Svensk.
Bot. Tidsk. 30:288-294.
Lanjouw, J., and Star.ev, F, A.
1964. Index Herbariorum. Part 1. The Herbaria of the world, ed. 5. Inter-
national Bureau for Plant Taxonomy and Nomenclature. Utrecht.
LINNAEvs, C,
1753. Species Plantarum. Stockholm,
L’OBEL, MarrHias DE
1581. Plantarum seu stirpium icones. Antwerp.
Makino, T.
1958. An illustrated flora of Japan. Tokyo.
Martineéz, M.
1959. Plantes Medicinales de Mexico, ed.4.. Ediciones Botas. Mexico D.F.
Morison, R.
1699. Plantarum historiae. Jacob Bobart. Oxford.
Ornvurr?, R., ET AL.
1963. Chromosome numbers in Compositae. III. Senecioneae. Amer.
Journ, Bot. 50:131-139.
Raisz, E.
1959. Landforms of Mexico (map, privately published). Cambridge.
Rickert, H. W.
1955. Materials for a dictionary of botanical terms. III. Inflorescences.
Bull. Torrey Bot. Club. 82:419-445.
RypBera, P. A.
1924. Some senecioid genera. Bull. Torrey Bot. Club 51:369-378, 409-420.
Scuuutz, C. H.
1845. Ueber einige mit Senecio zu verbindende Gattungen. Flora 28:497-
498,
SuHinners, L. H.
1950. The Texas species of Cacalia. Field and Lab. 18:79-83,
TouRNEFORT?, J. P.
1700. Institutiones rei herbariae. (Cacalia on 1:451.). Paris.

INDEX

(Synonyms in ztalics. Page numbers of principal entries in boldface.)

Adenostyles, 376, 377
Aulacophora (sect. of Cacalia), 377
Cacalia, 365, 366, 375, 376, 377, 378
albicans, 434
alpina, 375, 377
amplexicaulis, 434
amplifolia, 387
ampullacea, 409, 410
atriplicifolia, 377
sect. Aulacophora, 377
berlandiert, 434
brachycoma, 388
calotricha, 434
calva, 417
cervaritfolia (‘‘cervariaefolia’’),
390
chiapensis, 379
cirsiifolia, 403, 404
sect. Cissampelopsis, 377
sect. Conophora, 377
cordifolia, 411, 413, 414
coriacea, 402
decomposita, 392
digitata, 381
eriocarpa, 418
eriopoda, 434
sect. Eucacalia, 377
globosa, 394
goldsmithii, 395, 396
guatemalensis, 434
hastata, 377
holwayana, 419
incana, 434
jatrophoides, 378, 381
laevigata, 380
liebmannit, 434
longipetiolata, 380
lyrata, 434
macrophylla, 434
megaphylla, 421, 422
michoacana, 412
napeaefolia, 382
napellifolia, 398

Cacalia—Continued
nomenclature of, 376
nutans, 413
obtustloba, 421, 422
pachyphylla, 399
pallescens, 434
palmeri, 400
parasitica, 434
paucicapitata, 401
peltata, 424, 426, 434

var. conzatti1, 427
var. coultert, 426

peltigera, 428, 430
pinnatifida, 435
platylepis, 402
pratensis, 394, 395
prenanthoides, 435
pringlez, 403
pudica, 435
purpurascens, 435
purpusit, 405
radulifolia (“radulaefolia’’), 405,406
rumicifolia, 387
runcinata, 435
sarracenica, 435
schaffnert, 406
sessilifolia, 413, 414
silphiifolia, 407
sinuata, 408, 435
spathulata, 435
suaveolens, 375, 377
subdecurrens, 435
suffulta, 435
tabularis, 432
tepicana, 435
toluccana, 435
tridactylitis, 383
trigonophylla, 412, 413
tuberosa, 435
tussilaginoides, 409, 410
viscosa, 435

cacalioid condition, 368

Chaptalia nutans, 435

445

446

Cissampclopsis (sect. of Cacalia), 377
Conophora (sect. of Cacalia), 375, 377
Digitacalia, 365, 366, 367, 368, 370, 371,
372 (fig.), 374, 377, 378, 379
chiapensis, 379
heteroidea, 379, 380, 381
jatrophoides, 369 (fig.), 879, 381, 382
napeifolia, 379, 382
tridactylitis, 379, 383
FErechtites valerianacfolia f.
thoides, 435
Kucacalia (sect. of Cacalia), 375,
Matarique, 393
Mesadenia decomposita, 392
Mesoneuris bipinnatifida, 390
Mulgedifolii (sect. of Senecio), 374, 434,
435
Multinervii (sect. of Senecio), 375
Odontotrichum, 365, 367, 368, 370, 371,
372 (fig.), 377, 378, 379, 384, 406,
410, 434
amplifolium, 386, 387, 410, 435
amplum, 367, 368, 385, 387, 400
ampullaceum, 409
bipinnatifidum, 390
brachycomum, 368, 385, 388, 389,
401
caluum, 417
cervinum, 386, 390, 394, 399
chiapensis, 379
cirsiifolium, 368, 384, 385, 390
decompositum, 366, 368, 370, 386,
392, 393
delphinifolium, 435
filicifolium, 386, 390, 393, 399
globosum, 385, 394, 395
goldsmithii, 367, 395, 396, 405, 407
var. goldsmithii, 385, 396
var. rowellii, 385, 396
jatrophoides, 381
liebmannii, 436
multilobum, 368, 386, 397, 398
napeaefolium, 382
napellifolium, 386, 398
nephrophyllum, 401
pachyphyllum, 367, 368, 385, 387,
399
palmeri, 368, 385, 389, 398, 400, 401
paucicapitatum, 386, 401
platylepis, 367, 386, 402, 404
pratense, 394
pringlei, 368, 386, 396, 403

prenan-~

377

purpusii, 385, 405

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Odontotrichum—Continued
radulifolium, 378, 386, 405
rumicifolium, 387
scabrum, 394, 395
schaffnert, 406
silphiifolium, 367, 368, 385, 396,

405, 407
sinuatum, 367, 368, 369 (fig.), 386,
395, 398, 402, 404, 408
tridactylitis, 383
tussilaginoides, 386, 409, 410
Palmatinervii (sect. of Senecio), 374, 375
Pericalia, 365, 367, 368, 370, 371, 373
(fig.), 374, 377, 378, 379, 410, 429
michoacana, 368, 411, 412, 413, 415
ovatifolia, 413
sessilifolia, 367, 369 (fig.), 411, 413,
414, 415
Psacalium, 365, 367, 368, 370, 371, 373
(fig.), 374, 377, 378, 379, 384,
415, 424, 434
argutum, 426
calvum, 368, 417, 423
conzattt1, 427
coulteri, 426
eriocarpum, 416, 418, 421, 433
holwayanum, 416, 419, 420, 421,
433
langlasset, 419, 420
laxiflorum, 417, 420
megaphyllum, 417, 421
mollifolium, 427, 428
nanum, 368, 417, 422
nelsonii, 417, 423
obtustlobum, 421
peltatum, 416, 417, 424, 425, 434
var. adenophorum, 425, 427
var. conzattii, 425, 427, 428
var, peltatum, 369 (fig.), 425,
426
peltigerum, 428, 428
var. hintonii, 429, 430
var. latilobum, 430, 431
var. peltigerum, 429, 430
poculiferum, 417, 431, 433
species, 417, 433
tabulare, 416, 432
thyrsoideum, 436

Sciadoseris vaginata, 390

Senecio, 365, 368, 371, 377, 378, 434, 435
albo-lutescens, 408
amplifolius, 387
angulifolius 369 (fig.), 374, 434

Senecio—Continued
beecheyanus, 413
bracteatus, 434
callosus, 369 (fig.), 374, 435
calophyllus, 408
cardiophyllus, 413
cervartaefolius, 390
chiapensis, 379
cirstifolia, 403
cooperi, 375
cordobensis, 374
deformans, 374, 435
desortum, 434
eriopodus, 434
farfarus, 409
grayanus, 392
hartwegii, 375, 435
heteroideus, 380
jaliscana, 374
jatrophoides, 381
liebmannii, 484
moreliae, 420

INDEX 447

Senecio—Continued
sect. Mulgedifolii, 374, 484, 435
sect. Multinervii, 375
napeaefolium, 382
ovatifolius, 413
pallescens, 434
sect. Palmatinervii, 374, 375
parasiticus, 434
peltiferus, 424
purpurascens, 374, 435
radulaefolius, 406
roldana, 374
roseus, 435
rumictfolius, 387
runcinatus, 374
sclerophyllus, 399
seemanii, 375
sessilifolius, 413
subpeltatus, 374
tabulare, 432
tussilaginoides, 409

senecioid condition, 368

U.S. GOVERNMENT PRINTING OFFICE: 1968

BULLETIN OF THE UNITED STATES NATIONAL MUSEUM

CONTRIBUTIONS FROM THE UNITED STATES NATIONAL HERBARIUM

VoLuME 34, Part 7 (END oF VOLUME)

THE LICHEN GENERA CETRELIA
AND PLATISMATIA (PARMELIACEAE)

By Wriii1Am Louis CULBERSON
and Cuicita F. CULBERSON

SMITHSONIAN INSTITUTION PRESS e WASHINGTON, D.C. e 1968

Contents

Introduction .
Historical Background
The ‘‘Parmelioid’’ Cetrariae
Morphology .
Cortex
Pseudocyphellae
Puncta
Isidia .
Soredia
Lobulae .
Apothecia .
Ascospores .
Pyenidia.
Chemistry .

Biosynthetic Dichotomy Leading to ‘Aromatic and Aliphatic Sub-

stances
Orcinol- and B-Orcinol- -type Substitutions

Length and Oxidation State of the Side Chain in Oreinol- type Phenolic

Acids . .
Depsides and Depsidones
O-Methylation .
Relationship of Cetrelia Compounds to ‘Other Lichen Substances

Methods for the Microchemical Identification of Lichen Substanees in

Cetrelia and Platismatia .
Solutions
Microextraction
Microcrystal Tests
Paper Chromatography .
Hydrolysis of Depsides
Thin-layer Chromatography .

The Identification of Specific Substances
Alectoronic Acid and a-Collatolic Acid
Anziaic Acid .

Atranorin

Caperatic Acid .

Fumarprotocetraric Acid

Imbricaric Acid

Microphyllinic Acid .

Olivetoric Acid .

Perlatolic Acid . . .

An Unidentified Substance .
Chemical Criteria in the Taxonomy of Cetr elia and Platismatia .
Acknowledgments

Til

Page
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464

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465

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469
471
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IV CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Page

Taxonomic Treatment. 2. 2. 2. 2. ee 490
Cetrelia Culb. & Culb.. . . . Co 490
Artificial Key to the Species of Cetrelia Lo Co 49]
Cetrelia alaskana (Culb. & Culb.) Culb. & Culb._ Co 492
Cetrelia braunsiana (Mill. Arg.) Culb. & Culb.. 2. 2... 2... 493
Cetrelia cetrarioides (Del. er Duby) Culb. & Culb. re 498
Cetrelia chicitae (Culb.) Culb. & Culb. . 2 2 2. 2 ee eee 504
Cetrelia collata (Nyl.) Culb. & Culb.. 2. 2 2 2. we ee 505
Cetrelia davidiana Culb. & Culb. . . 2... 1 ee ee 507
Cetrelia delavayana Culb. & Culb.. 2. 2. 2 1. 1 ee ee es 509
Cetrelia isidiata (Asah.) Culb.& Culb, 2... 2. 7 ee ee 510
Cetrelia japonica (Zahlbr.) Culb. & Culb. 2... 2... 7 ee 511
Cetrelia nuda (Hue) Culb. & Culb, . 2... 2... ee 513
Cetrelia olivetorum (Nyl.) Culb. & Culb. . 2... 2... 2 ee 515
Cetrelia pseudolivetorum (Asah.) Culb.& Culb. . 2... 2 1 519
Cetrelia sanguinea (Schaer.) Culb.& Culb.. 2. 2... 0 2 we 521
Cetrelia sinensis Culb. & Culb. . 2. 2 2 2 2 2 ee ee 523
Platismatia Culb. & Culb. . 2. . . Co 524
Artificial Key to the Species of Platismatia . re 525
Platismatia erosa Culb. & Culb.. . . . Co 526
Platismatia formosana (Zahlbr.) Culb. & Culb. Co ee 529
Platismatia glauca (L.) Culb. & Culb. . 2. 2. 2 2. 7 ee ee es 530
Platismatia herrei (Imsh.) Culb. & Culb.. . 2. 2 2 2. ee ee 537
Platismatia interrupta Culb. & Culb, . . 2 1... ee ee 539
Platismatia lacunosa (Ach.) Culb. & Culb. . 2 2 2 2. ee ee 541
Platismatia norvegica (Lynge) Culb. & Culb. . . 2... 2. we 543
Platismatia regenerans Culb. & Culb, . . 1. 1. 1 ee ees 547
Platismatia stenophylia (Tuck.) Culb.& Culb, 2... 1. 7 7 ee 548
Platismatia tuckermanii (Oakes) Culb. & Culb. . . . 1... 7. 549
Literature Cited . 2. 2... 554

Index. . 2...) ee ee ee 557

THE LICHEN GENERA CETRELIA
AND
PLATISMATIA (PARMELIACEAE)

By Witu1aM Louris CuULBERSON
and Cuicita F, CULBERSON

Introduction

In 1960 we undertook a monographic study of the old genus Cetraria
in the traditionally broad sense that it was understood in Zahlbruck-
ner’s Catalogus Lichenum Universalis. It soon became obvious that this
assemblage contained several major constellations of species and that
these species groups were not particularly closely related to each
other. One of these major groups is of course Cetraria itself, and this
genus will be the subject of a future publication. A small, homoge-
neous, arctic-alpine segregate genus named Asahinea has already been
treated (Culberson and Culberson, 1965). Two groups remain—the
new segregate genera Platismatia and Cetrelia. The aim of the present
study is to describe and delimit these genera and to present a compre-
hensive treatment of the species which belong to them.

This study has the additional objective of examining the nature of
chemical variation in morphologically well defined species groups. It
was obvious early in our research that the plants under study showed
chemical variations strikingly correlated with morphologic ones.
Moreover, the chemical compounds themselves seemed to have closer
relationships to each other than those previously noted in other
genera of approximately the same size. We decided, therefore, to
treat chemistry and morphology in equal detail in our study. Of
course, the use of chemistry in lichen taxonomy is now so common-
place that it is impossible to imagine a serious treatment of any
group showing chemical diversity in which this aspect of the variation
was left unstudied. Few lichenologists, however, have tried to in-
tegrate chemical and morphological data or to place chemical varia-
tion in the same evolutionary frame of reference in which critical
taxonomy places morphological variation. But if we are to involve
chemistry in taxonomy, an attempt, however crude, must be made to
delineate the nature of chemical variation in lichens. In this mono-
graph we present two genera to serve as subjects for the kind of

449

450 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

theoretical speculation that will lead eventually to the structuring of
a framework of principles upon which practical taxonomic judgments
can rest.

Historical Background

From the very beginning, generic concepts have been one of the most
unsatisfactory aspects of the taxonomy of lichenized fungi. Linnaeus,
in his Species Plantarum of 1753, considered all the species as
members of the single genus Lichen and, to the detriment of lichen-
ology, the insensitive Linnaean treatment was selected as the starting
point for the nomenclature of the whole group. Even Acharius, the
first botanist to see the lichens with a discerning and critical eye, began
by keeping all the species in the genus Lichen (Acharius, 1798), ob-
viously simply in deference to tradition. A few years later though, in
1803, he divided the lichens into some two dozen genera, one of them
being Cetraria.

As originally described, the genus Cetraria contained eight species,
namely:

C. islandica (L.) Ach. C. fallax (L.) Ach.

C. cucullata (Bell.) Ach. C. glauca (L.) Ach.

C. nivalis (L.) Ach. C. sepincola (Ehrh.) Ach.
C. lacunosa Ach. C. juniperina (L.) Ach.

The genus soon became a general repository for all parmeliaceous
plants with marginal or submarginal apothecia. It was used in its
Acharian sense for the species listed above and for related species by
most of the major lichenological authors of the nineteenth century, for
example, Schaerer, Fries, Tuckerman, Korber, and Massalongo.
Nylander alone among the leading lichen students of the time arranged
the species differently. He took up the almost forgotten genus Pla-
tisma described by Hoffmann in 1790—although he changed the spelling
to ‘““Platysma’’—and used it to classify all but one of Acharius’ Cetra-
rae (Nylander, 1855). In so doing, he effectively typified the genus
Cetraria Ach. with Lichen islandicus L. by leaving only that species in
it (Culberson, 1966b). Nylander’s revival of Platysma reflected his
belief that Cetraria, as conceived by other botanists of the time, was
not homogeneous.

Although all these species had marginal or submarginal apothecia
and pycnidia, it was difficult to find any other major traits shared by
even the restricted list of species that Acharius originally put into the
genus. In the original complement of species listed above, Nylander
saw the erect, linear-lobed, subfruticose, soil-inhabiting Cetraria
islandica as an element too foreign from the other more distinctly
foliose species to be classified in the same genus with them. Nylander,
however, apparently did not question the generic affinity of all the
foliose species that he recognized in Platysma. He saw no obstacle to

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 45]

classifying the broad-lobed, Parmelia-like Cetraria glauca and the
narrow-lobed, erect, soil-inhabiting species such as the familiar C.
cucullata and C. nivalis in the same genus.

As time passed, other workers described many more species in
Cetraria, and the generic concept became more nebulous still. There
were species with marginal, reflexed apothecia, such as Cetraria
strachey: Bab., that in some later classifications were transferred to the
genus Nephromopsis Miill. Arg. There was also a whole array of very
broad-lobed forms, most of them foreign to Europe and superficially
resembling the Amphigymnia Parmelias, for example Cetraria sangui-
nea, C. collata, and C. japonica from the Orient. In addition, a few
particularly anomalous species, such as C. chrysantha Tuck. and C.
thomson Stirt., were put in Cetraria in spite of their lacking any con-
nection whatever with the bulk of the species already classified there,
except for having marginal apothecia.

The Catalogus Lichenum Universalis (Zahlbruckner, 1930) took a
particularly broad view of Cetraria and included all the plants men-
tioned above and most of their obvious relatives, excluding only a few
of the species as members of the genus Nephromopsis. Zahlbruckner’s
Catalogus had the effect of rigidly formalizing these generic concepts.
With the exception of this study, subsequent treatments have either
followed him in recognizing Cetraria and Nephromopsis (Riasénen,
1952) or have taken a still more conservative position and recognized
only Cetraria (Rassadina, 1950). Only Ejilif Dahl tried to evaluate the
major variations among the plants that had been referred to Cetraria
and to define the major natural groups in an assemblage of species that
years of casual taxonomy had left biologically meaningless.

The “Parmelioid” Cetrariae

Dahl challenged the monophyletic nature of Cetraria sens. lat. He
pointed out that Cetraria glauca (L.) Ach., C. norvegica (Lynge) Du
Rietz, C. collata Miill. Arg., and C. chrysantha Tuck. differed from the
true Cetrarias both morphologically and chemically. The cortex in
these species, he demonstrated, is composed of densely agglutinated
hyphae with tiny lumina, a type of cortex that he called “platysmoid”’
but which is more commonly called prosoplectenchymatous (Ozenda,
1963). The upper cortex of true Cetrarias, on the other hand, is com-
posed of thin-walled hyphae with inflated lumina, a type generally
termed paraplectenchymatous. These anatomical differences were
correlated with the distribution of some lichen substances. The dozen
true Cetrarias that Dahl analyzed chemically were characterized by a
predominance of usnic acid and fatty acids while the species with a
prosoplectenchymatous cortex were characterized by a predominance
of atranorin. On the basis of this morphological and chemical informa-
tion, Dahl postulated that Cetraria sens. lat. was polyphyletic. He

452 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

segregated the species with the prosoplectenchymatous upper cortex
and the presence of the substance atranorin from the true Cetrarias
and used the old genus Platisma Hoffm. to accommodate them.

When we started our study of the lichens traditionally assigned to
Cetraria, we made a preliminary survey to determine the general range
of the morphological and the chemical variation. It was soon obvious
that the difference that Dahl had pointed out between the “true”
Cetrarias and the members of the Cetraria glauca group was indeed a
basic one, and it held up as even more and more species were con-
sidered. We thereupon decided to put aside the true Cetrarias and to
study first the broad-lobed species that had traditionally been classi-
fied with them, a group which for convenient reference we can call the
“narmelioid”’ Cetrariae.

The “parmelioid”’ Cetrariae include the Cetraria glauca group as well
as such species as C. sanguinea, C. japonica, and C. collata. These
latter Asiatic species all have pseudocyphellae, a trait present in some
but not all members of the C. glauca group. Although the position of
C. sanguinea and C. japonica, for example, had never been seriously
disputed, apparently no one had noticed that these species were
extremely closely related to other species that have always been con-
sidered Parmelias, for example Parmelia cetrarioides and P. olivetorum.
(But the superficial similarity of P. cetrarioides Del. to the Cetrarias
was pointed out from the very beginning by Delise in his choice of the
specific epithet.) A species which several years ago was (erroneously)
being called Cetraria collata in North America is so similar to Parmelia
cetrarioides that the two species can be separated from each other with
certainty only by chemical tests. Although we were unsure of the
taxonomic disposition which should be made of all these species, it
was clear that P. cetrarioides and P. olivetorum were very closely re-
lated to at least some of the “parmelioid’” Cetrariae and that they
should be classified with them and not among the true Parmeliae.

Our knowledge of the “parmelioid”’ Cetrariae was summed up when
we compared these plants, the true Cetrarias, and the Amphigymnia
Parmelias to three very peculiar, distinctive, and closely related
species which had been classified in Cetraria and which we defined as
the new genus Asahinea (Culberson and Culberson, 1965). The traits
of the “parmelioid” Cetrariae and the three other species groups were
summarized then in a form which we present here with only slight
modifications (Table 1).

But the ‘“parmelioid”’ Cetrariae are themselves not a homogeneous
group. From the outset it was clear that the bulk of the “parmelioid”’
Cetrariae really consists of two distinct constellations of species rather
than of a single spectrum of forms. Although from Dahl’s treatment we
might have expected these species to be characterized by the presence
of fatty acids as the medullary lichen substances, only some of the

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 453

TABLE 1.—A comparison of the Major traits in Parmelia sect. Amphigymnia, Asahinea, the ‘‘parmelioid’’
Cetrariae (=Cetrelia+Platismatia), and Cetraria sens. str.

Parmelia sect. ‘“‘Parmelioid”’ Cetraria
Amphigymnia Asahinea Cetrariae sens. str.
(106 species) (3 species) (24 species) (about 75
species)
Cortex prosoplecten- prosoplecten- —_ prosoplecten- _—paraplecten-
chymatous chymatous chymatous chymatous
Rhizines present absent present present in
most
Marginal present in absent absent present in
ciliae many some
Pycnidia laminal laminal or marginal marginal
marginal
Apothecia laminal laminal or laminal or marginal
marginal submarginal
Disc perforate in imperforate perforate in imperforate
many most
Lichen sub- many; aro- 4 aromatic many; aro- many; aro-
stances matic and matic and matic and
aliphatic aliphatic aliphatic
Distribution nearly world- Northern Northern Northern
wide Hemisphere, Hemisphere, Hemisphere,
arctic- mainly arctic-
montane temperate temperate

species were actually found to contain fatty acids. More than half the
species produced (as medullary constituents) one or more aromatic
compounds, orcinol-type depsides (anziaic, imbricaric, microphyllinic,
olivetoric, and perlatolic acids), or orcinol-type depsidones (alectoronic
and a-collatolic acids). All the species producing aromatic compounds
exclusively as the medullary substances belong to the ‘Parmelia
cetrarioides’’ group while those producing a fatty acid (invariably
caperatic acid) as the medullary constituent all belong to the ‘‘Cetraria
glauca” group. The chemistry of the species of the two groups is sum-
marized in Table 2. An analysis of the chemical relationships of the
compounds in each group is given in a discussion farther on.

The two groups into which the species fall by similarity of general
habit as well as by chemistry of the medullary substances are marked
by suites of additional morphological and chemical traits (Table 3).
Internally, each of the two groups of the ‘“parmelioid’’ Cetrariae is
extremely uniform and yet each is so distinct from the other that they
cannot be considered congeneric. We propose the genus Platismatia for
the 10 members of the Cetraria glauca group and Cetrelia for the 14
members of the Parmelia cetrarioides group.

As characterized here, the genera Cetrelia and Platismatia both show
relatively conservative variation and extremely high levels of internal
homogeneity. The chemical variation here is far more precisely cor-

454 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

TABLE 2.—The distribution of the constant chemical substances in the species of Cetrelia and Platismatia and
the number of specimens tested of each species. The few unidentified ‘‘accessory” substances found are
not tabulated

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CETRELIA

alaskana + | + 14
braunsiana of + + 32
cetrarioides sens. str. + | | + | 35
cetrarioides sens. lat. + fy | 34
chicitae + + + | 24
collata + | + | | 17
davidiana + | + | 7
delavayana + | | | | + 8
isidiata > | + | 4
japonica + | + | 22
nuda + + + | 22
olivetorum + ot | 19
pseudolivetorum + | + 31
sanguinea + | + | | 67
sinensis | + | | - | | | 6

PLATISMATIA 1 | |
erosa +) + | | | 17
formosana + | + | | | | | | 2
glauca | + |} 4+ | | ul
herret b+} + / | | | | | | 5
interrupta +) + | | | | | — 20
lacunosa +) +) + | 8
norvegica | + } + | | | | | 7
regenerans p+ |] +. 4
stenophylla | +) 4) | | | | | 5
tuckermanti + | + | | | 7

related with visible traits (Table 3) than in most genera of the macro-
lichens. All the species of Platismatia produce the aliphatic fatty acid
caperatic acid as the medullary constituent and only one species
produces a B-orcinol-type depsidone (fumarprotocetraric acid) in
addition to the aliphatic substance. The species of Cetrelia, on the
other hand, never produce fatty acids as the medullary constituents
but only aromatic substances—orcinol-type depsides and depsidones,
substances all rather closely related to each other chemically.

One might wonder whether the genera Cetrelia and Platismatia, as

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 455

TABLE 3.—A comparison of the major traits in Platismatia and Cetrelia, the two genera constituting the “‘parme -
lioid”’ Cetrariae

Cetrelia Platismatia
(=the Parmelia cetrarioides group) (=the Cetraria glauca group)
Upper surface always pseudo- Upper surface usually pseudo-
cyphellate cyphellate
Soredia rare Soredia common
Isidia common Isidia rare
Lobulae absent Lobulae common
Upper cortex always IKI— Upper cortex often IKI+
Subhymenium thick, IKI— Subhymenium thin, IKI+
Thick zone of algae usually present Algae usually absent from im-
immediately below subhymenium mediately below subhymenium
Spores large, ellipsoid, 11-22 * 6-124 Spores small, subspherical, 5-8 X 3-5
Pycnidia common in most species Pycnidia rare in most species
Pycnoconidia with inflated ends Pycnoconidia without inflated ends
Medullarly chemistry: substances all Medullary chemistry: the aliphatic
aromatic orcinol-type depsides and substance caperatic acid in every
and depsidones species (one species also with a
depsidone)
14 species 10 species
Greatest species concentration in Greatest species concentration in the
eastern and southeastern Asia amphi-North Pacific

delimited by the morphological and chemical criteria explained above,
show phytogeographically meaningful distributional patterns, since
such distributions are expected in monophyletic genera. While many
parts of the world remain inadequately explored lichenologically, the
plants that we are concerned with here are large and conspicuous and
have often been collected, even by amateur plant hunters. Their
general conspicuousness and the frequency of their collection give
one a confidence in the significance of maps of their distribution that
one would not have for smaller, less noticeable cryptogams. In Figure 1
the number of species of Cetrelia and of Platismatia in floras in various
parts of the world is plotted. It is obvious from these maps that the
genera have very different centers of maximum species numbers
although both genera are circumpolar. Platismatia is represented
most abundantly in the amphi-North Pacific, especially western
North America. Cetrelia, on the other hand, is most richly represented
in Kast and Southeast Asia. From this phytogeographic evidence it
would seem indisputable that Cetrelia and Platismatia had dif-
ferent centers of origin. It is also noteworthy that neither genus is
represented in Australia or New Zealand. Cetrelia is not present in
Africa, and Platismatia is represented there only by a few isolated
mountain stations for the ubiquitous Platismatia glauca. The genus
Cetraria (in the strict sense) likewise does not exist in Africa but
Parmelia sect. Amphigymnia (Hale, 1965) is richly represented there.
Cetrelia is also absent from South America, and Platismatia is repre-

456 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

PLATISMATIA

Figure 1. The distribution of the genera Cetrelza (above) and Platismatia (below)
by numbers of species in floras in various parts of the world.

sented only in the southern part of that continent and there again by
P, glauca alone. Both Cetrelia and Platismatia are temperate, Northern
Hemisphere genera, but their different centers of origin and overall
ranges clearly indicate that the distinctive morphological and chemical
traits that characterize them are the product of different and in-
dependent evolutionary histories.

Morphology

The following morphological characteristics have proved to be the

most useful in the study of the species of the genera Cetrelia and
Platismatia:

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 457

CORTEX

The genera of macrolichens, with a few exceptions such as Collema,
have a true cortex. The cortex is recognizable as the outermost part of
the thallus and shows a greater range of anatomical variation than any
other vegetative tissue. It may consist of hyphae that are essentially
free, erect, and at right angles to the surface of the thallus (e.g.,
Roccella), or interwoven and parallel to the surface of the thallus (e.g.,
Anaptychia), or densely agglutinated throughout. If the hyphae are
agglutinated, they may be either inflated and have large lumina
(paraplectenchyma)—as in Peltigera, Leptogium, or Sticta—or they
may be very little if at all inflated and have tiny lumina (prosoplecten-
chyma)—as in Ramalina and Parmelia. The various cortex types have
been summarized by Ozenda (1963). In general the major cortex varia-
tions closely follow generic lines and the members of a single genus
generally all tend to show the same cortex type.

Dahl (1952) pointed out that the species of true Cetrariae which he
studied have a paraplectenchymatous cortex and that the species
which he assigned to the genus Platysma have a prosoplectenchymatous
cortex. This distinction between the true Cetrariae and the so-called
“parmelioid” Cetrariae has held up even after all the known species
have been examined. It is a basic difference of fundamental importance
between the “‘parmelioid” Cetrariae—the genera Platismatia (Plate 8,
Figures 43-44), Cetrelia (Plate 8, Figure 42), and Asahinea (Plate 7,
Figure 40)—and the true Cetrariae (Plate 7, Figures 37-38). In its
anatomy, the upper cortex of Cetrelia and Platismatia is very similar
to that of the species of Parmelia sect. Amphigymnia (Plate 7, Figure
39) and indeed to that of the whole genus Parmelia, it would seem,
although a detailed study of cortex structure throughout Parmelia has
never been made. The importance of cortex anatomy in defining the
“parmelioid” Cetrariae as distinct from Cetraria is so great that these
species would be considered isolated from Cetraria even without the
accompanying, correlated evidence based upon morphology and
chemistry (Table 1). But the fact that Cetrelia and Platismatia
actually have a large series of other traits separating them from the
true Cetrariae merely underlines the basic and deep-seated nature of
their taxonomic isolation from Cetraria.

Although cortex structure clearly separates Cetraria sens. str. from
Platismatia, Cetrelia, and Asahinea, it shows no variations of major
consequence among these three genera. The upper cortex in Platis-
matia and Cetrelia ranges from about 15 to 30u thick and the lower
cortex from about 10 to 30g thick. The lower cortex of these genera
bears rhizines here and there although naked areas may be extensive.
In producing rhizines from the lower cortex these genera differ strik-

458 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

ingly from Asahinea which does not (Culberson and Culberson, 1965).
In addition, although not strictly a cortex trait, a reddish-purple pig-
ment is also generally produced in the lower part of the medulla in
Asahinea, especially in moribund zones near the lower cortex. It is
notable that such pigmentation never occurs in Cetrelia or Platismatia.

PSEUDOCYPHELLAE

Pseudocyphellae are small pores in the cortex through which, if they
are big enough, the medulla is directly visible. The upper cortex of all
the species of Cetrelia and of about half the species of Platismatia is
pseudocyphellate.

In Cetrelia there are two kinds of pseudocyphellae, the small type
(Plate 12, Figure 52) and the large type (Plate 12, Figure 73). Most
species have pseudocyphellae of the small type. These little apertures
are less than a millimeter wide and punctiform or irregular in shape.
Cetrelia cetrarioides, C. chicitae, and C. braunsiana, for example, all
have small pseudocyphellae. Large pseudocyphellae, found in C. nuda
and (©. collata, exceed one millimeter in width and often appear to
result from the confluence of smaller pores. In the species producing
large pseudocyphellae, many specimens show at least some pseudo-
cyphellae of the small type as well.

The distinction between small and large pseudocyphellae was espe-
cially valuable in distinguishing Cetrelia chicitae and C. nuda in Japan.
Table 5, under the treatment for C. braunsiana (p. 495), shows the
distribution of pseudocyphellae of these two types as compared to the
distribution of other morphological traits among these three related
species in Japan.

PUNCTA

Perforations found in the lower cortex of some of the species may
have distinct taxonomic significance. These pores may not all be of
similar origin, and those in Platismatia appear to be somewhat dif-
ferent from those in Cetrelia. To stress the fact that the exact nature
of these peculiar pores or spots in the lower cortex is not clear, we call
them puncta rather than pseudocyphellae.

Puncta are best developed in the lower cortex of Platesmatia erosa
(Plate 11, Figure 51) and P. formosana where they furnish a valuable
trait for distinguishing the former species from the closely related P.
interrupta and the latter species from the morphologically somewhat
similar species P. lacunosa. Other members of Platismatia lack these
conspicuous interruptions in the lower cortex altogether.

A number of species of Cetrelia also shows similar spotlike modifica-
tions of the lower cortex. The species of Cetrelia which produce puncta

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 459

do not do so invariably, however, and in this genus the tendency to
produce a punctate lower cortex has little taxonomic value. Cetrelia
cetrarioides, C. collata, C. davidiana, and C. isidiata all consist of indi-
viduals that show at least some puncta in the lower cortex and others
that do not, and the occurrence or absence of the trait does not seem to
be correlated with any other variations. On the other hand, some
Cetrelia species—for example C. alaskana, C. braunsiana, and C.
sanguinea—never produce a punctate lower cortex.

In Cetrelia some of the puncta are actual pores in the lower cortex
and are apparently equivalent to the well developed ones seen in some
species of Platismatia. But in Cetrelia the lower cortex may also show
pale flecklike spots and every intergradation between these spots and
the true punctate pores. It may be that puncta originate as spots and
finally become an actual hole in the lower cortex. In general, though,
the abundance of the spots is greater in individuals of such species as
Cetrelia cetrarioides than is the abundance of true perforations.

In examining the lower cortex of Cetrelia species for puncta, one often
sees tiny holes in the cortex that are wounds which were made when
the specimen was removed from its substrate. Careful observation will
distinguish such irregular, sharp-edged wounds from the punctate
pores and the associated spotting discussed above, both of which are
entirely natural attributes of the plants. Puncta in the lower cortex of
Parmelia (=Cetrelia) olivetorum and P. cetrarioides were observed by
Gyelnik (1933), who was so impressed by these ‘‘pseudocyphellae”’
that he described two new species, Pseudoparmelia aradensis and P.
pseudofallax, based upon them. A year later he (Gyelnik, 1934) reduced
Pseudoparmelia pseudofallax to a form of Parmelia (= Cetrelia) cetra-
rioides after he discovered that these ‘‘pseudocyphellae” were common
and widespread in the latter species. Later still, in a study of the puta-
tive genus Pseudoparmelia, Santesson (1942) showed that the ‘‘pseudo-
cyphellae” in Pseudoparmelia cyphellata Lynge, the only species in the
genus prior to Gyelnik’s two additions, were nothing but wounds
where the lower cortex had been pulled off with the rhizines at the time
of collection. Santesson assumed that the ‘“‘pseudocyphellae” in the
lower cortex of specimens assigned to this genus by Gyelnik were
analogous to those which he described and illustrated in the South
American species P. cyphellata. But from Gyelnik’s descriptions it is
clear that the ‘“‘pseudocyphellae” which he saw in the European plants
that he studied were puncta, as defined in the present study, and not
wounds of casual origin.

ISIDIA

Isidia are rare in Cetrelia but relatively common in Platismatia. In
Cetrelia only C. braunsiana (Flate 13, Figure 55) and C. isidiata are truly

460 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

isidiate. In both these species, however, some individuals are almost
devoid of isidia altogether.

In an occasional plant of some ordinarily lobulate species, such as
C. pseudolivetorum or C. sinensis, some of the thallus appendages or
lobulae along the margins of the lobes may be so reduced in size that
they do not show the dorsiventrality typical of lobulae but resemble
isidia instead. These species should properly not be considered isidiate.

In the genus Platismatia the species P. norvegica, P. erosa, and P.
herrei are regularly isidiate as are many individuals of the polymorphic
P. glauca. Even in the habitually isidiate species, nonisidiate individ-
uals are sufficiently common to be found with some regularity and, as
one would guess, the extent of isidia production in isidiate individuals
varies greatly. Also, in some sorediate species the soredia become con-
solidated into structures that are distinctly isidioid. Platismatia inter-
rupta is a particularly variable species in which the soredia may be-
come isidioid ; some populations of P. glauca also show this tendency.

SOREDIA

Soredia are common in Cetrelia but relatively rare in Platismatia—
essentially the reverse of the situation for isidia in these genera. The
members of the Cetrelia cetrarioides group, with the single exception of
C. alaskana, are all sorediate. The soredia are very fine and powdery
and are produced, usually in abundance, at the edge of the lobes in
continuous marginal soralia (Plate 13, Figure 54). The soralia are often
so small that they can be seen easily only with some magnification.

In Platismatia there are no species in which soredia production is an
invariable diagnostic trait. Soredia are produced, however, in some
specimens of P. glauca and in many of P. interrupta. In these species
the soredia often become isidioid, especially in the older parts of the
thallus. In Platismatia the soredia are granular and laminal rather than
minute and marginal as in Cetrelia.

It should be mentioned that an occasional specimen of certain
species of Platismatia, such as P. norvegica, shows the development of
conspicuous, capitate soralia on the upper surface of the lobes. Soralia
of this sort are so infrequent yet so different from the ordinary soredia
in both Platismatia and Cetrelia that they do not seem to be a normal
thallus development at all. They are so anomalous that we regard
them as a pathological abnormality and give them no taxonomic
significance.

LOBULAE

Lobulae are minute, dorsiventral lobes on the upper surface and the
margins of the thallus. In some specimens they are produced in such
abundance on the surface of the thallus that the plants appear to be

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 461

cristate while in others they are more prominent on the margins so
that the thallus appears to be elegantly fringed with tiny lobes. As
we characterize them, lobulae are found in the genus Cetrelia but not
in Platismatia. In Cetrelia they are restricted to the C. japonica com-
plex of species, which includes C. sinensis and C. pseudolivetorum as
well as C. japonica (Plate 17, Figure 63). Since both lobulae and
isidia are similar in that they are entirely corticate and have an algal
zone and a medulla, it is not surprising that they seem to intergrade
in form in some specimens. In C. pseudolivetorum, for example, many
of the lobulae produced are so small and narrow that they no longer
show a distinct dorsiventrality and consequently closely resemble
isidia. Since no entirely and truly isidiate race of C. pseudolivetorum
is known, however, it would seem that the isidialike structures
sometimes encountered in this species would better be interpreted as
extremely reduced lobulae.

In Platismatia there are no structures analogous to the lobulae of
Cetrelia. In fact, the only modified appendages produced at all are the
special, branched, coralloid structures that develop from the expanded
thallus lobes in some specimens of Platismatia glauca (Plate 21, Figure
70) and, more rarely, in P. herrei. These fruticose structures, however,
are quite different from the tiny foliose lobulae produced by the species
of Cetrelia discussed above.

APOTHECIA

In both Cetrelia and Platismatia the apothecia (Plate 10) are usually
produced on the upper surface of the lobes and appear less frequently
at the margins. In the position of the apothecia, both genera differ
sharply from Cetraria in which the apothecia are strictly marginal.

With the exception of a few species, apothecial production in Platis-
matia is very infrequent. In Cetrelia, too, apothecia are very rare,
except in the C. collata group. In both genera the infrequency of
apothecial production contrasts strongly with the situation in Cetraria
where apothecia are generally very common. The rarity of apothecia in
the genera studied herein, however, recalls the Amphigymnia Parmelias
in which the total lack or at least high infrequency of apothecia is
typical of many species.

The only species of Cetrelia in which apothecia are produced with
high regularity is the collata group—the species C. sanguinea, C. davi-
diana, C. delavayana, and C. nuda, in addition to C. collata. In Japan,
83 percent of the herbarium specimens of C. nuda were found to bear
apothecia, and C. sanguinea in Java also often fruits. Cetrelia collata,
(. davidiana, and C. delavayana are known in fruit, but these Chinese
species have rarely been collected and the overall frequency of
apothecia in them is impossible to assess.

277-920 O - 68 - 2

46§2 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

In the Cetrelia cetrariordes complex—including C. cetrarioides, C.
alaskana, C. ridiata, C. chicitae, C. olivetorum, and perhaps C. braunsi-
ana—only C. chicitae has been found in fruit with anything approach-
ing commonness. In Japan about 17 percent of the plants are fertile,
but in North America apothecial production in this species is much
rarer. Although apothecia are known for C. cetrarioides, C. olivetorum,
and C. braunsiana, they are very uncommon. In C. cetrarioides, in
fact, only a single specimen bearing apothecia with apparently mature
spores was seen 1n this study; and in C. alaskana and C. isidiata apothe-
cia have never been observed at all.

In the C. gaponica group, C. japonica and C. pseudolivetorum are
very rarely known to fruit, while the other member of the complex, C.
sinensis, has never been found with apothecia.

In Platismatia, apothecia are also very rarely produced, with two
notable exceptions. The two North American species P. tuckermanii
and P. stenophylla commonly fruit and almost every well developed
specimen has apothecia. Two species, however, have never been found
in fruit—P. interrupta, a common species in Japan, and P. regenerans,
an apparent endemic of North Borneo. Apothecia in all the remaining
six species of Platismatia—P. erosa, P. formosana, P. glauca, P. herret,
P. lacunosa, and P. norvegica—are known, but their occurrence is rare.

Many of the apothecia of both Cetrelia (Plate 11, Figure 50) and
Platismatia become perforate, especially with age. In this trait they
recall the species of Parmelia sect. Amphigymnia in which perforate
apothecia are common. The true Cetrariae, however, have imperforate
apothecia.

Internally, the apothecia of Platismatia and Cetrelia differ in a
number of ways. In most species of Platismatia the subhymenium is in
direct contact with the medulla while in Cetrelia a thick layer of algae
separates the subhymenium from the medulla. The subhymenium of
Platismatia and that of Cetrelia seem to differ markedly with regard to
the products which make up the hyphal cell walls and, as a result, they
also differ with regard to the color reactions that they exhibit with
the IKI reagent. In Cetrelia the subhymenium remains uncolored in
the presence of IKI, but in Platismatia it becomes some shade of
purple, violet, lavender, or blue. There is, incidentally, no exception
to this general difference. These reactions of the subhymenium,
however, should not be confused with the dark blue reaction with
IKI of the asci in the hymenium. This latter reaction is very wide-
spread, probably universal, in both genera.

ASCOSPORES

Spores in the Parmeliaceae generally have been of very little system-
atic use. In Platismatia and Cetrelia, however, there is a very clear

CETRELIA AND PLATISMATIA 463

CULBERSON & CULBERSON

distinction in the spores at the generic level. In Platismatia (Plate 9,
Figure 46) the spores are small and short-ellipsoid, ranging in length
from 5 to 8u and in width from 3 to 5y, with many of the spores being
essentially subglobose. In Cetrelia (Plate 9, Figure 45), however, the
spores are twice as large as those of Platismatia and range in length
from 11 to 22u and in width from 6 to 12u. The large, more or less
thick-walled spores of Cetrelia are very similar to those of the Amphi-
gymnia Parmelias and less like the much smaller spores of the true
Cetrarias.
PYCNIDIA

Like the apothecia, the pycnidia in the two genera are infrequent in
general, only a very few species producing them with appreciable
regularity. In Cetrelia, in fact, pycnidia are unknown altogether in
five species, namely, C. alaskana, C. cetrarioides, C. isidiata, C. olive-
torum, and C. pseudolivetorum. Pycnidia are known, but are infrequent
to rare, in C. braunsiana, C. chieitae, C. japonica, and C. sinensis. Only
in the C. collata complex do they appear to be common, although some
of the Chinese species in this group—C. collata, C. damdiana, and C.
delavayana—have been collected too infrequently to permit generali-
zations. Cetrelia nuda (Plate 10, Figure 49), the commonest member of
this group in Japan, showed pycnidia in 100 percent of the specimens
examined in Japanese herbaria. It will be recalled that it is precisely
the C. collata complex, of which C. nuda is a member, which is the only
segment of the genus Cetrelia regularly bearing apothecia. A high pro-
duction of apothecia and a high production of pycnidia are very closely
correlated in this group. Of course, the explanation could simply be
that these species have a natural tendency to produce both these
structures. Nevertheless, one is tempted to assume an intimate rela-
tionship in the close correlation of apothecia and pycnidia. If pycnidia
and conidia (the pycniospores) which they produce were functioning
here in sexual reproduction, then the high frequency of pycnidia in
plants also bearing apothecia would be explained.

In Platismatia the situation is the same. The only two species in
which pycnidial production is frequent are Platismatia stenophylla and
P. tuckermanit. These are also the only two species of the entire genus
which produce apothecia with great regularity. In P. erosa, P. regen-
erans, and P. interrupta, pycnidia are unknown altogether. Platismatia
interrupta is noteworthy in this regard since it also has never been
found with apothecia in spite of its being common in many parts of
Japan. The other five species of Platismatia—P. formosana, P. herret,
P. lacunosa, P. norvegica, and P. glauca—all produce pycnidia very
infrequently. Pycnidia are so rare in P. glauca, in fact, that none was
seen in the hundreds of specimens of this common species examined
in this study.

464 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

In both Cetrelia and Platismatia the pycnidia are produced on the
margins of the lobes and not on the upper surface. In position, then,
the pycnidia of both genera are similar to those of Cetraria.

The conidia in both genera are rod-shaped and small and measure
about 1 by 4 to 5y. Conidia in Cetrelia seem to be produced in the
pycnidia with a higher frequency and in greater abundance than in
Platismatia where pycnidia, seemingly without conidia, are more com-
mon. It also appears to be a consistent difference between the two
genera that mature conidia in Cetrelia have inflated ends while those in
Platismatia do not.

Chemistry

Of the ten substances identified in the species studied, a fatty acid
(caperatic acid) and a £-orcinol-type depsidone (fumarprotocetraric
acid) occur only in Platismatia, seven orcinol-type depsides and depsi-
dones occur only in Cetrelia, and a B-orcinol-type depside (atranorin)
is found in all species of both genera (Table 2). In this section an
attempt will be made to suggest how these substances are related to
each other. The discussion will be based primarily upon a comparison
of the structures of these compounds in the light of what is known
about biosynthetic pathways.

All the compounds involved here are derived from acetate and
malonate units. An early branching point in the biosynthetic pathway
(Figure 2) leads either to aliphatic compounds like the fatty acids or
to: aromatic substances like the phenolic acids. Phenolic acids may be

ACETATE MALONATE
CONDENSATIONS
|

ALIPHATIC AROMATIC
fatty acids
A-ORCINOL-TYPE ORCINOL- TYPE
Oxidative Oxidative
Coupling \ Esterification Coupling
[ dibenzofurans | | B-orcinol depsides | |orcinol depsides | [| dibenzofurans |

Oxidative Coupling

A-orcinol depsidones | [ orcinol depsidones |

FicurE 2, A generalized diagram showing postulated biosynthetic pathways in the
elaboration of several major categories of taxonomically important lichen
substances.

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 465

substituted by one of two patterns: namely, orcinol-type compounds
like the medullary substances in Cetrelia or B-orcinol-type compounds
like atranorin and fumarprotocetraric acid in Platismatia. Simple
orcinol- and £-orcinol-type phenolic acids are esterified to depsides,
and some depsides are cyclized to depsidones by oxidative coupling.
They may also be methylated, chlorinated, hydroxylated, or decar-
boxylated (not shown in Figure 2). Various combinations of these
relatively few reactions lead to most of the known lichen depsides
and depsidones. Direct oxidative coupling of the phenolic acids can
give dibenzofuran derivatives known from some species of lichens.

BIOSYNTHETIC DICHOTOMY LEADING TO AROMATIC AND
ALIPHATIC SUBSTANCES

All the substances involved in this study appear to be of acetate-
malonate origin (Shibata, 1965). Biosynthesis of fatty acids from
acetate and malonate units has been the subject of very extensive
studies showing formation of straight-chain aliphatic acids by linear
condensations of malonate units after an initial condensation with an
acetate unit (Richards and Hendrickson, 1964). Figure 3 presents a
simplified version of the condensation of acetate and malonate units
with reduction of the carbonyl groups to give a simple, saturated, fatty
acid (left side of the figure). Caperatic acid has a slightly more complex
structure, with three carboxyl groups (one as a methyl ester) and at
least a superficial resemblance to citric acid.

Mosbach (1961, 1964) recently described the biosynthesis of the
aromatic substance orsellinic acid from acetate and malonate units,
but in this process an oxidized form of the chain of condensed units
must undergo cyclization to produce the aromatic-ring system. Figure
3 also shows how one acetate and three malonate units could cyclize to
orsellinic acid (right side of the figure). The carbonyl groups which are
reduced during fatty-acid synthesis are required in aromatic synthesis
to provide the double bonds in the aromatic ring—one of them by the
loss of water in the cyclization step and two of them by enolization.
Since the carbonyls are on alternate carbon atoms in acetate-malonate-
derived carbon chains, meta orientation of phenolic groups is observed
in the aromatic products.

ORCINOL- AND £-ORCINOL-TYPE SUBSTITUTIONS

An aromatic ring substituted with one alkyl group and two phenolic
groups, such as orsellinic acid, is related to orcinol, and a ring with an
additional carbon substituent para to the first is related to B-orcinol
(Figure 4). Phenolic units in lichen substances are generally substi-
tuted according to one of these two systems. The biosnythetic origin

466 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

CH,COOH + CH,COOH
COOH

Va condense

4 -CO»s
°) FH
CHC CH,COOH CHa Cay
O-G. {CH,COOH
~ CHa C™
\
reduce
CH,CH,C H,COOH
condense and CH,
reduce n-1 times
fo) COOH
CH4(C Ha )5,COOH O
A FATTY ACID
CHy(CH,))3CH COOH CH
HO-C-COOH > CHp HO COOH
CH,COOH OH
CAPERATIC ACID ORSELLINIC ACID

Figure 3. A comparison of the biosynthesis of aliphatic acids (on the left) and
aromatic phenolic acids (on the right) from acetate and malonate units. (Caper-
atic acid, the medullary constituent of all species of Platismatia, is an example of
a typical lichen fatty acid.)

of the extra carbon between the hydroxyl groups in 6-orcinol systems
is unknown.

If the 8-orcinol ring arises by direct carbon substitution onto a pre-
formed orcinol-type ring, then B-orcinol substances would be derived
from orsellinic acid by pathway ‘‘1” in Figure 4. For the complete
synthesis of a depside such as atranorin, several more steps would be
required including oxidation to an aldehyde and two esterification
reactions. It is also possible to imagine a branched carbon chain
which would produce the g-orcinol unit directly by cyclization (Figure
4, pathway ‘2”’). Such a branched chain could form if a propionic
acid unit replaced a malonic acid unit in the original condensation.
Similar suggestions have been advanced to explain the synthesis of
certain aromatic compounds occurring in other groups of plants (for

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 467

CH

3
0G)
OH

orcinol

R R

10-(O)-coo{O)-coot

OH OH

orcinol-type depside

CH,
HO
CH, OH
frorcinol
CH, CH
HO coo COOCH,

OHC OH CH,OH

-orcinol-type depside
(atranorin)

OOCCH=CHCOOH
CH3 CH ,OH

wo-{Oy-co0{O) coor
OHC fo) CH,

-orcinol-type depsidone
(fumarprotocetraric acid)

@

CH;
U
_ poary We)
=C° HCOOH
“CH —CS
CH, fe)
II
?
Q C-CH,
a “OH
ten,

FicureE 4. A comparison of the structures of orcinol- and 8-orcinol-type compounds
and two possible biosynthetic pathways (numbered “1” and ‘‘2’’) leading to
8-orcinol units. The precursor in pathway ‘2’’ could cyclize by pathway ‘‘3” to
the phenolic unit from which usnic acid is derived.

a discussion, see Whalley, 1963). Perhaps it is not just a coincidence
that the same branched carbon chain, cyclizing by a different mech-
anism, would yield the phenolic ketone required for the synthesis of
usnic acid (Figure 4, pathway ‘3’’). Usnic acid and atranorin are by
far the two most common lichen substances. Both are found mainly

468 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

in the upper cortex rather than in the medulla where most of the
depsides and depsidones accumulate. The production of a depside or
a depsidone in the medulla and either atranorin or usnic acid in the
cortex is the most striking pattern observed in the histological distri-
bution of chemical compounds in the macrolichens. Yet the chemical
structures of atranorin and usnic acid are so distinct that the possibil-
ity of their synthesis from similar starting materials was not immedi-
ately obvious.’

The production of 6-orcinol-type versus orcinol-type substances
may have distinct taxonomic significance since many groups of lichens
show a strong tendency toward the production of one type of compound
or the other. In the genera studied here, the medullary constituents of
Cetrelia are all orcinol-type depsides and depsidones while a §-orcinol
depsidone occurs in Platismatia. Atranorin, a -orcinol depside, 1s
found in the upper cortex of all the species of both genera.

LENGTH AND OXIDATION STATE OF THE SIDE CHAIN IN
ORCINOL-TYPE PHENOLIC ACIDS

Figure 3 shows how one acetate and three malonate units could
condense to form orsellinic acid, a precursor of many lichen and fungal
products. Starting with more than four units would give similar ring
systems with a longer side chain. Since each malonate unit contributes
two carbons to the final product, the side chain will increase in length
in two-carbon steps. Side chains known in lichen substances are C,, C,,
C,, and C;. Figure 5 shows the basic phenolic units of the orcinol-type
compounds. In the Cetrelia compounds, side chains contain five or
seven carbons except in ring B of imbricaric acid, which has a three-
carbon substituent. In these species there is a tendency toward pro-
duction of substances with longer side chains than are typical for
lichens in general. The orsellinic acid ring, so common in the sub-

1 After this section was written, three studies relating to the biosynthesis of
atranorin and usnic acid appeared. Taguchi, Sankawa, and Shibata (1966)
elegantly demonstrated that acetate-malonate-derived methylphloroglucinol
self-condenses to give usnic acid and that the extra methyl group is indeed in-
corporated before the aromatic ring is formed. Yamazaki, Matsuo, and Shibata
(1965) studied the labeling of lecanoric acid, atranorin, and chloroatranorin in
Parmelia tinctorum fed with “C-labeled acetate and formate. They found in-
corporation of “C in all three substances when the lichen was fed with labeled
acetate, but only in atranorin and chloroatranorin, which are the -orcinol-type
depsides, when the lichen was fed with labeled formate. The results indicate
that the extra C; substituents are supplied by formate. In a second study, Yama-
zaki and Shibata (1966) showed that *H-labeled orsellinic acid and g-orcinol-
carboxylic acid are incorporated exclusively into lecanoric acid and atranorin,
respectively, proving that the extra C; substituent is added before aromatization
during the biosynthesis of atranorin.

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 469

PHENOLIC ACIDS DEPSIDES

HO. COOH lecanoric acid, evernic acid,
erytnrin, gyrophoric acid, etc
OH
C3H;
° | divaricatic acid
H COOH .
| sekikaic acid, scrobiculin
OH
C5Hy, OH
o(O)-coo(Q)-coor
OH C3H,
R=H, unknown
ReCH,, imbricaric acid
ramalinolic acid, boninic acid,
homosekikaic acid, merochloro- META-DEPSIDES
Phaeic acid
CH,COC3H,
HO COOH — glomelliferic acid

H

C5Hy

noxCjrcoon
H

OH C5Hy4
R=H, anziaic acid
R=CHg, Perlatolic acid
cryptochioroph
yp phaeic acid DEPSIDONES
CH,COC.H,, CH COCéH, , OH
wo{O)-coos K0{O)-coo{O)-coo —_+ Physodic acid

CsHs, OH

n0{O)-co0{O)-coom

OH OH CsH,,

olivetoric acid

CH,COCSH), OH CH,COC.H,, OH
Om coo ~O}-<o oO) coo {Op-cooe
OH HgCOCSH., fe) CH,COCSH,,
RH, unknown R=H, alectoronic acid
R=CH,, microphyllinic acid R=CH,, collatolic acid
Cais
Ho{O)-coon { sphaerophorin — | grayanic acid
OH (with orsellinic acid?)

Figure 5. Chart showing the phenolic units (left column) incorporated in depsides
(middle column) and depsidones (right column). The heavier lines lead to sub-
stances found in Cetrelia and the chemical structures of these compounds are
given. The thinner lines lead to related substances known from other genera of
lichens and are indicated for purposes of comparison. The phenolic units are
arranged from top to bottom in order of increasing side chain length. Where two
phenolic units have side chains of the same length, the less derived (the B-keto
type) is given first. This arrangement shows that most of the medullary com-
pounds in Cetrelia are both closely related to each other and also more highly
derived within the framework of chemical variation shown by orcinol-type
depsides and depsidones.

470 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

stances in many groups of lichens, is not involved in any known medul-
lary constituents of Cetrelia.

Some orcinol-type phenolic rings in lichen substances retain a car-
bonyl group at the 6-carbon of the alkyl side chain. The position of this
carbonyl group corresponds to that of one of the original malonate
carboxyls, and it seems probable that the B-keto groups result when
there is no reduction at this position rather than by a new oxidation
step. Lobaric acid and norlobaridone each have an a-keto group which
probably forms by oxidation of a previously reduced side chain. Thus
compounds with a-keto side chains may be more derived than com-
pounds with fully reduced side chains, which in turn may require more
steps for synthesis than a substance with a chain of equal length which
retains a 6-keto carbon.

In the orcinol-type depsides and depsidones of Cetrelia, the C; side
chains retain a B-keto group while the C; and C, side chains are fully
reduced (Figure 5). Except for the one C;-substituted compound (imbri-
caric acid), biosynthesis of every phenolic unit in the Cetrelia com-
pounds must involve reduction of two carbonyls, either to the —CsHir
group or to the —CH,COC,H,, group. Oxidized C; alkyl groups are
unknown in lichen substances, and a 8-keto C, substituent is known
only in glomelliferic acid.

Thus the biosynthesis of the KC+ compound microphyllinic acid
may proceed through exactly the same steps required to synthesize
perlatolic acid (KC—) except that the initial condensation leading to
the phenolic rings in microphyllinic acid includes one extra malonate
unit. In olivetoric acid, one ring is substituted as in perlatolic acid and
the other as in microphyllinic acid.

DEPSIDES AND DEPSIDONES

For many years it was supposed that depsides were converted to
depsidones by the loss of water from phenolic groups properly placed
on each ring. There was, however, no known lichen depside with a
structure suitable to give a known lichen depsidone by this route. It is
now generally accepted that depsidones form from depsides by a loss
of hydrogen in an oxidative-cyclization process (Figure 6) (Erdtman,
1962; Erdtman and Wachtmeister, 1957). A number of depside-
depsidone pairs are known which show very similar ring substituents.
In Cetraria ciliaris sens. lat. and in some individuals of Pseudevernia
furfuracea (= Parmelia furfuracea), the depside-depsidone pair olive-
toric acid and physodic acid occur together (C. F. Culberson 1964,
1965). |

If the mechanism shown in Figure 6 is correct, then methylation of
any phenolic hydroxyl or even the carboxylic acid group would disrupt

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 47]

R - OH R OH R

TA, gfe -2H: /OH
HO coo COOH ——————+ -0 coo coo. —————+ 0 CN,
2
> OH © &R OH R O.
H’ °
R COOH
HiSs 9 |
Cc. R OH R
fe) 9 /OH
HO coo COOH ——————. © Cy
CH3O (0) fo)
CoHyy OH fe) R Yo)
OOH H
R fe)
picrolichenic acid COOH

FicureE 6. A possible biosynthetic route from depsides to depsidones. Picrolichenic
acid is formed when oxidative coupling is blocked by 2-O-methylation. (See the
text for fuller explanation.)

the formation of a depsidone from a depside. It has been pointed out
(Erdtman and Wachtmeister, 1957) that methylation of the phenolic
group ortho to the ester linkage can explain the formation of picro-
lichenic acid (Figure 6) by a mechanism similar to depsidone formation.
But methylation at either the 4- or the 2’-hydroxyl may block cycliza-
tion altogether. Microphyllinic acid, which occurs in Cetrelia, is a
depside closely related to alectoronic acid and a-collatolic acid, but it
is methylated at the 4-hydroxyl. The related unmethylated depside is
not yet known from lichens, but it would be the immediate precursor
of the common lichen substance alectoronic acid. The latter substance
could then be converted to a-collatolic acid by 4-O-methylation (Figure
5).

Blocking oxidative-cyclization reactions by methylation of hydroxyls
that would pass through a quinoid structure during the cyclization is
known in the biosynthesis of certain alkaloids where “it seems probable
that methylation precedes and controls the direction of oxidative
coupling. . .”’ (Scott, 1965).

O-METHYLATION

Methylation on oxygen is a very common and simple cause of
chemical variation in lichen depsides and depsidones. All the phenolic
groups in the molecule as well as the carboxylic acid group of ring B
of orcinol-type depsides are subject to methylation, but methyl sub-
stitution at the 4-hydroxyl is most frequently observed. Examples
among the Cetrelia compounds (Figure 5) are perlatolic acid, which is
4-O-methylanziaic acid, and a-collatolic acid, which is 4-O-methyl-
alectoronic acid.

As yet there is no experimental evidence to indicate when methyla-
tion occurs, but the biosynthesis of picrolichenic acid requires 2-O-

472 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

methylation before oxidative cyclization. On the other hand, if 4-O-
methylation or 2’-O-methylation can block depsidone formation as
suggested above, then depsidones must be methylated at these posi-
tions after oxidative cyclization.

It seems that the order of methylation may vary and perhaps the
taxonomic significance of substances which differ by methylation alone
will also vary. In Cetrelia, for example, anziaic acid and perlatolic acid
do not seem to occur together and the species in which they occur
appear to be very distinct, while alectoronic acid and a-collatolic acid
are always found together and seem to be more closely related
biosynthetically.

RELATIONSHIP OF CETRELIA COMPOUNDS TO OTHER LICHEN
SUBSTANCES

Figure 5 shows the orcinol depsides in Cetrelia and some closely
related substances from other lichens. The chart is arranged to show,
in a very general way, the relationships among these substances and
the extreme uniformity of the structures involved. All seven medullary
compounds in Cetrelia may be derived from three phenolic-acid units
and six of the substances from only two such units. Reactions probably
required in the biosynthesis of these compounds are (1) esterification of
the phenolic units, (2) 4-O-methylation (in imbricaric acid, perlatolic
acid, a-collatolic acid), and (3) oxidative cyclization to a depsidone (in
alectoronic acid and a-collatolic acid). (See the caption of Figure 5 for
additional explanation.)

The Cetrelia substances do not show some of the variations observed
in other orcinol-type depsides and depsidones. O-Methylation occurs
only at the 4-hydroxyl, for example, and more highly methylated sub-
stances, such as planaic acid (a methyl ether of perlatolic acid) and
confluentic acid (a trimethyl ether of olivetoric acid), are not found
in Cetrelia. Nor are meta depsides found, although many of the known
meta depsides could be derived from the same phenolic acid units in-
volved in the Cetrelia compounds. Perhaps some of the minor constit-
uents not identified in the present study will represent modifications
of chemical structure.

While the Cetrelia substances seem to be uniform, the synthesis of
a-collatolic acid, a monomethyl ether depsidone, represents a certain
degree of advancement over, say, the production of simple depsides
such as lecanoric acid. The production of relatively advanced but
highly uniform chemical substances would substantiate the concept
of these species as evolved from ancestors which produced such com-
plex compounds as alectoronic acid and a-collatolic acid. If this is
true, the other substances found here might have resulted from a

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 473

retrogressive evolution in which reactions leading to alectoronic acid
and a-collatolic acid became blocked or modified.

Considering the complete lack of direct evidence, other plausible
theories to account for the production of these substances could be
advanced, but the route suggested here is the simplest one in the light
of present knowledge. This hypothesis would support the concept of
the origin of Cetrelia from an Amphigymnia-type ancestor producing
alectoronic acid and a-collatolic acid. Hale (1965) found that alecto-
ronic acid (usually with a-collatolic acid) is indeed the most common
medullary substance in Parmelia subgenus Amphigymnia. As in
Cetrelia, all of these species also produce atranorin in the cortex, but
unlike Cetrelia the other species of Amphigymnia Parmelias produce a
wide variety of chemical substances including fatty acids, meta dep-
sides, orsellinic acid-derived depsides, and -orcinol depsidones. It is
significant that the common cortex substance usnic acid is produced in
Parmelia subgenus Amphigymnia but not in Cetrelia, and it is further
noteworthy that usnic acid never occurs in those Amphigymnia Par-
melias which produce alectoronic acid or olivetoric acid.

Methods for the Microchemical Identification of Lichen Substances in
Cetrelia and Platismatia

The aim of this section is to give a complete description of the most
reliable and convenient tests for determining the constituents of the
species treated in this monograph. The usual microcrystal tests (Asah-
ina, 1936-1940) and paper-chromatographic methods (Wachtmeister,
1959) have been used to identify the lichen substances in herbarium
specimens in this study, although in the time since the research
reported here was done thin-layer chromatography has become the
major chromatographic technique in lichen microchemistry. The
descriptions here are intended to give the taxonomist enough infor-
mation for reliable, practical identifications of the nine substances
involved in Cetrelia and Platismatia and are not meant as a complete
treatment of all the microcrystal or chromatographic tests which
might conceivably be used for these substances. The methods described
are sufficient to distinguish these substances from each other as they
occur in our genera, but alone they might not be sufficient to identify
the same substances in all the other lichens for which additional con-
stituents might introduce complications. In the research described
here, many other tests were actually used. These studies were neces-
sary to establish the data reported, but they are not required for
routine identifications in specimens of Cetrelia and Plastismatia now
that the major constituents are known.

474

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

SOLUTIONS

The following solutions are used in the microchemical procedures to
be described :

1.

9

10.

GE solution. A mixture of three parts of acetic acid and one
part of glycerin.

_ GAW solution. A mixture of equal volumes of ethyl alcohol,

water, and glycerin.

3. GAo-T solution. A mixture of one part o-toluidine, two parts

alcohol, and two parts glycerin. The solution should be
discarded when it turns red. A nearly colorless solution can
be prepared if the o-toluidine is first distilled from zinc
powder.

5 percent NaOH. A solution of 5 grams of sodium hydroxide
in 95 ml. of water.

Ammonium hydroxide-saturated n-butanol. Concentrated
ammonium hydroxide is added to n-butanol and the mix-
ture is shaken thoroughly in a separatory funnel or bottle
(the gas pressure frequently being released). Sufficient
ammonium hydroxide is added so that, after shaking, two
layers will separate. The upper layer and a smaller amount
of the lower layer are placed in the chromatographic tank.

Pyridine-ethyl acetate-water. A mixture of one part pyridine,
two parts ethyl acetate, and one part water.

Diazotized benzidine. Dissolve 1.7 grams of benzidine di-
hydrochloride in 250 ml. of water and add 3 ml. of con-
centrated hydrochloric acid. Prepare a separate solution of
25 grams of sodium nitrite in 250 ml. of water. Just before
use, one part of the sodium nitrite solution is added, by
drops, to one part of the benzidine solution.

. Stable PD. A solution of 10 grams of sodium sulfite and 1

gram of p-phenylenediamine in 100 ml. of water to which is
added 0.5 ml. or less of a wetting agent (such as ‘“‘Pho-
toflo’’).

0.1 m Na,;PQ,. A solution of 3.8 grams of trisodium phosphate
dodecahydrate in 100 ml. of water.

1 n H,SO,. A solution prepared by adding 3 ml. of concen-
trated sulfuric acid to 97 ml. of water with stirring.

In addition to these solutions, acetone, benzene, ethyl alcohol, and
ethyl ether, and the usual spot test reagents are used in the micro-
crystal and paper chromatographic procedures to be described.

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 475

MICROEXTRACTION

Thallus fragments equivalent to approximately one square centi-
meter are placed in a shell vial (1.2 by 3.5 cm.) and enough benzene is
added to cover the sample. After about 5 minutes the benzene is de-
canted onto a microscope slide on a slide-warming tray (about 55°C).
The extraction with benzene at room temperature is repeated twice
more, and the solutions are evaporated on the same heated slide. Then
the vial is placed on the warming tray and just enough acetone is
added to cover the sample. The fragments are extracted for 5 to 10
minutes and the solution is then evaporated on another heated slide.
This procedure is repeated twice more, as was done for the benzene
extraction. The lichen sample is discarded, and then the two slides with
the residues are ready for use.

MICROCRYSTAL TESTS

The microcrystal tests are performed on portions of the residues from
the acetone and benzene extracts. Small amounts of residue are re-
moved with a razor blade and transferred to another slide. The amount
of solid residue needed for a test depends primarily upon (1) the solu-
bility of the crystals in the test solution, (2) the volume of test solution
used, and (3) the concentration of the substance in the residue being
tested. Using a very small cover slip reduces the volume of test solution
applied and consequently the amount of solid residue required for a
crystal test. If the crystal test fails the first time, a second test with
more (or sometimes less) of the residue often gives better results.

Solutions for the crystal tests described below are the GE, GAW,
and GAo-T solutions and a 5-percent aqueous NaOH solution. A drop
of the test solution is applied to a cover slip which is gently placed on
the residue so that the liquid covers the solid. Tests with the GAo-T
solution and with 5 percent NaOH usually do not require heat. Tests
with the GE and GAW solutions depend upon a recrystallization proc-
ess and the slides should be warmed, preferably over a microflame.
Many of the substances treated here are more soluble in the GE solu-
tion than in the GAW solution, and usually less heat is needed for
tests in the GE solution. The solutions should not be allowed to boil,
and heating should be stopped at once if the solid seems to dissolve
rapidly. Most preparations are now ready for microscopic examination.
If crystals have not formed, the slides are reexamined later.

PAPER CHROMATOGRAPHY

Paper chromatograms are generally prepared on Whatman No. |
paper although other types of paper (e.g., Whatman No. 3 MM and

476 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Toyo Roshi No. 50) are equally satisfactory. The chromatograms are
developed by the ascending method, which is simpler than the descend-
ing method and requires less elaborate equipment. Any chromato-
graphic tank is satisfactory, but ideally it should be tall enough for
development of the chromatograms to 15 to 20 cm. and still not so
large that it becomes difficult to saturate the atmosphere in it with
the developing solvent. A filter-paper lining around the inside of the
tank helps to keep the atmosphere saturated.

The spots of the extracts are applied 1.5 cm. apart along a line 3 cm.
from the bottom of the paper. A small amount of the residue on the
slide is treated with a drop of acetone, and a portion of this solution is
drawn by capillarity into a narrow glass tube with an inside diameter
of about 1 mm. (thin-walled capillary tubing which has been pulled
out in a flame to a point). The tip of the tube is then touched several
times to a marked point on the starting line. With care, the spots can
be kept 3 to 5 mm. in diameter, thereby producing better results with
less material than larger spots would. The amount of substance needed
for the chromatogram depends largely on how its constituents sepa-
rate and the sensitivity of the spray reagent. The reagents used in
the present study are extremely sensitive, and the spots are easily
overloaded. The optimum quantity of material to be applied to a spot
can be judged easily after a few trials.

Paper chromatograms of all substances described here except fumar-
protocetraric acid are developed in ammonium hydroxide-saturated
n-butanol. With this solvent there are two liquid layers. The upper
layer is ammonium hydroxide-saturated n-butanol and the chromato-
gram should be adjusted so that it dips into only this layer. For paper
chromatograms of fumarprotocetraric acid, the single-phase solvent
pyridine-ethyl acetate-water (1:2:1) is used.

When the chromatogram has developed to a height of 15 to 20 cm.,
it is dried and then viewed with ultraviolet light. Both long- (366 my)
and short-wave (254 mu) lamps were used in the present study. The
short-wave light was superior largely because it delivered much less
visible light than the particular long-wave lamp available, and the
fluorescent spots are more easily distinguished in darkness. The spots
are encircled in pencil and labeled for color of fluorescence.

Chromatograms run in ammonium hydroxide-saturated n-butanol
are sprayed with a freshly prepared diazotized benzidine solution, and
after a few minutes the paper is washed with cold water and dried.
Chromatograms of fumarprotocetraric acid developed in pyridine-
ethyl acetate-water are sprayed with the stable PD solution. The
chromatograms should be sprayed in a hood or in some way so that
the mist of the reagents is not inhaled.

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 477

All paper chromatograms should be saved. Chromatograms sprayed
with stable PD can be kept for some time in a large plastic envelope.

HYDROLYSIS OF DEPSIDES

The depsides imbricaric acid and perlatolic acid can be confirmed by
chromatography of the hydrolyzed residues, and this procedure is also
particularly useful in distinguishing olivetoric acid from anziaic acid.
The methods described here are slightly modified from those suggested
by Wachtmeister (1956) and have been presented in detail elsewhere
for imbricaric acid and perlatolic acid (C. F. Culberson and W. L.
Culberson, 1965).

A portion of the extract containing the unknown substance is re-
moved from the residue slide with a razor blade and transferred to
a vial. The amount required is usually about two to four times as much
as is needed for a microcrystal test. The residue is covered with three
or four drops of 0.1 m NasPO,. The tube is kept stoppered at room tem-
perature or very gently warmed for 15 to 24 hours. After this time most
substances produce yellow, orange, pink, or red colors. A little ether is
added, and after a few minutes the tube is closed and shaken sufficiently
to mix the two layers. When the layers separate, the upper (ether)
solution is drawn off carefully to avoid including any of the lower
(aqueous) one. The ether solution is evaporated on a warmed slide and
the extraction is repeated twice. This extract will usually contain
phenolic substances and possibly excess starting material. Next, the
aqueous layer is acidified with two drops of 1 n H2SO, and extracted
with ether as before, evaporating the ether extracts on a second slide.
Both extracts are chromatographed in the ammonium hydroxide-
saturated n-butanol solvent and the chromatogram is viewed with
UV light and sprayed with diazotized benzidine.

Residues may also be hydrolyzed with concentrated sulfuric acid.
The solid is placed in a vial and covered with two or three drops of the
acid. After ten minutes at room temperature or (better) in a refrig-
erator, the tube is filled with crushed ice. When the ice melts the solu-
tion is extracted with ether three times as described above; the ether
solutions are evaporated on a slide; and the aqueous layer is discarded.
The residue is chromatographed as described above.

The appearance of the chromatograms from hydrolyzed extracts
will vary with the degree of decarboxylation and other side reactions
depending upon the particular conditions of the extraction, but the
results from this simple test can be particularly valuable.

THIN-LAYER CHROMATOGRAPHY

After the completion of the microchemical studies for this mono-
graph, the technique of thin-layer chromatography was developed

277-920 O - 68 - 3

478 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

(Stahl, 1965) and has subsequently been applied successfully to the
determination of many lichen substances. Thin-layer chromatography
was employed briefly using samples of Cetrelza previously studied by
the conventional microchemical methods, and in general good results
were obtained. For some species, additional spots could be observed
which were not found by paper chromatography, but these constit-
uents, present in minute quantities, could not be identified. Table 4
vives the R, values found for the eight substances known from Cetrelia,
The chromatograms were run on air-dried silica gel-G plates developed
with n-hexane-ethyl ether-formic acid and sprayed with 10 percent
H2SO,. The plates were dried in an oven at 70°C and the color of the
spots varied somewhat with the duration and conditions of drying.
On the same chromatograms, extracts of Platismatia lacunosa contain-
ing fumarprotocetraric acid showed a black spot at the origin.

TABLE 4.—R; values in thin-layer chromatograms for compounds known in Cetrelia. The chromatograms were
made on air-dried silica gel-G plates, developed with n-herane-ethyl ether-formic acid toa height of 10 cm.,
sprayed with 10 percent H,SO,, and then dried at 70°C.

Substance R: Color After Spraying

Alectoronic acid
Anziaic acid
Atranorin

22 brown
50 yellow
73 yellow-brown

0.

0.

0.
a-Collatolic acid 0. 28 brown
Imbricaric acid 0. 66 yellow
Microphyllinie acid 0. 35 red-brown
Olivetorie acid 0.35 yellow-brown
Perlatolic acid 0. 73 yellow

Identification of Specific Substances

ALECTORONIC ACID AND a-COLLATOLIC ACID

PLATE 1, FIGURES 1-3

The medulla of samples containing alectoronic acid or alectoronic
acid and a-collatolic acid shows a strong KC+ reaction and a brilliant
white fluorescence in UV light. Neither substance gives a color test
with K, C, or PD. The microchemical tests used to identify these com-
pounds in the present study have recently been described elsewhere
(W. L. Culberson and C. F. Culberson, 1965).

Benzene at room temperature dissolves some alectoronic acid and
a-collatolic acid. Since a-collatolic acid is slightly more soluble in this
solvent than alectoronic acid is, small quantities of a-collatolic acid in
the mixture can often be detected better in the benzene extract than
would be possible using a single acetone extraction.

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 479

Plate 1, Figures 1-2, shows examples of the variable crystal form of
alectoronic acid and a-collatolic acid cocrystallized from the GE
solution. Crystal morphology in the GAW solution is also variable and
a common crystal type is illustrated (Plate 1, Figure 3). The final
identification of these substances must be made by chromatography.
In ammonium hydroxide-saturated n-butanol, alectoronic acid shows
a spot at R, 0.24 which appears bluish in UV light and turns deep red
with diazotized benzidine. On the same chromatogram, a-collatolic
acid produces a long spot at about R, 0.56 which shows a pale fluores-
cence in UV light and turns dark red with diazotized benzidine. The
spot for atranorin runs between that for alectoronic acid and that for
a-collatolic acid.

ANZIAIC ACID

PLATE 1, FIGURES 4-5

All Cetreliae showing a C+ medullary reaction contain either anziaic
acid or olivetoric acid. Anziaic acid is less common. Like olivetoric
acid, it shows no color reactions with K or PD. Anziaic acid is slightly
more soluble in benzene than is olivetoric acid, which is only scarcely
soluble. The benzene extract contains atranorin along with anziaic
acid, and the subsequent acetone extract contains more anziaic acid,
often accompanied by more atranorin.

Frequently no crystals of anziaic acid form in the GE solution be-
cause the compound is very soluble there, but when recrystallization
does occur, there are clusters of small, curved needles (Plate 1, Figure
4). Anziaic acid recrystallizes more dependably from the GAW solu-
tion, but the curvature and size of the needles which form (Plate 1,
Figure 5) are variable. Unlike olivetoric acid, anziaic acid does not
produce very persisting crystals in the GAo-T solution. Colorless
needles which may be present in a fresh GAo-T preparation soon
dissolve.

Anziaic acid chromatographs well in ammonium hydroxide-satu-
rated n-butanol giving a spot at R, 0.67 which is blue in UV light and
bright red with diazotized benzidine. While there may be a slight red
coloration with diazotized benzidine just above the spot for anziaic
acid, the chromatogram is distinguishable from the typical double
spot produced by olivetoric acid near the same R; value. Determina-
tion of anziaic acid can be confirmed by hydrolysis since chromato-
graphic analysis then shows a single spot for olivetol-2-carboxylic acid
near R, 0.60.

480 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

ATRANORIN
PLATE 1, FIGURE 6; PLATE 2, FIGURES 7-10

A number of papers describe the microchemical identification of
atranorin. This compound turns yellow with K and pale yellow with
PD, although the latter reaction is rarely observed on the thallus.
Atranorin is somewhat soluble in benzene at room temperature but is
usually found in the subsequent acetone extract as well unless it is
present in the thallus in only very small concentration. Colorless
prisms in the GE solution (Plate 1, Figure 6) and yellow needles in
the GAo-T solution (Plate 2, Figures 7-8) are used to identify this sub-
stance. The crystal test in GAo-T is very sensitive (C. F. Culberson,
1963), and it is usually successful even in the presence of other com-
pounds. Large concentrations of alectoronic acid, a-collatolic acid,
and especially microphyllinic acid, however, may inhibit the crystal
formation of atranorin in the GAo-T solution and also sometimes may
decrease the size or alter the appearance of the prisms in the GE solu-
tion. The shade of yellow of the crystals in the GAo-T solution is
somewhat variable, perhaps due to the presence of chloroatranorin or
other compounds in the samples. Two types of atranorin crystals in
the GAo-T solution are shown (Plate 2, Figure 9). The smaller, finer,
more sharply curved needles are also a somewhat deeper yellow color.
Atranorin is sparingly soluble in alcohol and does not recrystallize
from the GAW solution.

Chromatography in ammonium hydroxide-saturated n-butanol
gives a good spot (R; 0.53) which is yellow in the ammonia atmosphere
of the developing tank, fluoresces yellow in UV light, and turns yellow
with diazotized benzidine. In the pyridine-ethyl acetate-water devel-
oping solvent, atranorin travels with the solvent front (R,;=1).

CAPERATIC ACID
PLATE 2, FIGURES 11-12; PLATE 3, FIGURES 13-18

The only fatty acid definitely proved to occur in the genera studied
here is caperatic acid, but microcrystal tests are not a good method for
identifying fatty acids, and it is certain that other constituents will be
found when better techniques are perfected.

Caperatic acid shows no thallus spot tests. It is sparingly soluble in
benzene at room temperature while protolichesterinic acid, another
common fatty acid of lichens, dissolves readily. By extracting the
thallus fragment first with benzene and then with acetone, these two
fatty acids would be separated from each other. This solubility dif-
ference is fortunate since experiments with pure caperatic acid ex-
tracted from Parmelia ceryptochlorophaea Hale (C. F. Culberson, 1965)

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 48]

show that under certain conditions this compound produces crystals in
the GE solution which are similar to those of protolichesterinic acid.

Normally, caperatic acid crystallizes from the GE solution as cir-
cular or cloud-shaped clumps appearing somewhat similar to
drops of oil (Plate 3, Figures 13-14). These crystals are very distinctive
and easily recognizable. But many GE preparations show plate-like
crystals, singly or in radiating clusters (Plate 3, Figures 15-16), similar
to those typical of protolichesterinic acid. Sometimes plate-like crys-
tals seem to emerge from the circular ones (Plate 3, Figures 17-18).
At first this gives the impression of a mixture of protolichesterinic acid
and caperatic acid. But, since the lichen fragment was extracted first
with benzene and this extract showed no protolichesterinic acid, the
presence of protolichesterinic acid in the second residue is not likely.

Chemical transformation of caperatic acid into protolichesterinic
acid under the influence of acetic acid in the GE solution is extremely
unlikely. A careful comparison of the crystal shapes indicates two
possible explanations: (1) either caperatic acid has two crystal forms or
(2) caperatic acid here is sometimes converted to some substance other
than protolichesterinic acid. The most obvious reaction which might
occur in the GE solution is the hydrolysis of the methyl ester of capera-
tic acid, giving norcaperatic acid. A sample of pure norcaperatic acid
showed crystals (Plate 4, Figure 19) in the GE solution that were very
similar to, but probably not identical with, the cloud-type crystals of
caperatic acid. The crystals of norcaperatic acid in the GAW solution
(Plate 4, Figure 20) are indistinguishable from those of caperatic acid.

Crystals of caperatic acid in GAW solution are very small circular
plates usually clustered into large radiating aggregates (Plate 2,
Figure 12). In some preparations there are curly crystals, especially at
the center of the aggregates of caperatic acid (Plate 2, Figure 11), per-
haps of an unidentified fatty acid.

Caperatic acid shows small, fuzzy, colorless balls in the GAo-T
solution, and it distorts the yellow crystals produced by atranorin
when the latter substance is also present (Plate 2, Figure 10).

When samples containing caperatic acid are chromatographed in
ammonium hydroxide-saturated n-butanol, a weakly fluorescing,
trailing spot may be found near R, 0.75. When the chromatogram is
sprayed with diazotized benzidine to reveal atranorin or other com-
pounds, the spot for caperatic acid turns very pale yellow and, while
the chromatogram is still wet, is barely visible. This is not, of course, a
reliable method for identifying caperatic acid.

FUMARPROTOCETRARIC ACID

In this study one species, Plastismatia lacunosa, gave a positive test
with PD and contained fumarprotocetraric acid. This substance is not

482 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

removed from the thallus by extraction with benzene at room tempera-
ture. Fumarprotocetraric acid dissolves in warm acetone but in this
species it is accompanied by atranorin and usually also caperatic acid.
From residues also containing caperatic acid, crystals of the o-toluidine
derivative of fumarprotocetraric acid are not readily identifiable in the
GAo-T solution, and all determinations of this substance were made
by chromatography.

Paper chromatograms developed with ammonium hydroxide-
saturated n-butanol show fumarprotocetraric acid as a dense streak
near the origin. The spot turns dark red with diazotized benzidine and
differs little in appearance from that of several other PD-+ depsidones.
Extracts from specimens showing such a spot were rechromatographed
in pyridine-ethyl acetate-water (1:2:1). With this solvent, fumarproto-
cetraric acid shows a distinctive spot (R; 0.39) which quenches UV
light and turns yellow after spraying with the stable PD reagent.

IMBRICARIC ACID
PLATE 4, FIGURES 21-23

The microchemical methods used to identify imbricaric acid are
described in detail elsewhere (C. F. Culberson and W. L. Culberson,
1966). A pure sample of this substance extracted from Cetrelia sinensis
was used to check the microchemical tests used on crude extracts from
thallus fragments. Imbricaric acid is negative with K, C, KC, and PD,
but the KC+ reaction of the thallus of many samples containing 1m-
bricaric acid requires that the latter substance be distinguished from
both KC+ and KC — compounds—in this study perlatolic acid,
alectoronic acid, a-collatolic acid, and microphyllinic acid.

Imbricaric acid is somewhat soluble in benzene at room tempera-
ture. The subsequent acetone extract is often gummy and less satis-
factory for the microcrystal tests. The large needles formed in the GE
solution (Plate 4, Figure 21) show oblique extinction (¢=45°) in
polarized light and in this respect differ from those of all other sub-
stances encountered here. The needles in the GAW solution (Plate 4,
Figure 22) also show oblique extinction (¢=45°). In the GAo-T solu-
tion, imbricaric acid forms colorless needles (Plate 4, Figure 23), probably
its o-toluidine salt. In the crude extracts where atranorin is also pres-
ent, fuzzy yellow crystals form (Plate 4, Figure 24). These crystals
may result from cocrystallization since very similar crystals were ob-
served when a mixture of pure imbricaric acid and pure atranorin was
treated with the GAo-T solution.

By chromatography in ammonium hydroxide-saturated n-butanol,
imbricaric acid undergoes slow hydrolysis giving a spot at R, 0.78,

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 483

blue-white in UV light and orange-red with diazotized benzidine,
which trails down from the spot to about R, 0.38 to 0.43 due to the
degradation product divarie acid. Imbricaric acid can be definitely
distinguished from perlatolic acid by hydrolysis with acid or base and
by observing the strong spot at R, 0.38 for divaric acid. In most cases,
however, the observation of crystals with oblique extinction in the
GE and GAW solutions and chromatography of the unhydrolyzed
material are sufficient to identify imbricaric acid.

MICROPHYLLINIC ACID

PLATE 5, FIGURES 25-27

Microphyllinic acid is negative with K, C, and PD but gives a
strong red coloration with KC. In at least one report in the literature,
microphyllinic acid was mistaken for alectoronic acid because both
substances give a positive KC test and needles in the GE solution.
But microphyllinic acid can be definitely identified by microcrystal
tests and chromatography.

Some microphyllinic acid is removed from the thallus fragment by
extraction with benzene at room temperature. Some atranorin is also
extracted then. The subsequent acetone extract contains less micro-
phyllinic acid and is often a dark gum that is difficult to work. The
microcrystal tests usually give best results with the benzene extract.
In the GE solution, straight radiating needles form (Plate 5, Figure
25) with an extinction angle of zero. Atranorin often crystallizes in the
same preparation, although the microcrystal test for atranorin in the
GAo-T solution is inhibited by microphyllinic acid. In the GAW
solution, microphyllinic acid recrystallizes as long, fine, sparsely
clustered needles with an extinction angle of zero (Plate 5, Figure 26).
These crystals are most similar to those of perlatolic acid but do not
resemble those of alectoronic acid. The most rapid method for dis-
tinguishing microphyllinic acid from the other KC+ substances in
Cetrelia is by the crystals of the sodium salt (Plate 5, Figure 27) which
precipitate from 5 percent aqueous sodium hydroxide.

Microphyllinic acid hydrolyzes completely during chromatography
in ammonium hydroxide-saturated n-butanol, and two characteristic
spots are observed. The upper one, (R,; 0.93), which shows a bright
blue-white fluorescence in UV light and turns orange with diazotized
benzidine, is due to the ring A fragment (Figure 5). The lower spot
(R, 0.49), which interferes with the chromatographic determination of
atranorin, appears blue in UV light, turns dark red with diazotized
benzidine, and is due to the ring B fragment. The same spots are ob-
served after base hydrolysis.

484 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

OLIVETORIC ACID

PLaTE 5, FIGURES 28-30; PLATE 6, FIGURE 31

Samples showing a positive thallus reaction with C may contain
either anziaic acid or olivetoric acid. Olivetoric acid gives no coloration
with K or PD, but turns red with C, and the color is still more intense
with KC. Olivetoric acid is reported to give a green color with aqueous
Ba(OH), (Asahina, 1936) while anziaic acid does not give such colora-
tion, but this reaction was not used in the present study. Olivetoric
acid is only slightly soluble in benzene and less soluble in it than
anziaic acid is. The benzene extract of thallus fragments with olive-
toric acid and atranorin encountered in this study is usually a small,
colorless crust which contains only a trace of olivetoric acid. The sub-
sequent acetone extract contains more atranorin and abundant olive-
toric acid.

In the GE solution, olivetoric acid is quite soluble and it crystallizes
slowly as small, fine, curved needles forming a dense mat (Plate 5,
Figure 28). Olivetoric acid is less soluble in the GAW solution, and it
usually produces beautiful, curved needles (Plate 5, Figure 29), which
are larger and thicker than those in the GE solution. The crystals of
anziaic acid in the GE and GAW solutions (Plate 1, Figures 4-5) are
sufficiently similar to those of olivetoric acid that tests using these
crystallizing reagents may require confirmation. Olivetoric acid also
produces crystals in the GAo-T solution without heating. These clus-
ters of straight, colorless needles or needle plates (Plate 5, Figure 30;
Plate 6, Figure 31) form rapidly and are usually mixed with the yellow
crystals formed by atranorin. Although anziaic acid may show small
colorless needles under the same conditions, these soon dissolve.

Olivetoric acid hydrolyzes during chromatography in ammonium
hydroxide-saturated n-butanol producing a double spot—one center
(R, 0.73) turns dark bluish-red with diazotized benzidine and the
other (R; 0.60) turns dark red with the same spray. These spots cor-
respond closely to those produced by base hydrolysis which gives a
spot for olivetol-2-carboxylic acid that is blue in UV light and turns
dark red with diazotized benzidine (R, 0.59) and a second spot, from
ring A, that is blue in UV light and dark bluish-red with diazotized
benzidine (R; 0.76). Anziaic acid also gives olivetol-2-carboxylic acid
by hydrolysis but not the second spot at R, 0.76.

PERLATOLIC ACID

PLATE 6, FIGURES 32-34

Perlatolic acid gives no color reactions with the usual reagents, but
some specimens containing this substance show a positive reaction

CETRELIA AND PLATISMATIA 485

CULBERSON & CULBERSON

with KC due to the presence of a still unidentified compound. This
situation is also encountered with samples containing imbricaric acid.
Because of the spurious KC+ reaction observed with perlatolic and
imbricaric acid-containing samples, the microchemical tests must
distinguish these substances from each other and from the known
KC-+ substances occurring in Cetrelia.

Extraction of the thallus fragment with benzene at room tempera-
ture removes a portion of the perlatolic acid along with atranorin.
The benzene extract is usually cleaner and more easily crystallized
with the microcrystal solutions than is the subsequent gummy acetone
extract.

When either extract is treated with the GE solution, perlatolic acid
recrystallizes as clumps of needles (Plate 6, Figure 32) with straight
extinction. Atranorin is usually observed in the same slide. In the
GAW solution, perlatolic acid forms needles (Plate 6, Figure 33)
which are usually shorter than those of imbricaric acid and their angle
of extinction is zero. Many residues containing perlatolic acid show
large colorless crystals (Plate 6, Figure 34) in the GAo-T solution when
a test is made for atranorin. These crystals are very different from those
of imbricaric acid in the same solution, but they have not been ob-
served regularly enough to provide a good test for perlatolic acid.

Perlatolic acid chromatographs in ammonium hydroxide-saturated
n-butanol with an R; of about 0.83 which is slightly higher than the
R, value for imbricaric acid. When the developing solvent is fresh,
perlatolic acid hydrolyzes slightly and trails to an R; of about 0.59 due
to the formation of olivetol carboxylic acid. The spot for perlatolic
acid appears blue-white in UV light and turns orange-red with diazo-
tized benzidine. Chromatographic confirmation of the presence of
perlatolic acid is obtained by acid or base hydrolysis and chroma-
tography of the products. The spot for olivetol carboxylic acid is at R,
0.59 as compared to that for divaric acid at R; 0.38, which would be
found by hydrolysis of imbricaric acid. This procedure has been
described in detail (C. F. Culberson and W. L. Culberson, 1966).

AN UNIDENTIFIED SUBSTANCE

PLATE 6, FIGURES 35-36

An unidentified substance in several species of Cetrelia gives yellow
crystal clusters (Plate 6, Figure 35) in the GE solution. Many of the
same specimens showed fine yellowish-tan needles (Plate 6, Figure 36)
in the GAo-T solution, and it is possible that the crystals in these two
solutions are caused by the same compound. The yellow crystals in
the GE solution usually form near the edge of the slide, and the solu-
tion in the center of the preparation does not appear yellow. This sug-

486 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

gests that the unidentified substance might not be a yellow pigment in
the lichen but rather a colorless substance which turns yellow and
deposits crystals in the GE solution where it is exposed to the air.

Chemical Criteria in the Taxonomy of Cetrelia and Platismatia

In the previous discussion of the generic traits of Cetrelia and Platis-
matia, the high level of chemical uniformity within each genus was
pointed oyt, an internal homogeneity all the more notable considering
the great magnitude of the chemical difference between the two genera
(Table 2). Chemistry here offers a striking correlation with the other
characteristics that define the genera. The use of chemistry in the
study of Cetrelia and Platismatia has considerable theoretical interest
for taxonomy because it shows (1) the value of chemistry in an
instance where it becomes a major characteristic defining genera,
(2) the usefulness at the species level of chemical variations that are
highly correlated with morphological variations in a genus which
has undergone evolutionary diversification in both chemistry and
morphology (Cetrelia), and (3) an example of a chemically conserva-
tive genus (Platismatia).

The involvement of chemistry in addition to morphology in the
formulation of taxonomic judgments in lichenology has had a limited
but nonetheless distinguished history, some of the most refined exam-
ples of its use being the elegant taxonomic systems constructed by
Yasuhiko Asahina in Japan and by the late Alexander Evans in this
country. Although today an increasing number of biochemists are
becoming interested in the unique natural products of the lichens,
lichen taxonomists by and large continue to ignore chemical variation.
The major objection that most lichenologists raise to chemical con-
siderations in lichenology is that chemistry, especially at the species
level, is not used in most other groups of plants and that consequently
it should not be used in lichenology either. The slight taxonomic use-
fulness of corolla pigmentation in the flowering plants is regularly
pointed to as proof of the inconsequential significance of chemical
variation as an indication of the natural relationships of plants at the
species level. In rejecting chemistry from taxonomic considerations,
Almborn (1965) probably summed up most present-day lichenolo-
gists’ views when he wrote: “It is essential that lichen taxonomy be
founded on firm principles not deviating too much from those gener-
ally accepted in other plant groups.”’

While Almborn’s view may appear to reflect the commendable cau-
tion that characterizes responsible scientific judgment, most taxono-
mists recognize that the taxonomic characters valuable in one group of
plants may be of absolutely no value whatever in another. In speaking

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 487

of vascular plants, Cronquist (1964) remarked: ‘‘We have found, to
our sorrow, that individual morphologic characters do not have a
fixed, inherent taxonomic importance wherever they occur. Instead,
they are only as important as they prove to be, in any individual in-
stance, in marking groups that have been perceived on the basis of all
the available evidence.” This observation is no less true for the crypto-
gamic plants than for the vascular plants, and it is no less true for
chemical characters than for morphologic characters. The best taxon-
omy is always eclectic. A blind rejection of a whole category of informa-
tion—in this case chemical information—is as extreme and as philo-
sophically unsound as the diametrically opposed view would be—
namely, to insist upon reflecting in the taxonomic system every minute
variation that a diligent chemist could find. In the actual practice of
classification, the taxonomist needs considerably more latitude and
freedom than either of these extreme positions would permit since the
different types of chemical variation to be found in the lichens do not
all seem to be equally important.

In order to explain how the chemical information found in the pres-
ent study was actually applied to the formulation of the taxonomic
judgments, it is necessary to recall what sort of chemical data were
actually found—although there is nothing unique about the chemical
information here. The nature of the chemical variation in Cetrelia and
Platismatia is essentially like that found in all the rest of the macro-
lichens. The chemistry of the species of these genera can be charac-
terized as the production of two histological categories of substances,
the cortex constituent and the medullary constituent (or constituents).

Individuals of Cetrelia and Platismatia all produce atranorin as the
upper-cortex constituent. No other lichen substance is produced in
this tissue and apparently every individual of every species produces
this constituent. The only other common upper-cortex substance,
usnic acid, a very widespread constituent in Parmelia for example, is
never produced in the two new genera. Since it shows no variation,
upper-cortex chemistry in Cetrelia and Platismatia is of no taxonomic
value.

The chemistry of the medulla is more variable, at least in Cetrelia;
the individual produces one substance (or occasionally two), and the
production of a given substance or of a given pair of substances pre-
cludes the production of other medullary compounds. If morpholog-
ically uniform or highly similar specimens show any qualitative
variation in medullary chemistry, it is a variation of the typical re-
placement type in which one compound is substituted for another,
usually with the chemically different populations showing different
geographic ranges—for example, Cetrelia japonica with microphyllinic

AS& CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

acid in Japan and C. sinensis with imbricaric acid in Formosa and
China.

In interpreting chemical data in lichen species, one is faced with two
questions: (1) To what extent are the data on the nonoccurrence of
given substances in specimens and in species really due to true absence
and to what extent are they due merely to the failure of technique to
reveal their presence? (2) What is the biological significance (and con-
sequent taxonomic significance) of the chemical variations that are
found?

The pattern of chemical variation in which every specimen produces
one medullary compound (or occasionally one pair of compounds) and
in which morphologically similar plants exist in one or several of
“chemical races” is so well known that it is now clear that it is
the major type of chemical variation in all the lichens. In the pres-
ent study every species matches this chemical pattern in that every
one has at least one distinct and constant medullary constituent.
There are no specimens without medullary compounds. In the present
study the constant medullary compounds are given high taxonomic
significance. Morphologically similar or identical populations with dif-
ferent but constant medullary constituents are considered distinct
species. (Cetrelia collata with imbricaric acid is considered distinct
from C. nuda with a-collatolic and alectoronic acids.) The constant
constituents of the species are listed in Table 2.

In addition to the constant constituent medullary compound, some
populations show an additional substance (or additional substances)
in some specimens but not in others. For example Platismatia erosa, P.
interrupta, P. formosana, and P. regenerans all produce caperatic acid
as the constant medullary constituent. In addition, in some individuals
of all these species, an unidentified substance could be demonstrated.
We do not know whether this substance occurred in all the specimens
and was not detected because of the limitations of our chemical tech-
niques or whether the compound is merely present in only part of the
populations. Since it occurs in addition to another substance (caperatic
acid) that is invariably present and since its real distribution is un-
known, this and the few other “accessory” substances found have not
been considered in the delimitation of species. Likewise, in Cetrelia
cetrarioides we are not sure whether some specimens contain both
perlatolic and imbricaric acids although it is certain that most specl-
mens contain only one substance or the other. Therefore, in this in-
stance we have not used chemistry in the taxonomy because we feel
that the chemical variation in this widespread species is still too
inadequately explained.

The above examples demonstrate how the apparent absence of
chemical constituents in individuals and in species has been evaluated.

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 489

The second problem, interpreting for taxonomic purposes the level of
biological significance of chemical variation, is far more difficult. Since
little has been definitively proved concerning the biosynthesis of the
substances involved, we cannot estimate how different biologically the
chemical races of morphologically similar lichens really are. Five main
categories of evidence, however, lead to the inevitable conclusion that
in morphologically similar or identical populations the largest chemical
variations (those based on the constant chemical components) reflect
major genetic differences; namely: (1) the orderly, nonrandom replace-
ment series that the major medullary compounds regularly display;
(2) the fact that chemically different populations usually show differ-
ent geographic distributions; (3) the observation, noted repeatedly in
many genera, that where geographic ranges overlap and where chem-
ically different populations become sympatric—and where conse-
quently chemically different plants may even grow side by side—no
chemically intermediate individuals are found; (4) the fact that in
localities where the chemical races are sympatric and where some
chemically different plants do exist side by side, the chemical races
as a whole select different habitats (W. L. Culberson and C. F. Cul-
berson, 1967); and (5) the lack of any indication whatever that en-
vironment has anything to do with the qualitative presence of the
major medullary constituents.

For a few lichen substances, there is some biosynthetic information
which helps in taxonomic evaluations. For example, it is believed that
stictic acid is derived from norstictic acid by a simple, one-step methyl-
ation reaction. In a recent study of the Ramalina siliquosa group
(W. L. Culberson, 1967), no taxonomic importance was attached
to the demonstration of norstictic acid in some of the stictic acid-
producing individuals because norstictic acid is the biochemically ex-
pected precursor of stictic acid. Unfortunately, much of our knowledge
of the biosynthesis of the substances produced in the genus Cetrelia is
less precise, but it should be pointed out that in the present study we
have not involved in the defining of species limits any pairs of sub-
stances believed to have such an intimate biosynthetic relationship as
that of norstictic and stictic acids. It is precisely in the area of inter-
preting the biological significance of chemical variation that the lichen-
ologist of the future, with a more precise knowledge of the chemical
relationships of the compounds, will hopefully be able to distinguish
significant from inconsequential chemical differences just as we hope
today to distinguish significant from inconsequential morphological
differences.

490 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM
Acknowledgments

This research was supported by National Science Foundation Grant
GB-1239 and by U.S. Public Health Service Research Grant GM-
08345 from the Division of General Medical Sciences, National Insti-
tutes of Health. We also wish to thank Professor Emeritus Dr.
Yasuhiko Asahina for the many kindnesses extended to us during the
months that we worked in his laboratory in Tokyo. The photographs
of anatomical sections were taken by Miss Barbara Fox and those of
herbarium specimens by Mr. Charles Racine.

Taxonomic Treatment
Cetrelia Culb. et Culb., gen. nov.

Thallus (5—)7-24(-26) cm. latus; laciniae 0.5-2.0(-2.5) cm. latae.
Superficies superior pallide cinerea, albida, vel apud exemplaria vetera
herbarii pallide citrella, pseudocyphellata, isidiata vel sorediata.
Superficies inferior picea, rhizinata, saepe punctata. Cortex superior
15-30(—34) crassus, prosoplectenchymatosus; medulla 85—230(—264) u
crassa, alba; cortex inferior (10—)12—-25(-31)y crassus. Apothecia
(0.1-)0.5-2.0(—4.2) em. lata, submarginalia, vulgo perforata; hyme-
nium (53—)62-110(-124) crassum, asci IKI-+ caerulescentes; strata
subhymenialia (35-)40-78(-84)yn, IKI—, stratum conspicuum al-
garum fere tegentia. Sporae 8-nae, ellipsoideae, 11-22(-25) X6-12u.
Pycnidia marginalia; conidia 1-1.5%(3-)4-6y, recta, extremis
nonnihil inflatis. Typus generis C. cetrariovdes est.

Thallus (5-)7-24(-26) em. broad; lobes 0.5-2.0(-2.5) cm. broad.
Upper surface ashy white or tan, pseudocyphellate, isidiate or soredi-
ate. Lower surface black, rhizinate, punctate in some species. Upper
cortex 15-30(-34)y thick, prosoplectenchymatous; medulla 85-230
(264) u, white; lower cortex (10) 12-25(-31)y thick. Apothecia (0.1—)
0.5-2.0(-4.2) em. broad, submarginal, usually perforate; hymenium
(53-)62-110(-124)y, the asci IKI+ blue; subhymenial layers (35-)
40-78(-84)y, IKI—, usually underlain by a conspicuous layer of
algae. Spores 8, ellipsoid, 11-22(-25)6-12y. Pyenidia marginal;
conidia 1—1.5X (3—)4-6y, rod-shaped, the ends slightly inflated.

Constituents: atranorin and alectoronic, anziaic, a-collatolic, imbri-
caric, microphyllinic, olivetoric, or perlatolic acids.

None of the species of the Parmelia cetrarioides group, recognized
here as the genus Cefrelia, is the type of a validly described genus
which could be used for them. Although Cetraria sanguinea Schaer., a
member of the new genus, was included along with a Parmelia in the
genus Aspidelia Stirt., that generic name is invalid (W. L. Culberson,

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 49]

1966a). The name Cetrelia is a fanciful concoction from Cetraria and
Parmelia, the genera in which most of the species had previously been
placed.

Artificial Key to the Species of Cetrelia

1. Thallus sorediate or isidiate or both.
2. Thallus sorediate, primarily along the margins of the lobes, not isidiate.
3. Medulla C+ pink or pinkish-orange, olivetoric acid present; widely
distributed... . .. . . 11. C. olivetorum
3. Medulla C—, olivetoric acid absent; widely distributed.
4. Medulla KC— or KC+ dirty pinkish, alectoronic acid and a-collatolic
acids absent; widely distributed, common in Europe but rare in North
America.
5. Medulla usually KC—, perlatolic acid present.
3. C. cetrarioides sens. str.
5. Medulla usually KC+ arty pinkish or rarely KC—, imbricaric acid
present. . . . . 38. C. cetrarioides sens. lat.
4. Medulla KC+ bright pink or - red, alectoronic and a-collatolic acids
present; widely distributed, common locally in eastern North America
but rare in Europe. ............. 4 C. chicitae
2. Thallus isidiate, not sorediate.
6. Isidia accompanied by conspicuous dorsiventral lobulae in tufts on the
thallus surface or as a fringe along the margins of the thalluslobes . . 9
6. Isidiate plants without dorsiventral lobulae.
7. Medulla C+ red, anziaic acid present; isidia often poorly developed;
Formosa and Japan... .. . 8 C. isidiata
7. Medulla C—, KC+ pink or red, alectoronic and a-collatolic acids pres-
ent; isidia well developed and not at all sorediose through most of the
range but granular, poorly developed and sorediose in the southern
part (Philippines); Asia. ........ .. 2. C. braunsiana
1. Thallus neither sorediate nor isidiate.
8. Thallus with dorsiventral lobulae in tufts or clusters on the upper surface or
as a fringe along the margins of the lobes.
9. Medulla C— ; lobulae well developed and abundant.
10. Microphyllinic acid present; Japan and southeastern Asia.
9. C. japonica
10. Imbricaric acid present; China and Formosa . . 14. C. sinensis
9. Medulla C+ pink or red; lobulae well developed or poorly developed,
infrequent, granular, and isidioid.
11. Olivetoric acid present; Japan, Formosa, China, and the Himalayas.
12. C. pseudolivetorum
11. Anziaic acid present; mountains of southern and southeastern Asia
and Japan. ................ 18. C. sanguinea
8. Thallus without lobulae.
12. Pseudocyphellae very small, rarely reaching 1 mm. long, simple and punc-
tiform (Plate 12, Figure 52).
13. Medulla C+ pink or red.
14, Olivetoric acid present; mountains of southwestern China.
6. C. davidiana
14, Anziaic acid present; mountains of southern and southeastern Asia
and Japan. ... . . . . 18. C. sanguinea (some forms)
13. Medulla C— ; perlatolic or r imbriearic acid present.

492 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

15. Lobes relatively thick and uniformly colored; pyenidia conspicuously

distributed along margins cf the lobes; on trees, southwestern China.

7. C. delavayana

15. Lobes relatively thin, the margins darker than the center; pycnidia

unknown; on tundra soil in Alaska. . . . . . 1. C. alaskana

12. Pseudocyphellae very large, some more than 1 mm. long, compound and

irregular (Plate 12, Figure 53).

16. Medulla KC-+ pink, alectoronic and a-collatolic acid present; Japan,

Formosa, and southwestern China... ... . . 10 C. nuda

16. Medulla KC—, imbricaric acid present; southwestern China and Nepal.

5. C. collata

1. Cetrelia alaskana (Culb. & Culb.) Culb. & Culb., comb. nov. PuaTeE 14

Cetraria alaskana Culb. & Culb., Bryologist 69:200. 1966. Type: U.S.A.,

Alaska, Cape Sabine, Thomson, Lich. Arct. 13 (DUKE, holotype; COLO,
LD, MSC, WIS, WIS-Herb. Thomson, isotypes).

Thallus medium, 5-7 em. broad, pale, yellowish-tan, or brownish,
often mottled with black, the margins distinctly brown. Lobes 1-1.5
em. broad, smooth, not pruinose, esorediate, pseudocyphellate, the
pores minute. Lower surface jet-black throughout or the margins
chestnut brown, smooth, not punctate, rhizines few, often very short
and papilliform. Upper cortex 16-29 thick; medulla 86-135 thick;
lower cortex 16-23, thick. Apothecia and pycnidia unknown.

MEDULLARY REACTIONS.—K—, C—, KC— or KC4+ pale pink,
PD-.

ConstituENTs.—Imbricaric acid and atranorin.

Distripution (Map, Figure 10).—The west coast of Alaska on the
Bering Sea and Arctic Ocean; in the tundra.

Cetrelia alaskana is a conspicuous, large-lobed lichen inhabiting plant
debris in the Alaskan tundra. It has doubtless been collected more
frequently than the present records indicate and has consistently been
mistaken for other species. The fact that it may be locally abundant is
clearly shown by the fact that at Cape Sabine, the type locality,
Thomson collected enough of the species to issue in his exsiccat
series, Lichenes Arectict.

Cetrelia alaskana resembles C. cetrarioides more than it does any
other species. Both have characteristic, minute, easily overlooked
pseudocyphellae in the upper cortex. Cetrelia alaskana, however, can
easily be distinguished from C. cetrarioides by its lack of soredia.

Cetrelia alaskana has also been mistaken for Asahinea chrysantha
(Tuck.) Culb. & Culb. which occurs in the same region and the two
species can probably be found together in the field. Cetrelia alaskana
always has some rhizines on the lower surface even though they may
not be produced in abundance while the lower surface of A. chrysantha
is invariably naked. Asahinea chrysantha also contains usnic acid in the
upper cortex and is consequently yellow-colored in contrast to the
usnic acid-lacking, tan-colored upper cortex of C. alaskana. The

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 493

medulla of A. chrysantha contains alectoronic acid and is consequently
strongly fluorescent in ultraviolet light while the medulla of C. alaskana
contains imbricaric acid and is not fluorescent. A detailed account of
Asahinea and its relationships to other groups in the Parmeliaceae has
already been given (W. L. Culberson and C. F. Culberson, 1965).

CHEMICAL ANALYSIS.—All 14 samples of C. alaskana tested micro-
chemically contained atranorin and imbricaric acid. The identity of
imbricaric acid was definitely established by the extraction and de-
tailed analysis of a small sample (C. F. Culberson and W. L. Culber-
son, 1966). The substance responsible for the KC+ reaction observed
on the thallus could not be identified.

ADDITIONAL SPECIMENS EXAMINED:

U.S.A.: ALasKa: Pitmegea River, 15 mi. upstream from Cape Sabine, 68°48’N,
154°20’W, Thomson, Lich. Arct. 13. (Type collection, see above.) Ogotoruk Creek
Drainage, 68°05’N, 165°32-47’W, Johnson et al. 21 (COLO). 40 mi. east of Cape
Lisburne, Cantlon & Gillis 57-136A (MSC). Kuskakwin, Goodnews Bay, Geist,
1933 (Herb. Degelius, LD). Lower Telephone Creek, Palmer 585 (US, WIS).
Peace River, Palmer 614 (US, WIS).

2. Cetrelia braunsiana (Miill. Arg.) Culb. & Culb., comb. nov. PuatE 15

Parmelia braunsiana Mill. Arg., Flora 64:506. 1881. Type: Japan, Tokyo,
Brauns 2 (pro parte) (G, holotype).

Platysma braunsianum (Mill. Arg.) Hue, Nouv. Arch. Mus., IV, 1:210. 1899.

Cetraria braunsiana (Mill. Arg.) Zahlbr., Bot. Mag. [Tokyo] 41:353. 1927.

Parmelia braunsiana f. isidiosa Mill. Arg., Flora 64:506. 1881. Type: Japan,
Tokyo, Brauns 2 (pro parte) (G, holotype).

Platysma braunsianum f. isidiosum (Mill. Arg.) Hue, Nouv. Arch. Mus., IV,
1:210. 1899.

Cetraria braunsiana f. isidiosa (Mill. Arg.) Zahlbr., Catal. Lich. Univ. 6:284.
1930.

C. collata f. isidiata Asah., Journ. Jap. Bot. 10:475. 1934. Type: Japan, Mus-
ashi Prov., Mt. Buko, Asahina 71b (TNS, lectotype).

Thallus medium to large, 12-24 cm. broad; lobes 0.5-1.7 cm. broad.
Upper surface gray or ashy-white, uniformly brownish or tan in old
herbarium specimens, the margins + ascendent and commonly isidi-
ate, at least in part, the tips pruinose, pseudocyphellate, the pores
small, punctiform to irregular, rarely more than 1 mm. broad, often
associated with tiny, granular, delicately coralloid isidia. Lower surface
black, the margins brown or grayish like the color of the upper surface,
not punctate; rhizines black. Upper cortex 19-25y thick; medulla
109-264 thick; lower cortex 16-22y thick. Apothecia rare, submar-
ginal, perforate, about 0.5 mm. broad; hymenium 93-109, thick;
subhymenial layers 40-53y thick, underlain by a dense layer of algae.
Spores 8, ovoid, 12-158-9u. Pycnidia rare, marginal, black, prui-
nose; conidia 1><4—6u, rod-shaped, the ends slightly enlarged.

MEDULLARY REACTIONS.—K —, C—, KC+ pink, PD—.

ConsTITUENTs.—Alectoronic and a-collatolic acids and atranorin.

277-920 O - 68 - 4

494 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Fiaures 7-11. Distributions. 7. Cetrelia olivetorum (Nyl.) Culb. & Culb. 8. Cetrelia
isidiata (Asah.) Culb. & Culb. 9. Cetrelia pseudolivetorum (Asah.) Culh. & Culb.
10. Cetrelia alaskana (Culb. & Culb.) Culb. & Culb. 11. Cetrelia braunsiana (Mill.
Arg.) Culb. & Culb.

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 495

Distribution (Map, Fig. 11)—Widespread in Hokkaido and
Honshu, Japan, in the high mountains of Formosa, Luzon, Philippines,
Yunnan, China, and in the Himalayas; on tree trunks and over
boulders.

Cetrelia braunsiana, C. chicitae, and C. nuda have been recognized in
recent years by Japanese botanists as Cetraria collata {. isidiata Asah.,
C. collata sens. str. (=f. collata), and C. collata f. nuda Hue respectively.
They are, however, quite distinct morphologically, although chemi-
cally all produce atranorin and alectoronic and a-collatolic acids.
Cetrelia braunsiana is isidiate, C. chicitae is sorediate, and C. nuda
produces neither soredia nor isidia, but it has pseudocyphellae different
from those of the other two species.

The differences among these three species, and the consequent justi-
fication for regarding them as distinct and of specific rank are sum-
marized in Table 5.

TABLE 5.—The percentages of Japanese specimens of Cetrelia braunsiana, C. chicitae, and C. nuda showing
various morphological traits and the geographic distribution of the species

braunsiana chicitae nuda
Total number of specimens examined 74 35 51
Small pseudocyphellae (less than 1 93 100 10
mm.)
Large pseudocyphellae (more than 1 7 0 90
mm.)
Isidia 100 0 0
Soredia 0 100 0
Apothecia 4 17 83
Pycnidia 7 14 100
Pyenidia pruinose 0 0 24
Lobes pruinose (at least in part) 69 43 14
Lower surface punctate 1 6 59
Japanese range Hokkaido, Hokkaido, Southern
Honshu Honshu Honshu,
Shikoku,
Kyushu

Japanese specimens belonging to this group never produce both
isidia and soredia and most (90%) of the specimens with neither isidia
nor soredia have large pseudocyphellae rather than the small ones
characteristic of the other two species. It should be pointed out, how-
ever, that,the isidia in Cetrelia braunsiana are often produced in very
small numbers. Invariably, however, if Japanese specimens belonging
to this group and having small pseudocyphellae and no soredia are
examined carefully under the dissecting microscope, at least a few

496 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

delicate marginal isidia will be found. The type specimen itself of C.
braunsiana has very few isidia.

It is apparent from Table 5 that Cetrelia braunsiana and C. chicitae
are more similar to each other than they are to C. nuda. The nonisidi-
ate, esorediate C. nuda always produces pycnidia, which are often
pruinose, and commonly has apothecia and a punctate lower cortex.
It actually belongs to the Cetrelia sanguinea group and is not partic-
ularly closely related to C. braunsiana and C. chicitae at all, although
its taxonomy has been confused with theirs.

As pointed out above, it is generally possible to distinguish Japanese
specimens of Cetrelia braunsiana from those of C. chicitae by careful
examination of the plants for isidia. Farther south, and especially in
the Philippines, the distinctions may become more difficult. In some
Philippine specimens the margins of the lobes are densely covered with
soredia-like structures that become isidioid and granular, apparently
by compaction and the formation of a cortexlike surface. These
anomalous specimens are otherwise well developed and may represent
a unique local race even though essentially typical specimens of both
C. braunsiana and C. chicitae are also known from the Philippines.
This local problem in distinguishing between some specimens of C.
braunsiana and C. chicitae, however, should not be taken to indicate
that these two species differ only slightly because in reality they differ
quantitatively in the expression of many traits (Table 5). Neverthe-
less, the identification of certain isolated specimens from the Philip-
pines may depend entirely upon the presence of isidia as opposed to
soredia.

The type specimen of Parmelia braunsiana Mill. Arg. is isidiate
and the type of P. braunsiana f. isidiosa Mill. Arg., really part of the
same collection, is somewhat more isidiate. In view of the great varia-
tion in the production of isidia in the Japanese population of this
species, there is no basis for recognizing the forma.

Cetrelia braunsiana is part of the Sino-Japanese floristic element
with a range extension into Formosa and the Philippines. In spite of
its relatively close relationship to the very widespread species C.
chicitae, no specimens of it from outside the Orient are known. In Japan
(. braunsiana is widely distributed and is found at lower elevations as
well as in the mountains. Its broader amplitude of ecologic tolerance,
as compared to that of most of the Japanese species of Cetrelia, is also
seen in its general abundance in Japan as well as in its more extensive
vertical geographic range there.

CHEMICAL ANALYsIs.-—Alectoronic acid, a-collatolic acid, and atra-
norin were identified in the 32 samples tested. The tests were made
without difficulty and, in most cases, directly on the acetone extract,
the concentration of a-collatolic acid being sufficient to give a good

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 497

chromatogram. Although many of the residues were yellow, no known
pigment could be identified and all tests for usnic acid were negative.

NoMENCLATURE.—In the Japanese botanical literature the name
Cetraria braunsiana (Mill. Arg.) Zahlbr. has been used consistently to
refer to the species treated here as Platismatia interrupta Culb. & Culb.
A simple but widespread misunderstanding of the application of the
name apparently accounts for this.

This species was named for David Brauns (1827-1893), Professor
of Geology at the University of Halle, who made the original collection
near Tokyo in 1881.

REPRESENTATIVE SPECIMENS EXAMINED:

JAPAN: Hoxxkarpo: Sapporo, Faurie 9212 (PC). Prov. Ishikari: Daisetsu,
Kuro-dake, Asahina, 1937 (TNS, two collections); Hot Spring Matsuyama,
Asahina, 1935 (TNS). Prov. Kitami: Lake Abashiri, Asahina & Togashi, 1953
(TNS). Prov. Kushiro: Lake Akan, Asahina, 1953 (TNS).

Honsuuv: Hirosaki, Faurie 196 (PC). Prov. Bingo: Sera-gun, Kamiyama-mura,
Sato, 1932 (TNS). Prov. Echigo: Iwafune-gun, Arakawa-cho, Togashi, 1956
(TNS). Prov. Echizen: Imadate-gun, Okamoto-mura, Hosoi & Ui, 1935 (TNS).
Prov. Ettyu: Magawa Pass, Asahina, 1936 (TNS, two collections). Prov. Harima:
Shiso-gun, Mikawa-mura, Funakoshi, Utsum7z, 1951 (TNS). Prov. Inaba: Karo,
Ikoma, 1953 (TNS). Prov. Iwaki: Matsukawa-ura, Kurokawa 58086, 58087
(TNS). Prov. Iwashiro: Ura-bandai, Asahina, 1954 (TNS). Prov. Izu: Nirayama,
Honryu-ji Temple, Asahina, 1956 (TNS). Prov. Kaga: Enuma-gun, Hot Spring
Yamanaka, Togashi, 1959 (TNS). Prov. Kai: Mt. Minobu, Togashz, 1961 (TNS,
three collections). Prov. Kii: Mt. Koya, Numajiri 699 (TNS). Prov. Kozuke:
Tano-gun, Ueno-mura, Shio-no-sawa Pass, Togashi, 1956 (TNS). Prov. Mikawa:
Mt. Horaiji, Asahina, 1956 (TNS, two collections). Prov. Mino: Seki-ga-hara,
Asahina, 1938 (TNS). Prov. Musashi: Chichibu, Ryogami-mura, Hyugaotani,
Kurokawa 550534 (TNS); Mt. Buko, Asahina 71b (TNS). Prov. Owari: Inuyama,
Asahina, 1938 (TNS). Prov. Rikuchu: Isawa, Mt. Shishihana, Suzuki 491 (TNS).
Prov. Rikuzen: Tamatsukuri-gun, Naruko-cho, Asahina & Togashi (TNS). Prov.
Sagami: Odawara, Mt. Daiyuzan, Saijo Temple, Togashi, 1956 (TNS). Prov.
Settsu: Arima-gun, Dojo-mura, Togashi, 1952 (TNS). Prov. Shimotsuke: Nikko,
Lake Yuno-ko, Asahina, 1931 (TNS, two collections). Prov. Shinano: Lake
Shirakaba, Asahina & Togasht, 1958 (DUKE); Mt. Kimpu, Asahina, Togashi &
Nuno (TNS, 2 collections); Suwa-gun, Izumino-cho, Asahina & Togashi, 1958
(TNS); Mt. Yatsu-ga-take, Hot Spring Shibu-no-yu, Kurokawa 58198 (TNS) and
58199 (DUKE); Mt. Tadeshina, Kurokawa 51746, 71864, 58374, 58375 (TNS) and
58375 (MSC); Mt. Norikura, Asahina, 1952 (TNS). Prov. Suo: Hikarishi, Muro-
zumi, Asahina & Togashi, 1959 (TNS). Prov. Suruga: Lake Saiko (Nishi-no-umi)
near Mt. Fuji, Culberson & Culberson 10684 (DUKE). Prov. Tango: Miyazu-shi,
Nariai Temple, T’ogashi, 1956 (TNS). Prov. Uzen: Goshiki-numa, Suzuki, 1954
(TNS). Prov. Wakasa: Mt. Aoba, Togashi, 1956 (TNS). Prov. Yamashiro: Atago-
gun, Hanaze-mura, Mt. Taihi, Togashi, 1955 (TNS).

SHIKOKU: Prov. Tosa: Hata-gun, Okata-cho, Kurokawa 550231, 550232 (TNS);
Mt. Yokogura, Fujikawa, 1931 (TNS).

FORMOSA: Mt. Ali, 2,200 m., Kurokawa 13 (TNS). Mt. Ba-Hsen, 2,180 m.,
Jen-rong Wang 133 (DUKE). Piyanan Pass, 1,600-1,900 m., Kurokawa 821 (TNS).

PHILIPPINES: Luzon: Benguet Prov.: Mt. Pulog, McGregor, 1909 (TUR).
Mountain Prov.: Mt. Data, Hale 26019, 26150, 26401 (US).

498 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

CHINA: Yunnan: Kou-toui, above Mo-so-yn,? Delavay 3009 (PC).

NEPAL: Mewakhola Valley, 11,000 ft., Awasthi 2265 (UPS). Above Sauwala
Khola, 10,500 ft., Stainton et al. 4379b (pro parte) (BM).

INDIA: Northwestern Himalayas: Kamaon, Almora District, Dhakuri Ridge,
9,500 ft., Awastht & Awasthi 633 (UPS).

3. Cetrelia cetrarioides (Del. ex Duby) Culb. & Culb., comb. nov. PLaTE 15

Parmelia perlata 6 cetrarioides Del. ex Duby, Bot. Gall., ed. 2, 601. 1830.
Type: France, Mont d’Or, Delise, 1828 (STR, lectotype). The lectotype,
designated by Hale (1960), was not seen, but an extract of a fragment of it
prepared by Hale was studied microchemically by us (see below).

P. cetrarioides (Del. ex Duby) Nyl., Flora 52:289. 1869.

P. monochorum Zahlbr., Symb. Sin. 3:191. 1930. Type: China, Szechwan,
Doko Pass southwest of Muli, Mazzetti 7399 (WU, holotype).

P. cetrarioides f. bisoralifera Laven, Repert. Spec. Nov. Fedde 50:294. 1941.
Type: Bavaria, prope lacum Lautersee, Laven (not seen).

Thallus medium to large, 7-16 cm. broad; lobes (0.5-)1—1.5 cm.
broad. Upper surface uniformly tan or pale, pseudocyphellate, the
pores very small, rarely reaching 0.5 mm., the margins of the lobes
finely and usually abundantly sorediate. Lower surface jet-black, the
margins brown, usually punctate with many small pores; rhizines few,
black, rarely pale. Upper cortex (16-)19-28(-31)u thick; medulla
(62-)90-155(-171)p thick; lower cortex 16-25(—28)u thick. Apothecia
extremely rare, 1-2 mm. broad (immature, the only ones seen in situ).
Spores ovoid, 12-14 19-24 (seen in a single apothecial fragment).
Pyenidia unknown.

MEDULLARY REACTIONS.—K—, C—, KC— or KC+ pinkish, PD—.

ConstITvENTS.—Perlatolic or imbricaric acid and atranorin.

Distrisution (Maps, Figures 12-13).—Eastern North America,
Japan, the mountains of southern Asia, and western Europe; usually
in shady places, on the bark of trees, rock faces, and over boulders.

The Cetrelia cetrarioides complex includes four chemically distinct
populations. The population producing atranorin and olivetoric acid
in the medulla is slightly but distinctly different morphologically from
the other three and clearly represents a different taxon, here treated as
the species Cetrelia olivetorum (see page 515). The remaining three
chemical types consist of plants which produce atranorin and (1) perla-
tolic acid, (2) imbricaric acid, or (3) alectoronic and a-collatolic acids.
In examining a large series of herbarium specimens one may be tempted
to see subtle morphological “tendencies”? among the chemical races,
but the accurate identification of individual specimens ultimately
depends upon chemical analysis.

2The spelling of Chinese place names in the citation of specimens taken by
Delav ay follows Bretschneider’s (1898) excellent summary of Delavay’s botanical
work. Our only modification of Bretschneider’s spellings has been to hyphenate
compound names.

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 499

*
AY
——

wy
oI

x
i
i

ier a
9
O.
x
al

y?
ans
eR

Pane
OE
Py

id.

Figures 12-14. Distributions. 12-13. Cetrelia cetrarioides (Del. ex Duby) Culb. &
Culb. (two chemical strains). 12. Cetrelia cetrarioides (Del. ex Duby) Culb. &
Culb. sens. str., the strain with perlatolic acid. 13. Cetrelia cetrarioides (Del. ex
Duby) Culb. & Culb. sens. lat., the strain with imbricaric acid. 14. Cetrelia chi-
citae (Culb.) Culb. & Culb.

In the present study the plants containing alectoronic and a-colla-
tolic acids are assigned to Cetrelia chicitae and the perlatolic and imbri-
caric acid-containing plants to C. cetrarioides sens. lat. It may be
thought arbitrary and untenable, in the absence of a unique morphol-
ogy, to consider the plants that produce alectoronic and a-collatolic
acids to represent a distinct species while keeping together taxonomi-

500 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

cally those that produce perlatolic acid and those that produce imbri-
caric acid. As pointed out in a previous study of the chemistry and geo-
eraphic distribution of the components of the cetrariordes complex
(Culberson, 1958), plants assigned to C. chicitae represent the bulk of
the P. cetrarioides complex in North America, while in Europe the
complex is made up almost entirely of perlatolic and imbricaric acid-
producing populations. While the three chemical races might be nearly
sympatric purely from the point of view of qualitative range (although
the worldwide distribution here is poorly known), the quantitative
distributions are clearly different. For this reason and since the produc-
tion of the two compounds alectoronic acid and a-collatolic acid seems
to be a relatively large departure from the chemistry of the other
chemical types, this type is regarded as a distinct species.

The retention of the remaining two types, the perlatolic acid-con-
taining plants and the imbricaric acid-containing ones, within the
single species Cetrelia cetrarioides is less satisfactory. Unlike the chemi-
cal species C. chicitae, these two races do not seem to be significantly
more abundant in one part of their range than in other parts. Although
by proper chemical analyses it is possible to identify which of the acids
is present in a given specimen, there are a few imbricaric acid-contain-
ing specimens in which our microchemical and chromatographic tests
cannot assure with absolute certainty the absence of perlatolic acid
(although an alternate explanation is possible; see the chemical dis-
cussion below). In view of the essentially sympatric distribution of
these two chemical types and the possibility at least that some indi-
viduals may actually produce both perlatolic and imbricaric acids, it
does not seem desirable at present to try to accommodate the chemical
variation in the taxonomy.

Some nonchemical aspects of the taxonomy in the Cetrelia cetrart-
oides complex are also still not completely explored. For example, two
anomalous, well developed, imbricaric acid-producing specimens from
Sikkim in the Himalayas, where the genus Cetrelia is exceptionally well
represented, lacked soredia altogether. The plants are relatively small-
lobed and by habit alone are obviously closely allied to C. cetrarioides.
Since typical specimens of the imbricaric acid-producing race of C.
cetrarioides are found in the same Himalayan region, these specimens
have provisionally been assigned to C. cetrarioides in spite of their lack
of soredia. They are pointed out here only to call attention to the fact
that additional studies on new collections of the C. cetrarioides complex
from the Himalayas might necessitate additional modifications in the
taxonomy.

The tundra-inhabiting species Cetrelia alaskana might be mistaken
for C. cetrarioides because superficially it looks similar. Like many
individuals of C. cetrarioides, C. alaskana produces imbricaric acid, but

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 5()1

it is never sorediate while soredia are usually well developed in most
specimens of C. cetrariovdes. The two species also have strikingly dif-
ferent ecologies, and their geographic ranges are widely separated.

The soredia in Cetrelia cetrarioides are typically confined to the
margins of the lobes in long soralia. An anomalous form with conspic-
uous capitate soralia has been named as an infraspecific variation
(Laven, 1941). An analogous form bearing capitate soredial masses has
also been described for Platismatia norvegica. These rare, peculiar
forms are not considered taxonomically in this study because they
seem to represent pathological rather than natural growths.

In its broad sense, including both the chemical races that compose it,
Cetrelia cetrarwides is the most widely ranging species in the genus.
It is, of course, the very fact of this broad range and the accompanying
internal chemical variation that would speak against considering C.
cetrarwides a single species as it is regarded in the present study.

Cetrelia cetrarioides, like the closely related species C. olivetorum and
C. chicitae, is generally restricted to places with relatively moist, cool
climates. It is probably most abundant and best developed in the
British Isles and in other oceanic parts of western Europe. As would be
expected, the southernmost stations of its range on all continents are
in mountains.

Throughout much of its range Cetrelia cetrarioides is very infrequent.
Its extreme rarity in North America has already been reported (Cul-
berson, 1958), a rarity all the more noteworthy in view of the extreme
local abundance of the closely related species C. chicitae (see the dis-
cussion for that species). In parts of Europe C. cetrarioides is also very
rare. For example, in the southern parts of Norway (Lynge, 1921),
Sweden (Du Rietz, 1924), and Finland (Hakulinen, 1963)—regions
which have been intensively explored by lichenologists—it has been
found in only a few localities.? In Asia too the records are widely
scattered. The extreme rarity of the species of course means that it has
often gone unnoticed by collectors. The distribution maps given here
(Figures 12-13) are consequently only roughly approximate.

CHEMICAL ANALYSIS.—Sixty-nine samples of this species were tested;
35 contained atranorin and perlatolic acid and 34 contained atranorin
and imbricaric acid mixed with at least one unidentified phenolic sub-
stance. Perlatolic acid was identified by the crystals observed in the

3 No Scandinavian specimens of Cetrelia cetrarioides were seen in this study.
Although the species unquestionably occurs in southern Scandinavia, no localities
are given on the maps (Figures 12-13) because the chemical race to which such
specimens belong is unknown. The presence of C. cetrarioides sens. lat. in Scandi-
navia is acknowledged on the map in Figure 1 which shows the geographic distri-
bution of species numbers in the genus Cetrelia. The two species recorded there for
Scandinavia are C. cetrarioides and C. olivetorum.

502 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

GE and GAW solutions and by the straight extinction of these crystals
with crossed Nicols. Imbricaric acid crystals show oblique extinction
and a somewhat different morphology in these solutions.

All residues were chromatographed in NH,OH-n-butanol, and many
of them were hydrolyzed with 0.1 n Na:HPO, to detect imbricaric
acid. While perlatolic acid-containing samples were always KC—
(with the possible exception of one specimen), a definite to pale KC+
red reaction was observed on many imbricaric acid-containing thalli.
Imbricaric acid was identified by comparison with imbricaric acid in
Cetrelia alaskana (C. F. Culberson and W. L. Culberson, 1966). A
KC-+ reaction which has also been observed with C. alaskana is due to
an unidentified KC+ substance or to a low concentration of some C+
substance. Further studies are necessary to identify this material.

Asahina and Fuzikawa (1935) have suggested that imbricaric acid
and perlatolic acid may occur together, but their evidence is not con-
clusive. Hydrolysis of several imbricaric acid-containing residues
showed a spot by chromatography at an R, corresponding to that for
olivetol-2-carboxylic acid. This could be explained if perlatolic acid
were also present, but such a mixture would still not be KC+. Either
the samples containing imbricaric acid may also contain perlatolic acid
and still a third constituent to give the KC+ reaction, or these samples
contain imbricaric acid and a KC+ substance that produces olivetol-
2-carboxylic acid upon hydrolysis.

NomeEncLAtTuRE.—After a careful study of the appropriate speci-
mens from Delise’s herbarium, Hale (1960) was able to typify Parmelia
perlata B cetrarioides Del. ex Duby in the sense that the binomial P.
cetrarioides (Del. ex Duby) Nyl. has generally been used. Although we
were not able to examine the lectotype, Hale’s (1960) photograph of it
is unmistakable. From a microextract that he had made from it at the
time of his study and which he subsequently gave to us for testing, we
confirmed his report that this specimen contains perlatolic acid. In the
quite possible eventuality that a future interpretation based on addi-
tional evidence should recognize the imbricaric acid-producing race as
a distinct species, it should be noted that P?. monachorum Zahlbr. offers
apparently the oldest specific epithet for this variation; we have identi-
fied imbricaric acid and atranorin in the holotype.

Santesson (1942) noted that Pseudoparmelia pseudofallax Gyeln. is
also a synonym of “Parmelia cetrarioides,” but we have not seen the
type specimen and do not know to which chemical race it belongs.
Even if it belongs to the variant producing imbricaric acid, it could
not be used, since the epithet monachorum is older.

ADDITIONAL SPECIMENS EXAMINED:

Specimens of C. cetrarioides, the perlatolic acid-producing race:

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 5(03

U.S.A.: West Viratnra: Pendleton Co.: Panther Knob, Hale 14264 (DUKE).

WasHINGTON: Snohomish Co.: Marysville, Grant, 1926 (MICH), 1927 (MICH).

CHINA: Yunnan: Kou-toui, above Mo-so-yn (Lan-kong), 3,000 m., Delavay
1887 (PC); Yen-tze-hay, 300 m., Delavay, 1887 (PC); Yen-tze-hay, Lan-kong.
Delavay, s.d. (PC) ; Koua-la-po, Delavay 1595 (pro parte) (PC); Koua-la-po, Ko-kin,
3,000 m., Delavay, 1885 (PC, 2 collections).

SIKKIM: Tonglo, 16,000 ft., Hooker 2083 (K).

POLAND: Mont. Carphatica, Mont. Gorganae, near Skole, Aasamaa Cet-302
(DUKE).

AUSTRIA: Hallstadt, ex herb. Bory de St-Vincent, 1805-1806 (PC).

GERMANY: Bayern: Obere Hochebene, Wornbrunn, Schwind, 1925 (BP).

SWITZERLAND: Vaugeron, near Genéve, Schleicher 80 (DUKE). Ticino:
Fusio, Maggia Valley, Barkman 4290 (DUKE).

FRANCE: CétTeEs-pvu-Norp: Forét de Lorges, des Abbayes, 1929, 1930 (Herb. des
Abbayes, 2 collections).

FInisTérE: Forét de Coatloc’h, Piquenard, 1900 (PC); the same, des Abbayes,
1930, 1933 (Herb. des Abbayes, 2 collections).

AVEYRON: Forét de Serre, Barreau, 1831 (PC, 3 collections).

SaRTHE: Forét de Perseigne, des Abbayes, 1936 (Herb. des Abbayes).

VoscEs: Docelles, Harmand, Lich. Loth. 288 (pro parte) (PC).

Puy-pE-D6mE: Bois de Job, near Ambert, Gasilien, 1883 (Herb. des Abbayes).

HavuTeEs-PyrénfEs: Pauze, near Cauterets, collector unknown (PC).

WALES: MERIONETHSHIRE: Penmaenpool, Culberson & Culberson 11564
(DUKE).

Specimens of C. cetrarioides sens. lat., the imbricaric acid-producing
race:

CANADA: Quésec: Labelle Co.: North of La Conception, Hermann 15783
(DUKE).

JAPAN: Fujiyama; Ityigome-Magajisi, Almquist, 1879 (S); the same, Umaga-
yeshi, Almquist, 1879 (S). Togakushi, Faurie 777 (PC).

CHINA: Yunnan: Hee-chan-men, Delavay, 1887 (PC); Santo-ha-ho, 3,000 m.,
Delavay, 1888 (PC, 2 collections); Lenago Pass between Mekong and Yangtse,
27°43'N, Gebauer, 1914 (W). SzEcHwaN: Doko Pass SW of Muli, Handel-M azzetti
7399 (UW).

SIKKIM: Lachan, 13,000 ft., Hooker 2086 (K). Wallanchoon 13,000 ft., Hooker
1814, 2082 (K).

POLAND: Mont. Carphatica, Mont. Gorganae, near Skole, Aasamaa Cet-301
(DUKE).

GERMANY: Miinchen, Arnold, Lich. Monac. Exs. 324a, (pro parte), 324b, (pro
parte) (PC).

CZECHOSLOVAKIA: Nizke Poloniny, Nov4 Sedlica, PiSwt, Lich. Slovak. Ezs.
13 (DUKE).

FRANCE: Catvapos: Forét de St-Tever, collector unknown, 1890 (PC).

SEINE-ET-MarneE: Fontainebleau, Nylander, s.d. (PC).

Puy-pE-D6meE: Besse, des Abbayes, 1933, 1935 (Herb. des Abbayes, 2 collections).

HAvuTE-VIENNE: Montagnes de Goullet, Lamy de la Chapelle, 1874 (PC);
Goullet, bei St. Sylvestre, Lamy de la Chapelle, as Arnold, Lich. Exs. 582 (PC).

PyRE£NE£EsS-ORIENTALES: La Massane, des Abbayes, 1933 (Herb. des Abbayes).

Hautes-Pyrinfes: Pauze, near Cauterets, collector unknown, s.d. (PC).

SCOTLAND: Aray.usuire: Ardnamurchan Peninsula, Salen, Culberson &
Culberson 11816 (DUKE).

504 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

4. Cetrelia chicitae (Culb.) Culb. & Culb., comb. nov. PLATE 14
Cetraria chicitae Culb., Bryologist 68:95. 1965. Type: U.S.A., West Virginia,
Hale, Lich. Amer. Exs. 56 (DUKE, holotype).

Thallus medium to large, 10-16 cm. broad; lobes 0.7-1.7 cm. broad.
Upper surface light gray or ashy, uniformly brown or tan in old her-
barium specimens, smooth, often pruinose at the tips, the margins
erupting with fine soredia, pseudocyphellate, the pores small, rarely
exceeding 1 mm. Lower surface jet-black, the margins brown or colored
like the upper surface and in some specimens wrinkled, the crests of
the folds becoming punctate. Upper cortex 16-26(—31)y thick; medulla
85-186 (-260) u; lower cortex, 10-26 thick. Apothecia very uncommon,
perforate, 4-5 mm. broad; hymenium 62-78u thick; subhymenial
layers 62-78u thick. Mature spores unknown. Pycnidia rare, marginal ;
conidia 1 <5—6u, rod-shaped, the ends somewhat enlarged.

MEDULLARY REACTIONS.—K —, C—, KC+ pink, PD—.

ConstiTuENTS.— Alectoronic acid and a-collatolic acid and
atranorin.

Distrisution (Map, Figure 14).—Southern Ontario and New
Brunswick, Canada, and the adjacent United States, south in the
Appalachian Mountains to North Carolina; southern Sakhalin and
Japan, and the mountains of Formosa, Luzon (Philippines), Sabah
(Borneo), and Java; on the trunks of trees and on rocks and over
boulders. A discussion of the range of this species is included in the
treatment for C. cetrarioides (p. 501).

Cetrelia chicitae is essentially indistinguishable morphologically
from C. cetrarioides, although the species differ in chemical constituents
and in their geographic distributions. For a full discussion of the rela-
tionship of C. chicitae to C. cetrarioides and to C. olivetorum, see the
treatment of C. cetrarioides (p. 498). The relationship of C. chiertae to
other species of the same chemistry for which it might be mistaken is
explained in the taxonomic discussion of C. braunsiana (pp. 495-496).

CHEMICAL ANALYSIS.—The 24 samples tested all contained atrano-
rin, alectoronic acid, and a-collatolic acid. Early in the study, 14 of
these specimens were tested by means of a single acetone extraction,
but later the method of double extractions with benzene and then ace-
tone was employed. Microcrystal tests in the GE, GAW, and GAo-T
solutions were always confirmed by chromatography to prove the
presence of a-collatolic acid. In the fragments tested the concentra-
tions of all three substances were always sufficient to give a good test
without the difficulties sometimes encountered in other species in
demonstrating small concentrations of a-collatolic acid or atranorin in
the presence of alectoronic acid.

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 5()5

ADDITIONAL SPECIMENS EXAMINED:

U.S.A. anp CANADA: Fora long list of selected specimens examined, see the
original description of this species (W. L. Culberson, 1965).
U.S.S.R.: Sakuatin Istanp: Takinosawa, Asahina, 1932 (TNS, 2 collections) ;
Aikawa, Asahina, 1932 (TNS).
KOREA: Locality unknown, Asahina, 1934 (TNS).
JAPAN: For a complete list of the Japanese specimens examined at the time
that the species was described, see W. L. Culberson (1965).
FORMOSA: Mt. Ali, 2,200 m., Kurokawa 12, 38 (TNS); Mt. Shin-Kao-San,
Mt. Chien-San, 3,100-3,300 m., Kurokawa 251 (TNS).
SABAH: Kinabalu National Park, near East Mesilau River, 1,700 m., Hale
28431 (US).
PHILIPPINES: Mountain Proy.: Mt. Data, 2,312 m., Hale 26358 (US).
JAVA: Res. Pasoeroean, Goenoeng Ardjoena, Tretes-Lalidjiwa-track, Du Rietz
95:15 (UPSV).
FRANCE: Vosges: Docelles, l’Encerf, Harmand, Lich. Loth. 288 (DUKE, PC).
5. Cetrelia collata (Nyl.) Culb. & Culb., comb. nov.
Platysma collatum Nyl., Flora 70:134. 1887. Type: China, Yunnan, Delavay
1590 (H-Herb. Nyl. No. 36116a, holotype).
Cetraria collata (Nyl.) Mill. Arg., Nouv. Giorn. Bot. Ital. 24:192. 1892,

Thallus medium, to 15.0 em. broad; lobes 0.8-2.5 cm. broad. Upper
surface pale brownish in old herbarium specimens (whitish when
fresh?), smooth, pseudocyphellate, the pores large, many becoming
confluent and exceeding 1 mm. Lower surface jet-black, punctate in
some specimens, the margins dark brown or colored like the upper
surface, relatively smooth, ascendent; rhizines few. Upper cortex
16-34 thick; medulla 109-2644 thick; lower cortex 16-25, thick.
Apothecia 0.7—-2.0 cm. broad, cup-shaped at first and expanded with
age, perforate or entire, the thalloid exciple wrinkled and abundantly
pseudocyphellate; hymenium 99-124, thick, subhymenial layers 35-
56u thick. Spores 8, ellipsoid, 15-20 X7-12y, the spore wall about 1.54
thick. Pycnidia large, marginal, unstalked; conidia about 1.5X6.0u,
rod-shaped, the ends often somewhat inflated.

MEDULLARY REACTIONS.—K—, C—, KC— or KC+ pinkish, P—,
UV-.

ConstTITUENTS.—Imbricaric acid and atranorin.

DistRIBUTION (Map, Figure 19).—The high mountains of Szechwan
and Yunnan, China, and Nepal; on the trunks and stems of trees and
shrubs.

Cetrelia collata might be mistaken for C. davidiana, C. delavayana, C.
nuda, or C. sanguinea. It differs from all these species, however, in
producing imbricaric acid rather than some other substance in the
medulla. It can easily be distinguished by its C— medullary reaction
from C. davidiana (with olivetoric acid) and C. sanguinea (with anziaic
acid), both of which are C+ pink or red in the medulla. From C. nuda
(with alectoronic and @-collatolic acid) and C. delavayana (with perla-

506

CONTRIBUTIONS FROM THE NATIONAL

HERBARIUM

15

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Figures 15-21. Distributions. 15. Cetrelia sanguinea (Schaer.) Culb. & Culb. 16. C.
japonica (Zahlbr.) Culb. & Culb. 17. C. nuda (Hue) Culb. & Culb. 18. C., sinensis
tulb. & Culb. 19. C. collata (Nyl.) Culb. & Culb. 20. C. davidiana Culb. & Culb.
21. C. delavayana Culb. & Culb.

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 507

tolic acid) it is best distinguished by the appropriate microchemical
tests.

CHEMICAL ANALYSIS.—AIl 17 samples tested contained imbricaric
acid and the residue from extracts gave straight needles of oblique
extinction in the GE and GAW solutions. Atranorin was easily identi-
fied by microcrystal tests. Most extracts were chromatographed before
and after base hydrolysis. The hydrolyzed residues frequently showed
an additional weak spot at an R; corresponding to olivetol-2-carboxylic
acid and most samples tested showed a definite KC+ pink reaction
apparently due to the presence of at least one unidentified substance.

ADDITIONAL SPECIMENS EXAMINED:

CHINA: Szecuwan: Mt. Lose-chan, south of Ningyiien, 3,950 m., Handel-
Mazzetti 1441 (WU).

Yunnan: Lidjiang, Mt. Ndaza Ko, 4,000 m., Rock, as Zahlbr., Lich. Rar. Exs.
322 (S, UPS, W); Snow Range, 4,300-12,000 ft., Rock, 1931 (W); Mt. Yilung-
schan, Handel-Mazzetti 4254 (WU); between Dali (Talifu) and Hodjing, above
Hsiang-schuiho, 26°10’N, 3,400 m., Handel-Mazzetti 7398 (pro parte) (W); Lopin-
chan Pass, above Lan-Kong, Delavay, 1888 (PC, 2 collections) ; Mt. Tsang-chan,
above Tali, 400 m., Delavay, 1885 (PC), 4,000 m., Delavay, 1885 (PC, 2 collections),
Delavay, 1887 (PC); Fang-Yang-Tschang, above Mo-so-yn, Delavay, 1887 (WU);
Koua-la-po, 3,000 m., Delavay, 1885 (H-Herb. Nylander, PC).

NEPAL: Above Sauwala Khola, 10,500 ft., Stainton et al. 4379b (pro parte)
(BM).

6. Cetrelia davidiana Culb. & Culb., sp. nov. PLATE 16

Thallus mediocris, 9-14 cm. latus; laciniae 0.7-2.2 cm. latae. Super-
ficies superior sordide citrella (albida apud plantas vivas?), levis,
pseudocyphellata, pseudocyphellis parvis, ca. 0.5 mm. crassis vel con-
fluentibus et ad 1 mm. Superficies inferior picea, marginibus castanea
vel pallida, levis vel subrugosa, fere punctata. Cortex superior 19-31,
crassus, medulla 115-2264 crassa; cortex inferior 12—-22y crassus.
Apothecia 0.5-2.0 cm. lata, perforata, primum cupuliformia deinde
expansa et radiatofissa, excipulo thallino rugoso et pseudocyphella-
tissimo; hymenium 71-115 crassum; subhymenium hyalinum vel
subluteum, 40-81, crassum, stratum crassum algarum tegens. Sporae
8-nae, ellipsoideae, 14-19 7-114. Pycnidia numerosa, marginalia, raro
in superficie superiore ad margines laciniarum; conidia 13-5 recta,
extremis nonnihil inflatis.

Thallus medium, 9-14 em. broad; lobes 0.7-2.2 em. broad. Upper
surface uniformly light tan (whitish when fresh?), smooth, pseudo-
cyphellate, the pores small, about 0.5 mm. in diameter, or becoming
confluent and reaching 1 mm. Lower surface jet-black, the margins
brown or colored like the upper surface, smooth or somewhat wrinkled,
especially at the ascendent margins of the lobes, if punctate, the puncta
restricted to the crests of the ridges. Upper cortex 19-31 thick;

508 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

medulla 115-226 thick; lower cortex 12—22y thick. Apothecia 0.5-2.0
cm. in diameter, perforate, cup-shaped at first then expanded and split
longitudinally, the thalloid exciple much wrinkled and abundantly
pseudocyphellate; hymenium 71-1154 thick; subhymenial layers
colorless or pale yellowish, 40-81 thick, immediately underlain by a
thick zone of algae. Spores 8, ellipsoid, 14-19 X 7-11, the spore wall
relatively thick, 1.0-1.7y. Pyenidia abundant, marginal or a few
laminal toward the margin of the lobes; conidia 13-5y, rod-shaped,
the ends of some somewhat enlarged.

Typr.—“In montis Yiilung-schan prope urbem Lidjiang [‘Liki-
ang’’], Yunnan,” China, Handel-Mazzetti 4254 (W, holotype).

MEDULLARY REACTIONS.—K —, C+ pink, PD—.

Constituents.—Olivetoric acid and atranorin.

DistripuTion (Map, Figure 20).—Mountains of Szechwan and Yun-
nan, China; on the bark of trees.

Cetrelia davidiana is a Chinese species without soredia, isidia, or
lobulae. It is similar to C. olivetorum chemically and in that it has
minute pseudocyphellae, but differs in that it is esorediate. It is also
similar to C. pseudolivetorum but differs in that it lacks the marginal
lobulae.

Cetrelia davidiana must be distinguished chemically from similar
esorediate and nonisidiate species most closely related to it. The posi-
tive reaction of the medulla with C will suffice to separate it from C.
collata and C. nuda and even from C. alaskana and C. delavayana, the
latter two of which also differ morphologically in possessing small and
inconspicuous pseudocyphellae rather than large and obvious ones.
The medullary reaction of C. davidiana with C is generally a pink or
rose due to the presence of olivetoric acid while that of C. sanguinea,
containing anziaic acid, is usually bright red. But definitive micro-
chemical tests are obviously preferable here to the thallus spot tests.

CHEMICAL ANALYSIS.—The extracts from fragments of seven samples
were tested. Atranorin was usually determined by crystals in the GE
and GAo-T solution and by chromatography. Olivetoric acid was
crystallized from the GE, GAW, and GAo-T solutions and was con-
firmed by chromatography to distinguish it definitely from anziaic
acid. Physodic acid did not occur with olivetoric acid in any of the
samples tested.

SPECIMENS EXAMINED:

CHINA: Szechwan: Above Fumadi, Wolo-ho River between Yenyiien and
Yungning, ea. 3,300 m., Handel-Mazzetti 3039 (WU).

YuNNAN: Between Dali (Talifu) and Hodjing, above Hsiang-schuiho, 26°15'N,
ca. 3,400 m., Handel-Mazzetti 7398 (pro parte) (W, 2 collections); Mt. Yilung-
schan near Lidjiang, Handel-Mazzetti 4254 (W, WU); Ti-ou-ze Pass, above Ta-pin-
tze, Delavay, 1888 (PC); Koua-la-po (Ho Kin), 3,000 m., Delavay, 1885 (PC).

PLATES

277-920 O - 68 - 5

PLATE 1

Crystals produced by substances in the microcrystal-test solutions: 1. Alectoronic
acid and a-collatolic acid cocrystallized in GE. The smaller crystals are atra-
norin. Prepared from the acetone extract after benzene extraction of Cetrelia
chicitae (X 125). 2. Alectoronie acid and a-collatolie acid cocrystallized in GE.
Prepared from the acetone extract after benzene extraction of Celrelia braun-
stana (110). 3. Alectoronic acid and a-collatolic acid cocrystallized in GAW.
Prepared from the acetone extract after benzene extraction of Cetrelia chicitae
(X 200). 4. Anziaic acid recrystallized in GE. Prepared from the acetone extract
after benzene extraction of Cetrelia isidiata (450). 5. Anziaic acid recrystallized
in GAW, Prepared from the acetone extract after benzene extraction of Cetrelia
isidiata (X 165). 6. Atranorin recrystallized in GE. Prepared from the acetone
extract after benzene extraction of Cetrelia sanguinea (140).

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 1

—

~~,
RX AN A
re WY!
WO)

+)

/

ed
~
r
=
+
oe
e

b-

PLATE 2

Crystals produced by substances in the microcrystal-test solutions: 7. The yellow
atranorin condensation product with GAo-T. Prepared from the benzene extract
of Platismatia erosa (X 120). 8. The yellow atranorin condensation product with
o-toluidine in GAo-T. Prepared from the benzene extract of Platismatia tucker-
manii (110). 9. The yellow atranorin condensation product with o-toluidine in
GAo-T showing two crystal types, the larger needles typical of the atranorin
derivative. The smaller, more densely clustered needles may be due to an un-
identified substance. Prepared from the benzene extract of Platismatia interrupta
(130). 10. Yellow crystals of the condensation product of atranorin and o-
toluidine in GAo-T distorted by the accompanying colorless caperatic acid.
Prepared from the acetone extract after benzene extraction of Platismatia tucker-
manit (1385). 11. Caperatic acid recrystallized in GAW. Prepared from the
acetone extract of benzene-extracted Platismatia erosa (115). 12. Caperatic
acid recrystallized in GAW showing typical crystals and abnormal curly plates
toward the center of the clusters. Prepared from the acetone extract of benzene-
extracted Platismatia interrupta (X 100).

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 2

~*~

9s j

Ati

PLATE 3

Crystals of caperatic acid in the microcrystal-test solutions: 13. The ‘‘cloud” form
of caperatic acid from recrystallization in GE. Prepared from ecaperatic acid
isolated from Parmelia cryptochlorophaea (240). 14. The “cloud”? form of
caperatic acid from recrystallization in GE. Prepared from the acetone extract
of benzene-extracted Platismatia glauca (X 140). 15. The “‘plate’’ form of eapera-
tic acid from reerystallization in GE. Prepared from a pure sample of caperatic
acid isolated from Parmelia cryplochlorophaea (X 120). 16. The ‘plate’? form of
sauperatic acid from reerystallization in GE. Prepared from the acetone extract
of benzene-extracted Platismatia glauca (180). 17. Mixed erystal forms of
caperatic acid from recrystallization in GE. Prepared from pure caperatic acid
isolated from Parmelia cryptochlorophaea (120). 18. Mixed crystal forms of
‘aperatic acid from recrystallization in GE. Needles and prisms of atranorin are
also present. Prepared from the acetone extract of benzene-extracted Platismatia
erosa (X 110).

CULBERSON & CULBERSON—PLATE 3

CONTR. NAT. HERB. VOL. 34, PT. 7

PLATE 4

Crystals produced by substances in the microcrystal-test solutions: 19. Pure
noreaperatic acid recrystallized in GE (X 165). 20. Pure noreaperatice acid re-
crystallized in GAW (X110). 21. Imbricaric acid (large needles) and atranorin
(small prisms) recrystallized in GE. Prepared from the benzene extract of
Cetrelia cetrarioides (X 110). 22. Imbricaric acid recrystallized in GAW. Prepared
from the benzene extract of Cetrelia cetrarioides (115). 23. The o-toluidine salt
of imbricaric acid (colorless needles) in GAo-T. The fuzzy crystals barely visible
are yellow and may be a cocrystallization with the atranorin condensation
product with o-toluidine, Prepared from the benzene extract of Cetrelia sinensis
(130). 24. Small yellow crystals beside the atranorin condensation product
with o-toluidine in GAo-T; commonly observed in samples containing imbri-
earic acid. Such crystals may be due to cocrystallization or to an unidentified
substance. Prepared from the benzene extract of Cetrelia cetrarioides containing
imbriearic acid (x 180).

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 4

NMP AO 19 : :

PLATE 5

Crystals produced by substances in the microcrystal-test solutions: 25. Micro-
phyllinie acid recrystallized in GE. Prepared from the benzene extraet of Cetrelia
japonica (X 120). 26. Microphyllinie acid reerystallized in GAW. Prepared from
the benzene extract of Celrelia japonica (135). 27. The sodium salt of micro-
phyllinie acid precipitated in 5% aqueous sodium hydroxide. Prepared from the
benzene extract of Cetrelia japonica (110). 28. Olivetorie acid reerystallized in
GE. Prepared from the acetone extraet of benzene-extraeted Celrelia olivetorum
(X 490). 29. Olivetorie acid reerystallized in GAW. Prepared from the acetone
extract of benzene-extracted Cetrelia davidiana (x 120). 30. The colorless o-
toluidine salt of olivetoric acid in GAo-T. Prepared from the acetone extract of
benzene-extracted Cetrelia olivetarum (X 110)

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 5

PLATE 6

Crystals produced by substances in the microcrystal-test solutions: 31. The color-
less o-toluidine salt of olivetoric acid in GAo-T. Prepared from the acetone ex-
tract of benzene-extracted Cetrelia pseudolivetorum (140). 32. Perlatolic acid
recrystallized in GE. Prepared from the benzene extract of Cetrelia cetrarioides
(X 115). 33. Perlatolic acid recrystallized from GAW. Prepared from the ben-
zene extract of Cetrelia delavayana (115). 34. The o-toluidine salt of perlatolic
acid (large colorless erystals) and atranorin (small yellow clusters) in GAo-T.
Prepared from the benzene extract of Cetrelia delavayana (170). 35. Yellow
crystals of an unidentified substance crystallized in GE. Prepared from an ex-
tract of Platismatia erosa (X 380). 36. Unidentified yellowish-tan crystals formed
in GAo-T from an extract of Platismatia interrupta (X 100).

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 6

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 7

, te
mate LU

Anatomical structure of the upper cortex in Cetraria, Parmelia, and Asahinca:
37. Parapleectenchymatous cortex of C. oakesiana Tuck. 38. Same, C. ericetorum
Opiz. 39. Prosoplectenchymatous cortex of P. (Amphigymnia) stuppea Tayl.
40. Same, A. chrysantha (Tuek.) Culb & Culb. All photographs 300,

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON-—PLATE 8

Anatomical structure of the upper cortex in Cetrelia and Platismatia: 41. Proso-
plectenchymatous cortex of C. chicitae (Culb.) Culb. & Culb. 42. Same, C.
nuda (Hue) Culb. & Culb. 43. Prosoplectenchymatous upper cortex of P. glauca
(L.) Culb. & Culb. 44. Same, P. tuckermanit (Oakes) Culb. & Culb. All photo-
graphs 300.

CULBERSON & CULBERSON—PLATE 9

CONTR. NAT. HERB. VOL. 34, PT. 7

MOF.

bre

Lia

ee

tt

YS

g

ae

Le

NO RNAS Oe

re

’

Af

vr

MOF? *

ee ee Se

CONTR. NAT. HERB. VOL. 34, PT. 7

CULBERSON & CULBERSON—-PLATE 10

Laminal and submarginal apothecia in Cetrelia: 47. Immature apothecia of C.
cetrartoides (Del. ex Duby) Culb. & Culb. (Culberson & Culberson 11563, from
Merionethshire, Wales) 8. 48. Apothecia of C. nuda (Hue) Culb. & Culb.
(Asahina, 1934, from Mt. Fuji, Japan) *4. 49. Apothecium and marginal

pyenidia of C. nuda (Tue) Culb, & Culb. (Togashi, 1955, from Yamoto Provinee,
Japan) 6,

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 11

50. Perforate apothecia of Cetrelia davidiana Culb. & Culb. (Handel-Mazzetti
4254, from Yunnan Provinee, China) 3.7. 51. Punetate lower cortex of
Platismatia erosa Culb. & Culb. Clwasthi 2292, from Nepal) <6.

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 12

92

The two types of pscudocyphellae in Cetrelia; 52. The small type of pseudoeyphel-
lac: Cetrelia olivetoruwm (NylL) Culb. & Culb. (Faron, 1882, from New England).
o3. The large type of pseudoeyphellae: Cetrelia nuda (Hue) Culb, & Culb. (Toga-

shi, 1955, from Musashi Provinee, Japan). Both photographs 3.5.

CONTR. NAT. HERB. VOL. 34, PT. ? CULBERSON & CULBERSON—PLATE 13

Soredia and isidia in Cetrelia; 54, Soredia in marginal soralia in Cetrelia olivetorum
(Nyl.) Culb. & Culb. (Lichenotheca Polonica 174, from Poland) 7. 55, Isidia
in Cetrelia braunsiana (Mill. Arg.) Culb. & Culb. (T'ogashi, 1956, from Eehigo

\

Provinee, Japan) & 7.

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 14

96

56. Cetrelia alaskana (Culb. & Culb.) Culb. & Culb. (Holotype, Thomson, Lichenes
Arctict 13, from Cape Sabine, Alaska) x 1.3. 57. Cetrelia chicitae (Culb.) Culb.
& Culb. (Kurokawa 59222, from Shinano Province, Japan) x 1.

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 15

98

58. Cetrelia braunsiana (Mill. Arg.) Culb. & Culb. (Togash?, 1956, from Echigo
Provinee, Japan) 2. 59. Cetrelia cetrarioides (Del. ex Duby) Culb. & Culb.
(Culberson & Culberson 11564, from Merionethshire, Wales) 0.8.

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON-—PLATE 16

60

60. Cetrelia davidiana Culb. & Culb. (Handel-Mazzetti 4254, from Yunnan Prov-
ince, China) 1.3. 61. Cetrelia delavayana Culb. & Culb. (Isotype, Delavay,
1888, from Yunnan Province, China) X 1.5.

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 17

62

62. Cetrelia isidiata (Asah.) Culb. & Culb. (Kurokawa 331, from Formosa) X 1.
63. Cetrelia japonica (Zahlbr.) Culb. & Culb. (Kurokawa 58374, from Shinano
Province, Japan), showing detail of the lobulae X 5.

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 18

64. Cetrelia nuda (Hue) Culb. & Culb. (Asahina, 1934, from Suruga Province,
Japan) X 1.3. 65. Cetrelia olivetorum (Nyl.) Culb. & Culb. (Lichenotheca Polonica
174, from Poland) x 1.2.

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 19

66. Cetrelia pseudolivetorum (Asah.) Culb. & Culb. (Kurokawa 137, from Formosa)
x1. 67. Cetrelia sanguinea (Schaer.) Culb. & Culb. (Du Rietz, 1927, from
Java) < 1.2.

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 20

68. Cetrelia sinensis Culb. & Culb. (Kurokawa 449, from Formosa) 1.5. 69.
Platismatia erosa Culb. & Culb. (Poelt L37, from Nepal) x 2.

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 21

Platismatia glauca (L.) Culb. & Culb. (two forms): 70. With fruticose branchlets
(Stickney 1005b, from Montana) 2.5. 71. With sorediate lobes (Imshaug
26501, from Quebec Provinee, Canada) X 1.1.

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 22

72. Platismatia herret (Imsh.) Culb. & Culb. (Lectotype, Hall, 1871, from Oregon)
x1. 73. Platismatia interrupta Culb. & Culb. (Culberson & Culberson 10669,
from Suruga Province, Japan) 1.5.

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 23

74. Platismatia lacunosa (Ach.) Culb. & Culb. (Hyerdam, 1947, from Alaska) * 1.3.
75. Platismatia norvegica CLynge) Culb. & Culb. GSzezawinsk? V.1./558, from
Vancouver Island, British Columbia, Canada) 1.

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 24

2
/

76. Platismatia regenerans Culb. & Culb. (Holotype, Hale 28207, from Sabah)
x2. 77. Platismatia stenophylla (Tuck.) Culb. & Culb. (Tolotype, Bolander
384, from California) x 0.9.

CONTR. NAT. HERB. VOL. 34, PT. 7 CULBERSON & CULBERSON—PLATE 25

18

78. Platismatia tuckermanti (Oakes) Culb. & Culb. (Moore 1372, from North Caro-
lina) 1.3. 79. Platismatia formosana (Zahlbr.) Culb. & Culb. (Suzukr, 1935,
from Formosa) X 2.

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 5()9

7. Cetrelia delavayana Culb. & Culb., sp. nov. PLATE 16

Thallus mediocris, 7-10 cm. latus;laciniae 1.3-1.7 em. latae. Super-
ficies superior sordide citrella apud exemplaria vetera herbarii (albida
apud plantas vivas?), levis, pseudocyphellata, pseudocyphellis puncta-
tis, paucis, parvissimis. Superficies inferior picea, marginibus castanea,
nonnihil punctata, leviter rhizinata. Cortex superior 16—22u crassus;
medulla 101-186u crassa; cortex inferior (12—)16—23u crassus. Apothe-
cia ad 0.8 cm. lata, perforata, excipulo thallino dense pseudocyphel-
lato; hymenium 62-1094 crassum; strata subhymenialia 43-78y
crassa, supra hyalina et subtus aurea. Sporae 8-nae, ellipsoideae, 16-24 «
9-12y. Pycnidia magna, marginalia, non stipitata; conidia ca. 1.5
6.0u, recta, extremis nonnihil inflatis.

Thallus medium, 7-10 cm. broad; lobes 1.3-1.7 em. broad. Upper
surface pale brownish in old herbarium specimens (whitish when
fresh?), smooth, pseudocyphellate, the pores few, very small, rarely
reaching 0.5 mm. Lower surface jet-black, the margins dark brown,
somewhat punctate in some specimens; rhizines few. Upper cortex
16-22, thick, medulla 101—186u thick; lower cortex (12—)16-23,u thick.
Apothecia to 0.8 cm. broad, perforate, the thalloid exciple densely
pseudocyphellate; hymenium 62-109 thick; subhymenial layers
43-78, thick, hyaline above and golden yellow below. Spores 8, ellip-
solid, 16-24 « 9—-12u. Pyenidia large, marginal, unstalked; conidia about
1.5 6.0u, rod-shaped, the ends often somewhat inflated.

Type.—‘‘Au col de Lopin-chan,’”’? Yunnan, China, Velavay, 1888
(PC, holotype).

MEDULLARY REACTIONS.—K—, C—, KC— or KC-+ pinkish, P—.

ConsTITUENTS.—Imbricaric acid and atranorin.

DistRiBuTION (Map, Figure 21).—In the mountains of Szechwan and
Yunnan, China; on the bark of trees and shrubs.

Cetrelia delavayana can be distinguished from most other esorediate
and nonisidiate species of the genus both morphologically, by the
presence of small pseudocyphellae rather than large aggregate ones, and
chemically by the presence of perlatolic acid. Cetrelia collata produces
imbricaric acid and inhabits much the same range but has very large
pores exceeding 1 mm. Cetrelia alaskana has small pores and imbricaric
acid, its habit is completely different, and its range is the western coast
of Alaska.

CHEMICAL ANALYsIs.—Eight samples of this species were tested. All
showed perlatolic acid and atranorin. Four residues were hydrolyzed
and showed no trace of divaric acid, indicating the absence of imbri-
caric acid, but at least two samples gave a KC+ reaction which could
not be associated with identifiable crystals or chromatographic spots.
It is possible that this unidentified KC+ accessory compound is the

277-920 O - 68 - 6

510 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

same as that which gives a KC+ reaction in other Cetfelia species
which contain acid.

SPECIMENS EXAMINED:

CHINA: SzecHwan: Mt. Lose-chan, south of Ningyiien, 3,950 m., Handel-
Mazzetti 1441 (WU).

YUNNAN: Yangtze watershed, Likiang, Likiang Snow Range, Rock 11775 (W);
Lenago Pass between Mekong and Yangtse, 27°43'N, Gebauer, 1914 (TNS, W,
WU); between Dali (Talifu) and Hodjing, above Hsiang-schuiho, 26°15’N, 3,400
m., Handel-Mazzetti 7398 (W); Lopin-chan Pass, Delavay, 1888 (PC, 2 collections).

8. Cetrelia isidiata (Asah.) Culb. & Culb., comb. nov. PuaTE 17

Cetraria sanguinea Schaer. f. isidiata Asah. in Sato, Parmeliales (I), in Nakai

& Honda, Nova Fl. Jap. 73. 1939. Type: Japan, Honshu, Titibu [-=Chi-
chibu], Mt. Kobushi, Asahina, 1933 (TNS, holotype).

Thallus small to large, 5-19 em. broad; lobes 0.5-1.5 cm.broad.
Upper surface grayish or uniformly light brown in old herbarium speci-
mens, pseudocyphellate, the pores tiny and infrequent, isidiate, the
isidia mostly marginal, globose to somewhat coralloid or slightly
branched, or poorly developed and very infrequent altogether. Lower
surface jet-black, the margins brown, punctate or not; rhizines con-
spicuous but unevenly distributed. Upper cortex 16-314 thick; medulla
84-217 thick; lower cortex 13-314 thick. Apothecia and pycnidia
unknown.

MEDULLARY REACTIONS.—K—, C+ red, PD—

ConstITUENTs.—Anziaic acid and atranorin.

Distripution (Map, Figure 8).—Honshu, Japan, and Formosa; on
trees.

The only other C+ isidiate species that Cetrelia isidiata might be
mistaken for is C. pseudolivetorum which contains olivetoric acid and
reacts to produce a pink color with this reagent. A chemical test is
desirable for the identification of these species since the thallus spot
tests may be somewhat variable.

The original classification of the present taxon as a variety of the
Javanese species sangvinea is not tenable even though both produce
anziaic acid in the medulla. The Javanese plants lack isidia, have very
large pseudocyphellae that become confluent, and usually bear apothe-
cla, while the Japanese and Formosan ones have isidia and minute
pseudocyphellae and are unknown in fruit. Cetrelia isidiata seems to be
a very well defined species, more closely related to C. chicitae, for
example, than to C. sanguinea.

The single Formosan collection examined (Plate 17) is rich and well
developed, but it shows a very meager production of isidia as do the
Japanese specimens. Since only a few collections of this species are
known, it is impossible to give a definitive assessment of the impor-
tance of isidia as a diagnostic trait; in fact they may be of little conse-

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 511

quence. It is, therefore, with some hesitation that we take up isidiata
as the specific epithet, a name given by Asahina to the taxon when he
considered it a forma of the species treated in this monograph as
Cetrelia sanguinea. Isidia are much less well developed in the present
species than in C. brawnsiana, for example.

CHEMICAL ANALYsIS.—The four known specimens of this species
were tested, and all contained atranorin ( proved in the GAo-T solution)
and anziaic acid (identified by paper chromatography).

ADDITIONAL SPECIMENS EXAMINED:

JAPAN: Honsuu: Chichibu, Mt. Kobushi, Asahina, 19383 (TNS, original
collection; 3 specimens).

FORMOSA: Mt. Shin-Kao-San, Mt. Shi-San, 3,300-3,600 m., Kurokawa 311
(TNS).

9. Cetrelia japonica (Zahlbr.) Culb. & Culb., comb. nov. PLATE 17
Cetraria japonica Zahlbr., Annal. Mycol. 14:60. 1916. Type: Japan, Prov.
Kotsuke, Mt. Akagi, Yasuda (W, lectotype).
Platysma collatum f. microphyllinum Hue, Nouv. Arch. Mus., IV, 1:209. 1899.
Type: Japan, Fujiyama, Faurie 519 (PC, lectot ype).
Cetraria collata f. micropiyllina (Huc) Zahibr. Catal. Lich. Univ. 6:285. 1630.

Thallus medium to very large, 10-28 em. broad; lobes 0.5-1.5 em.
broad, the margins densely fringed with multibranched lobulae, narrow
and isidioid or more broadly expanded and lobelike, in some specimens
becoming so abundantly developed as to make the lichen appear
cristate. Upper surface grayish-white or ashy-white (or light brownish
in old herbarium specimens), smooth, pseudocyphellae conspicuous,
punctiform to elongate but rarely reaching 1 mm. in size. Lower sur-
face jet-black, a marginal zone near the tips of the lobes chestnut
brown or even whitish, not punctate; rhizines black. Upper cortex
18-23y thick, interspersed with abundant crystals; medulla 83-1714
thick; lower cortex 16-23 thick, without crystals, darkening. Apothe-
cla very rare, submarginal, often perforate, 0.4-1.3 cm. broad; hyme-
nium 65-70y thick; subhymenial layers 47-57 thick, underlain by a
thick continuous zone of algae. Spores 8, ellipsoid, 14-21 8-I1p.
Pyenidia very rare, large, black, emergent, about 0.1 mm. broad,
restricted to the marginal lobulae, the ostiole prominent; conidia
1.5 X5u, rod-shaped, the ends somewhat enlarged.

MEDULLARY REACTIONS.—K —, C—, KC— or KC-+4 pink, PD—.

ConstTITUENTS.—Microphyllinic acid and atranorin.

Distripution (Map, Figure 16).—The mountains of Hokkaido and
Honshu in Japan, South Korea, Formosa, northern Sabah (Borneo),
and Java.

Cetrelia japonica is chemically distinct from all the other species of
the genus in the production of microphyllinic acid. Morphologically,
however, it could be mistaken for C. sinensis (with imbricaric acid), C.

512 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

sanguinea (with anziaic acid), or C. pseudolivetorum (with olivetoric
acid). While its distinction from C. sinensis Is probably purely chem1-
cal, the morphology of the whole population of C. japonica is appar-
ently somewhat different from that of the populations of C. sanguinea
and ©. pseudolivetorum, although it must be admitted that many
individual specimens of all these species cannot be identified without
chemical tests. As a rule the margins of the lobes of C. japonica are
moderately to densely fringed with tiny lobulae or dorsiventral isidia.
Cetrelia pseudolivetorum and C. sanguinea also have similar lobulae, but
as a rule the lobulae are less well developed, and many thalli, especial-
ly in C. sanguinea, may lack them altogether.

The population assigned herein to Cetrelia japonica may represent
two taxa. The only known Javanese specimen has only true isidia
rather than lobulae. It may be a rare Javanese endemic, but it 1S as-
signed to C. japonica provisionally because such isidia can be found
(with lobulae) on some specimens of C. japonica from Japan, because
the Javanese plant produces microphyllinic acid (a rare chemical trait
in the genus), and because typical specimens of C. japonica are known
from nearby Sabah.

CHEMICAL ANALYSIS.—This species contains the rare lichen sub-
stance microphyllinic acid. Twenty-two samples were tested, and all
but two showed atranorin in either the GE or the GAo-T solutions.
But microphyllinic acid seems to interfere with the erystal tests for
atranorin so that often it is necessary to allow the slides to stand over-
night before crystals of atranorin are observable in the GAo-T solution.
The two samples which seem to give a negative test for atranorin
undoubtedly do contain this substance since the GAo-T slides became
bright yellow, and atranorin should be considered a constant compo-
nent of this species. Atranorin was not determinable by chromatography
in the presence of microphyllinic acid by the methods used in this study
because a decomposition product of microphyllinic acid has the same
R, value as atranorin. Microphyllinic acid was proven by its crystals
in the GE solution and in 10 percent NaOH solution as well as by
chromatography.

REPRESENTATIVE SPECIMENS EXAMINED:

JAPAN: Hoxxarpo: Prov. Ishikari: Mt. Ashibetsu, Asahina, 1935 (TNS, 2
collections).

Honsnu: Togakushi, Faurte 777, 795 (PC, 2 collections). Towada, Faurie 14300,
14304 (PC, 2 collections). Prov. Echigo: Iwafune-gun, Takanosu, Togashi, 1959
(TNS). Prov. Hitachi: Tsukuba, Asahina, 1924 (TNS). Prov. Inaba: Yazu-gun,
Mt. Misumi, [koma 2512 (TNS). Prov. Iwashiro: Makukawa Hot Spring, Auro-
kawa 58127 (TNS, US). Prov. Kai: Mt. Fuji, Yoshida-guehi, Hisauchi, 1928
(TNS). Prov. Kii: Mt. Koya, Asahina & Togashi, 1957 (TNS); the same, Nwma-
jirt, 1926 (TNS, 3 collections). Prov. Kotsuke: Mt. Akagi, Yasuda (W). Prov.
Mikawa: Mt. Horaiji, Park Iwagoya, Asahina et al., 1960 (TNS). Prov. Musashi:

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 513

Mine-no-Yakushi, Asahina, 1939 (TNS); Chichibu, Mt. Ryogami, Kurokawa
550571 (TNS). Prov. Rikuchu: Mt. Hayachine, Monma-guchi, Kurokawa 59275
(TNS). Prov. Sagami: Hakone, Mt. Futago, Asahina, 1952 (TNS); Motohakone,
Asahina, 1924 (TNS). Prov. Settsu: Mt. Myoken, Ui, 1935 (TNS). Prov. Shinano:
Mt. Yatsu-ga-take, Kurokawa 51196, 51197, 51198 (TNS); Pass Kitazawa, Hattyo-
zaka, Asahina, 1937 (TNS); Mt. Nyugasa, Yoshioka, 1937 (TNS). Prov. Suruga:
Mt. Fuji, Omiyama-guchi, 3-2-gome, T'ogashi, 1961 (TNS); Fujiyama, Faurie 519
(PC). Prov. Tanba: Minamikuwata-gun, Sogabe-mura, Togashi, 1956 (TNS).
Prov. Wakasa: Mt. Aoba, Togashi, 1956 (TNS, 2 collections). Prov. Yamato:
Omine-san, Mt. Sanjo-ga-take, Togash7, 1952 (TNS).

SHIKOKU: Prov. Awa: Mt. Ken-zan, Fujikawa, 1934 (TNS). Prov. Tosa: Be-
tween Mt. Tsutsujo and Mt. Tebako, Iwatsuki 1804 (TNS).

Kyusuu: Prov. Bungo: Mt. Urahiko, Komura, 1951 (TNS). Prov. Buzen: Mt.
Hikosan, Komura 984 (TNS). Prov. Higo: Mt. Ichibusa, Fujikawa, 1933 (TNS).
Prov. Hizen: Mt. Unzen, Nodake, Asahina & Togashi, 1958 (TNS). Yakushima,
Kosugidani, Omura, 1950 (TNS).

KOREA: Prov. Kei-nan: Mt. Chi-san, Sato 331a (MSC).

FORMOSA: Prov. Ilan: Piyanan Pass, 1,100-1,300 m., Kurokawa 741 (TNS).

SABAH (BORNEO): Kinabalu National Park, from Mesilau River to Kunda-
son, along Mesilau Trail, Hale 29269 (US).

JAVA: Res. Pasoeroean, Goenoeng Ardjoena, Tretes-Lalidjiwa-track, Du
Rretz 95:16 (UPSV).

10. Cetrelia nuda (Hue) Culb. & Culb., comb. nov. PLATE 18
Platysma collatum f. nudum Hue, Nouv. Arch. Mus., IV, 1:208. 1899. Type:
Japan, Fujiyama, Faurie 569 (PC, lectotype).
Cetraria collata f. nuda (Hue) Zahlbr., Catal. Lich. Univ. 6:285. 1930.
Parmelia yunnana Hue f. subnuda Zahlbr., Hedwigia 74:210. 1934. Type:
China, Lidjiang Snow Range, 11,300-12,000 ft., Rock, 1931 (W, holotype).
Thallus large, 16-24 cm. broad; lobes 0.6-1.8 cm. broad. Upper sur-
face grayish or ashy-white, smooth, pseudocyphellate, the pores large,
conspicuous, irregular, becoming confluent, the largest aggregates ex-
ceeding 1 mm. Lower surface black, the margins of some lobes similar
in color to the upper surface, wrinkled in part and, if very rugose, with
puncta on the crests of the folds. Upper cortex 16-31, thick; medulla
130-186, thick; lower cortex 16-22y thick. Apothecia very common,
submarginal to superficial, perforate, irregular at maturity, 1.3-1.5
em. broad. Spores 8, ovoid, 14-16 X6-10u. Pycnidia always present,
large, often pruinose, marginal or at times superficial; conidia 1<5~7y,
rod-shaped, the ends slightly inflated.
MEDULLARY REACTIONS.—K —, C—, KC+ pink, PD—.
ConstituENts.—Alectoronic and a-collatolic acids and atranorin.
Distr1BuTION (Map, Figure 17).—Widely distributed in the islands
Honshu, Shikoku, and Kyushu in Japan and the high mountains of
Formosa and Yunnan; locally common on tree trunks and over
boulders.
In Japan this species is often conspicuous because of its broad lobes
speckled with aggregate pseudocyphellae and its large thallus. The
negative reaction of the medulla with C will distinguish it from Cetrelia

514 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

davidiana and C. sanguinea; from C. collata, which contains imbricaric
acid, the only certain identification is based upon the microchemical
demonstration of the alectoronic and/or a-collatolic acids of C. nuda.
Cetrelia alaskana and C. delavayana, which may superficially resemble
C. nuda, differ chemically in producing imbricaric acid and morpho-
logically in having tiny pseudocyphellae rather than large, aggregate
ones. Some specimens of C. braunsiana with poorly developed isidia
may resemble C. nuda, but they have very small pseudocyphellae.

This taxon has been recognized in Japan as Cetraria collata f. nuda
(Hue) Zahlbr., and its taxonomy has been confused with that of Cetrelia
chicitae (the ‘‘Cetraria collata f. isidiata”’ of the Japanese literature).
The reasons for recognizing these three variations as species are dis-
cussed in the treatment of Cetrelia braunsiana (see p. 495).

Cetrelia nuda is the only species in the genus for which any informa-
tion on the physiological ecology is available. Hosokawa and his co-
workers (1964) reported that in this lichen, called Cetraria collata f.
nuda in their paper, apparent photosynthesis increases both with in-
creasing light and increasing relative humidity, the maximum photo-
synthesis as measured by carbon dioxide uptake being at 50,000 Lux
and 90 percent relative humidity. They believe that these results cor-
relate well with their observation that in closed forests Cetrelia nuda
occurs primarily in the crowns as opposed to the trunks or the bases
of the trees. Cryptogams habitually more abundant at lower levels
in the same study sites, they showed, have maximum rates of photo-
synthesis at much lower light intensities—for example, at about 10,000
Lux for the moss Thuidium cymbifolium Doz. & Molk., an inhabitant of
the tree base. Furthermore, at high light intensities, C. nuda carries on
appreciable photosynthesis even when the relative humidity is reduced
to 70 to 80 percent. These data would seem to indicate that C. nuda
has very high light requirements and yet is also at least somewhat
adapted to effective photosynthesis at the lower relative humidities
that are encountered in places of high illumination.

CHEMICAL ANALYSIS.—AII 22 samples of this species tested contained
atranorin and alectoronic acid. Twenty of the samples tested also con-
tained a-collatolic acid. Crystals of atranorin were observed in the
GAo-T solution and sometimes in the GE solution. The substance was
also demonstrated chromatographically. Alectoronic acid and a-col-
latolic acid in mixture were detected by chromatography after observ-
ing the crystals in the GE and GAW solutions. Many samples were
also checked with 10 percent NaOH to verify the absence of micro-
phyllinic acid.

NoMENCLATURE.—Hue considered this to be a forma of Platysma
collatum, which he believed to be a sorediate species.

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 515

REPRESENTATIVE SPECIMENS EXAMINED:

JAPAN: Honsuu: Onikobe, Faurie 283 (PC). Togakushi, Faurie 777 (pro parte)
(PC, 2 specimens). Prov. Aki: Yamagata-gun, Furuyashiki-Mt. Osorerankan,
Kotake, 1956 (TNS). Prov. Izu: Mt. Amagi, Asahina, 1927 (TNS). Prov. Kai: Pass
Syosen, Kurokawa 521285 (TNS). Prov. Kii: Mt. Koya, Numajiri 699 (TNS).
Prov. Kozuke: Mt. Akagi, Asahina, 1935 (TNS). Prov. Musashi: Chichibu, Mt.
Ryogami, Kurokawa 550572 (LD, TNS); Nippara, Pass Sengen, Asahina 71a
(TNS); Mt. Shiroiwa, Kurokawa 50380B, 50431B, 510274 (TNS). Prov. Mutsu:
Kise, Murai, 1933 (TNS). Prov. Sagami: Hakone, Owaku-dani, Watanabe, 1955
(TNS). Prov. Shimotsuke: Nikko, Faurie 577 (PC); Nikko, Yumoto, Asahina,
1952 (TNS). Prov. Shinano: Shibunoyu, Mt. Yatugatake, Kurokawa 58197
(MSC). Prov. Suruga: Fujiyama, Faurie 508 (pro parte), 569 (PC); Mt. Fuji,
Omiya-guchi, 2-gome, Asahina, 1925, 1926, 1932, 1934 (TNS, 7 collections); Mt.
Fuji, Umagayeshi, Almquzst, 1879 (S). Prov. Yamato: Mt. Omine, Yosigayado,
1,600 m., Tagawa 269 (TNS).

Suikoxv: Prov. Awa: Mt. Ken-zan, Togashi, 1956 (TNS). Prov. Iyo: Kitauwa-
gun, Mt. Nametoko, Ogata, 1928, 1930 (TNS, 2 collections). Prov. Tosa: Mt.
Tsutsujo-Mt. Tebako, Iwatsuki 1787, 1806, 1811 (TNS).

Kyusuvu: Prov. Bungo: Mt. Kuro-dake, Omura, 1950 (TNS). Prov. Buzen:
Shimoge-gun, Mt. Ura-hiko, Omura, 1951 (TNS). Prov. Higo: Mt. Ichibusa,
Fujikawa, 1933 (TNS).

FORMOSA: Mt. Ali, 2,200 m., Kurokawa 13, 37 (TNS); Mt.Tsu-Tson-San,
Mt. Ali, 2,300-2,900 m., Kurokawa 122 (TNS).

CHINA: Yunnan: Kou-toui, above Mo-so-yn, 3,000 m., Delavay, 1887 (PC);
Schweli-Salween-Scheidegebirge, 25°45’N, 2,000-2,800 m., Gebauer, 1914 (WU);
Mekong-Salween-Scheidegebirge, 26°10’N, Gebauer, 1914 (W; WU, 2 collections).

11. Cetrelia olivetorum (Nyl.) Culb. & Culb., comb. nov. PLaTE 18

Parmelia olivetorum Nyl., Not. Siallsk Fauna Fl. Fenn. Forhandl., n. ser.
5:180. 1866. Type: Switzerland, collector unknown, s.d. (H-Herb. Acharius,
lectotype).

P. perlata f. (?) rubescens Th. Fr., Lich. Scand. 112. 1871. Type: Switzerland,
collector unknown, s.d. (H-Herb. Acharius, lectotype).

P. rubescens (Th. Fr.) Vain., Bot. Mag. [Tokyo] 35:47. 1921.

P. cetrarioides var. rubescens (Th. Fr.) Du Rietz, Nyt Mag. Naturv. 62:75.
1924,

Pseudoparmelia aradensis Gyeln., Acta Fauna Fl. Univ., ser. 2 Bot. [Bucaresti]
1(5-6) :6. 1933. Type: Rumania, Arad, Kiimmerle & J dvorka (BP, holotype).

Parmelia olivaria f. subvenosa Gyeln., Repert. Spec. Nov. Fedde 36:302. 1934.
Type: Hungary, prope Budapest, Timké (BP, holotype).

[Not P. perlata 8 olivaria Ach., Meth. Lich. 217. 1803.]

[Not P. perlata 8 olivetorum Ach., Lichenogr. Univ. 458. 1810.]

Thallus small to medium, 6-14 em. broad; lobes 0.5-1.8 cm. broad.
Upper surface light tan or brownish, smooth, sometimes slightly con-
vex, pseudocyphellate, the pores abundant and very small, rarely ex-
ceeding 0.5 mm.; the margins of most lobes with dense, fine soredia.
Lower surface black, the margins chestnut brown or colored like the
upper surface, rarely punctate; rhizines few, black. Upper cortex 19-
31y thick; medulla 96-239, thick; lower cortex 16—25y thick. Apothecia
rare (unknown in Asian specimens), 0.1-0.7 cm. broad, not perforate;

516 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

hymenium 62-109y thick; subhymenial layers 62-84, thick, the proper
exciple underlain by a thick layer of algae. Spores ellipsoid, 12-15
7-10u. Pyenidia unknown.

MEDULLARY REACTIONS.—K—, C+ pink or red, PD—.

ConsTITUENTS.—Olivetoric acid and atranorin.

DistrriBuTION (Map, Figure 7).—Appalachian Mountains and north-
eastern United States and adjacent Canada; southern Scandinavia to
the high mountains of southern Europe; high mountains of south-
western China, Formosa, and Japan; over boulders and on tree trunks.

Although Cetrelia olivetorum is very closely related to C. cetrarioides
and C. chicitae, it can almost invariably be distinguished from them by
the experienced eye. The sum total of its small, difficult-to-describe yet
distinctive traits—the slightly convex lobes, the abundant but ex-
tremely minute pseudocyphellae, the extremely fine but abundant mar-
ginal soredia, and the total lack of laminal soredia—makes it possible
to name most specimens at sight. In the southern Appalachians, where
this species can be found growing on the same boulders with C. chicitae,
it almost invariably shows tinier pseudocyphellae and a slightly dif-
ferent habit than the latter species. Nevertheless, some individuals of
C. chicitae and C. olivetorum are so similar that chemical tests are re-
quired for positive identifications. Cetrelia olivetorum can be distin-
euished by its distinctive, pinkish-rose medullary reaction with C due
to the presence of olivetoric acid. Cetrelia cetrarioides produces perlato-
lic acid or imbricaric acid and C. chicitae produces alectoronic and
a-collatolic acids, explaining why neither of the species gives a positive
reaction with C.

The lack of information about the precise range of this species is in
part due to the fact that it is relatively rare and in part because it has
been mistaken for the commoner Cetrelia cetrarioides. The species
seems to be restricted to regions of relatively cool summers and ample
rainfall. In all but the northern part of its range, these conditions are
met with only at higher elevations and all the southern stations of the
species are in mountains. The species usually grows with bryophytes
over shaded boulders or on the trunks of trees in moist forests.

The American range is the most accurate part of the map of the total
world distribution (Figure 7). Ahti (1964) made a detailed study of the
region between the north shore of Lake Superior and Hudson Bay. He
records the species from a single locality a little over 100 miles north of
the northernmost locality mapped and notes that the lichen there is
“rare.”’ Lynge (1921) records a few localities in southern Norway and
Du Rietz (1924) and Hakulinen (1963) attribute it to Finland but
without citing precise localities. Records for the southern part of the
range in Europe are doubtless nearly accurate, the localities corre-
sponding to the higher mountains there.

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 517

Generalizations about the distribution in Asia are impossible, yet it
seems significant that very few specimens from Japan are present in
herbaria in spite of intensive lichenological collecting in that country.
On the other hand, a fairly large number of well developed specimens
are known from the high mountains of Yunnan Province, and the
species may achieve its maximum abundance in Asia in southwestern
China.

The frequency with which Cetrelva olivetorum has been found mixed
with C. cetrarioides in packets in herbaria indicates that these species
often grow together in nature and doubtless have a very similar physio-
logical ecology.

CHEMICAL ANALYSIS.— All 19 samples tested contained atranorin and
olivetoric acid. Olivetoric acid was readily identified in most samples
by its crystals in the GE, GAW, and GAo-T solutions. To avoid con-
fusion with anziaic acid, all extracts were also checked chromatograph-
ically. Physodic acid, known to occur with olivetoric acid in Cetraria
ciliaris (C. F. Culberson, 1964), was not found in any of the samples of
Cetrelva olivetorum tested.

NOMENCLATURE.— The specific epithet olivetorum is a variation of
the earlier Acharian epithet olwvaria which alluded to the olive trees in
southern Europe from which the (different) plants originally called
olwarva reputedly came. The present species does not grow in the olive
region of Europe and the name is wholly inappropriate. It might be
noted, however, that olivetoric acid, the characteristic constituent of
the medulla of this species, was extracted for the first time from Cetrelva
olivetorum and was named for it.

The name oliwetorum has been a source of considerable confusion
inasmuch as Acharius and Nylander both used it at different ranks for
different taxa. Acharius (1803) named a broad-lobed Parmelia from
olive trees P. perlata g olivaria Ach. and later, for some reason, referred
to the same taxon as P. perlata B olivetorum Ach., placing the earlier
epithet olivaria as a synonym (Acharius, 1810). In 1866 Nylander used
the epithet olivetorum at the species rank and indicated Acharius as the
originator of the name although he used it for a taxon different from
the one for which Acharius had coined it.

The appropriate sheet in the Acharian herbarium contains three
specimens, the illegitimate varietal name olivetorum of Acharius being
written closest to one which appears to be Parmelia reticulata Tayl., a
species which could have been collected on olive trees in Europe al-
though its geographic origin is not given. The medulla of this specimen
is C—. Both of the other specimens belong to the species under con-
sideration here; their medullary reaction is C+ red, and the larger
specimen is labeled “Helvetia.” Since Cetrelia olivetorum does not grow
in the olive zone of Europe, neither of the two Swiss specimens can be

518 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

associated with the Acharian epithets olivaria or olietorum. These two
specimens, with their distinctive medullary reactions with C, are surely
the ones Nylander referred to in his diagnosis of Parmelia olivetorum:
“Ach, ex hb. ipsius, cui speciei medulla thalli ope hypochloritis calici
erythrinice tingitur.” The binomial Parmelia olivetorum must be attrib-
uted to Nylander alone (Art. 72) and typified upon one of the C+
specimens. The larger specimen labeled “Helvetia” is the lectotype.
This specimen is also designated as the lectotype of P. perlata f. (?)
rubescens, a taxon described apparently as a forma by Th. Fries who
seems to have understood the nomenclatural history of the olivetorum
names in spite of the solution that he offered by proposing another
name altogether.

REPRESENTATIVE SPECIMENS EXAMINED:

U.S.A.: Marne: Knox Co.: Rockland, Merrill, Lich. Exs. 99 (DUKE). York
Co.: Kennebunkport, Pier, 1917 (UC, DUKE).

New Hampsuire: Cheshire Co.: Alstead, Seymour, 1921 (UC, DUKE). Graf-
ton Co.: North Woodstock, collector unknown (DUKE).

VerMONT: Caledonia Co.: McIndoe’s Falls, Bliss, 1918 (DUKE).

New York: Essex Co.: Wilmington, Brodo 499 (DUKE).

PENNSYLVANIA: Monroe Co.: Mt. Pocono, Kaiser 1910 (DUKE).

West Viroeinia: Pocahontas Co.: Gaudiner Knob, Hale, Lich. Amer. Exs. 56
(UC).

Virainta: Giles Co.: Mountain Lake, Culberson 11180, 11238 (DUKE). Madi-
son Co.: Shenandoah National Park, Hale 19213 (pro parte) (US). Page Co.: Little
Stony Man Cliffs, Shenandoah National Park, Hale 19219 (US); Skyline Drive,
Crescent Rocks, Hale 18167 (US).

Norru Carouina: Avery Co.: Near the Avery-Caldwell County line, Culberson
11770 (DUKE). Rockbridge Co.: Mile 35, Blue Ridge Parkway, Hale 17813
(US).

Ouro: Locality unknown, Sullivant (CB).

Micuican: Isle Royale: Rock Harbor, Harper 160 (UC, DUKE).

Wisconsin: Langlade Co.: Langlade, Culberson 2007 (DUKE). Price Co.: Park
Falls, Culberson 1814 (DUKE).

JAPAN: Prov. Echigo: Sasa-ga-mine, Togashi, 1950 (DUKE).

FORMOSA: Mt. Ali, Mt. Tsu-Tson-san, 2,300-2,900 m., Kurokawa 133 (TNS).
Prov. Nautoh, Lisan, Kurokawa 880 (TNS).

CHINA: Szecuwan: Mt. Lose-schan, south of Nengyiien, 3,950 m., Handel-
Mazzetti 1441 (W).

YuNNAN: Lenago Pass between Mekong and Yangtze, 27°43" N, Gebauer, 1914
(W, WU); Fang-Yang-Tchang, above Mo-To, Delavay, 1887 (PC); Long, above
Ta-pin-tze, Delavay, 1887 (PC); Koua-la-po, Delavay 1595 (pro parte) (PC); Hee-
chan-men, Delavay, 1887 (PC); Yen-tze-hay, Delavay, 1887 (PC, 2 collections).

NEPAL: Above Sauwala Khola, 10,500 ft., Statnton et al. 43796 (pro parte)
(BM).

RUMANIA: Transytvania: Hunyad, Lojka (PC); Arad, Kiimmerle & Jdvorka,
s.d. (BP).

POLAND: Bieszezady Zachnodnie Mountains, Puszeza Bukowa, Mt. Paportna,
810 m., Glanc, as Tobol., Lichenoth. Polonica 174 (DUKE).

HUNGARY: Near Budapest, m. Nagyhdrshegy, Timké (BP).

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 519

SWITZERLAND: Léman, collector unknown (PC).

GERMANY: Bavaria: Miinchen, Arnold, Lich. Monacensis Ezs. 324a,b,
Arnold, Lich. Monacensis Exs. 6 (PC).

FRANCE: Cores-pu-Norp: Forét de Lorges, des Abbayes, 1929 (Herb. des
Abbayes).

FInIsTERE: Forét de Cranou, des Abbayes, 1933 (Herb. des Abbayes).

HauTEs-Pyr&én&Es: Cauterets, Lamy de la Chapelle (PC).

12. Cetrelia pseudolivetorum (Asah.) Culb. & Culb., comb. nov. PLATE 19
Parmelia (Amphigymnia) pseudolivetorum Asah., Journ. Jap. Bot. 27:16.
1952. Type: Japan, Mt. Fuji, Asahina (TNS, holotype).

Thallus small to large, 0.7-18.0 em. broad; lobes 0.5-1.5 cm. broad.
Upper surface grayish or grayish-white or uniformly light brownish or
tan in old herbarium specimens, smooth or becoming cracked, pseudo-
cyphellate, the pores small, to 0.5 mm. broad, punctiform or slightly
elongate, the margin and occasionally the surface with granular, simple
or coralloid isidia, often grading into dorsiventral, dissected lobulae.
Lower surface black, the margins brown or colored like the upper sur-
face, not punctate; rhizines few, black. Upper cortex 16-30u thick;
medulla 150-200u thick; lower cortex 11-17p thick. Apothecia very
rare, 0.4—-1.7 em. broad, mostly perforate: hymenium 53-78u, hyaline;
subhymenial layers 40-53 thick. Spores 8, ellipscid, (11—)16-22
9-12u. Pyenidia unknown.

MEDULLARY REACTIONS.— K—, C+ pink, P—, UV—.

ConstITUENTS.— Olivetoric acid and atranorin.

DistriBuTIon (Map, Figure 9).—The mountains of central Japan,
Formosa, and Yunnan, China, and the Himalayas; usually on rock or
over boulders with bryophytes.

Cetrelia pseudolivetorum has a pink medullary reaction with C due to
the presence of olivetoric acid thereby distinguishing it from the two
morphologically similar species for which it would most probably be
mistaken, C. japonica (with microphyllinic acid) and C. sinensis (with
imbricaric acid). In reality, though, the bulk of the specimens of C.
pseudolwetorum differ morphologically from these species (see the dis-
cussion of C. japonica, pp. 511-512).

Some forms of Cetrelia pseudolivetorum with isidia or isidioid lobulae
instead of the expanded dorsiventral lobulae more typical of the species
might be mistaken for the common Japanese species C. braunsiana.
The latter species has very delicate coralloid isidia and is C— but
KC+ because of its production of alectoronic acid. The two species
can usually be distinguished without the need of chemical thallus spot
tests, however.

Cetrelia pseudolivetorum might also be mistaken for some rare forms
of C. sanguinea with well-developed lobulae. If it has small marginal
isidia, it might be mistaken for C. isidiata. Both these species give a red

520 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

(rather than a pinkish) medullary reaction with C and both produce
anziaic acid rather than the olivetoric acid.

Although chemically identical, Cetrelia olivetorum and C. pseudo-
livetorum are morphologically distinct. The margins of C. olvvetorum
have extremely fine, powdery, white soredia while those of C. pseudo-
livetorum have isidioid lobulae or similar outgrowths. There are no
integradations between the two species although badly damaged
specimens of C. pseudolivetorum, in which the delicate marginal ap-
pendages have been broken off exposing the white medulla, might be
mistaken for C. olivetorum.

Cetrelia pseudolivetorum shows considerable variability, but the
variation is so continuous that there is no reason to suspect that the
population is not sufficiently uniform to be considered a single species.
Forms with coralloid marginal isidia recalling those of C. brawnsiana
may be found, while in other extreme variants the marginal outgrowths
take the form of the exaggerated, expanded, dorsiventral lobulue of
certain forms of C. japonica and C. sinensis. But all the intergrada-
tions are present as well, and these minor variations need not be re-
flected in the taxonomy.

This species is restricted to the mountains of southern and south-
eastern Asia and Japan. The elevation data for the known localities
indicate a vertical range of about 1,500-3,600 m. with most of the
specimens coming from about 2,000--3,000 m. Unfortunately the labels
of the specimens give few habitat data, but when a substrate has been
noted, it is usually rock. In reality, however, the habitat ecology of
this species is probably very similar to that of others in the genus.
For example, a particularly rich collection made on shrubs at about
10,500 feet in Nepal by Stainton, Sykes, and Williams consisted of a
mixture of Cetrelia pseudolivetorum, C. olivetorum, C. braunsiana, and
C. collata. Or it might be that distinct ecological races with regard to
substrate preference exist in some correlation with the morphological
plasticity noted above. Systematic field studies might be expected to
elucidate this point.

CHEMICAL ANALYSIS._—Thirty-one samples of this species were tested
in the same manner as described for Cetrelia olivetorum. All contained
atranorin and olivetoric acid.

REPRESENTATIVE SPECIMENS EXAMINED:

JAPAN: Prov. Shinano: Lake Shirakaba, Asahina 58505 & Togashi (DUKE);
Mt. Norikuratake, Asahina 52805 (TN8&). Prov. Shimotsuke: Nikko, Chuzenji,
Nakaji 2510 (TNS). Prov. Suruga: O-miya-guchi, 1-go-han, Mt. Fuji, Asahina
34125 (TNS); Lake Saiko (Nishi-no-umi), near Mt. Fuji, Culberson 10078,
10643 (DUKE).

FORMOSA: Mt. Ali, Mt. Tsu-Tson-San, 2,300-2,900 m., Kurokawa 121, 134,
137 (TNS). Between Su-Su-Lu and Mt. Ali, 200-2,200 m., Kurokawa 611 (TNS).
Mt. Nan-Fu-Ta-San, 1,500-2,400 m., Kurokawa 928 (TNS). Mt. Shin-Kao-San,

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 52]

Mt. Shi-San, 3,300-3,600 m., Aurokawa 332 (TNS); the same, 3,100-3,300 m.,
Kurokawa 253 (TNS); Pianan to Sunlien, 8S. Asahina 3608 (TNS).

CHINA: Moupin (eastern Tibet). Without precise locality, David 1869 (PC, 2
collections).

YUNNAN: Tsang-chan, above Ta-li, 4,000 m., Delavay, 1885 (PC, 2 coilections) ;
the same, Delavay, 1888 (PC, 3 collections); Yen-tze-hay, 300 m., Delavay, 1887,
(PC); Hee-chan-men, Delavay, 1887 (PC); Fang-Yang-Tchang, above Mo-so-yn
Delavay, 1887 (PC); Lopin-chan Pass, 3,200 m., Delavay, 1888 (PC); Lopin-chan
Pass, above Lan-Kong, Delavay, 1888 (PC, 2 collections); Lopin-chan Pass,
Delavay, 1888 (PC).

SIKKIM: Wallanchoon, Hooker 2087 (K).

INDIA: Chakrata, Awasthi 501 (H).

NEPAL: Above Sauwala Khola, 10,500 ft., Stainton et al. 4379b (pro parte)
(BM).

13. Cetrelia sanguinea (Schaer.) Culb. & Culb., comb. nov. PLATE 19
Cetraria sanguinea Schaer. in Moritzi, Zoll., Syst. Verz. Java Pflanz. 129.
1845-1846 (1846). Type: Java, Mt. Pangerano, Zollinger, 1842-1844 (G,

holotype).
Imbricaria sanguinea (Schaer.) Schwend. in Naegeli, Beitr. Wiss. Bot. 3:159.
1863.

Platysma sanguineum (Schaer.) Hue, Ann. Jard. Bot. Buitenzorg 17:177. 1901.

Parmelia megaleia Nyl., Synops. Lich. 1:378. 1860. Type: Java, Junghuhn
(PC, lectotype).

Platysma megaleium (Nyl.) Nyl., Flora 52:290. 1869.

Imbricaria megaleia (Nyl.) Jatta, Nuov. Giorn. Bot. Ital., n. ser. 9:469. 1902.

Platysma subperlatum Nyl., Flora 49:130. 1866. Type: Java, Togoe, Kurz,
1861 (H-Herb. Nylander, lectotype).

Aspidelia wattii Stirt., Trans. & Proce. New Zealand Inst. 32:82. 1899 (1900).
Type: Himalayas, Watt 7070 (BM). (Invalidly published; cf. Culberson,
1966a).

Thallus medium to large, 11-24 em. broad; lobes 0.5-1.5 em. broad.
Upper surface whitish or tan, brownish in old herbarium specimens,
sometimes brownish or blackish toward the margins, smooth to mi-
nutely wrinkled, pseudocyphellate, the pores small, rarely more than 0.5
mm. broad and often inconspicuous or even lacking altogether on some
lobes, the margins of the lobes entire to sometimes minutely lacerate
or beset with lobulae. Lower surface jet-black, the margins often dark
brown, not punctate; rhizines few, black. Upper cortex 16-25 thick,
impregnated with a noncrystalline brown pigment; medulla 93-1704
thick; lower cortex 16-28 thick. Apothecia frequent, very large, per-
forate, cup-shaped when young, irregularly expanded and radially
lacerate with maturity, 1.5-4.2 cm. broad, the thalloid exciple densely
pseudocyphellate; hymenium 60-93 thick; subhymenial layers 43-
61u thick, impregnated with a yellow pigment turning reddish brown
with FeCl;. Spores 8, 17-229-14y, the spore wall about 2p thick.
Pycnidia large, marginal; conidia 14,, rod-shaped.

MEDULLARY REACTIONS.—K —, C+ bright red, PD—.

ConstiTuENTs.—Anziaic acid and atranorin.

522 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Disrripution (Map, Figure 15).—In the high mountains of Japan,
southwestern China, Java, and Sumatra; on the bark of trees and
shrubs.

While Cetrelia sanguinea resembles a number of other species in
habit, the one for which it might most easily be mistaken is C. davi-
diana, the only closely related C+ species. In C. davidiana, however,
the reaction produces a pink coloration instead of the red one char-
acteristi¢ of C. sanguinea and reflects the presence of olivetoric acid
rather than anziaic acid. Of course, microchemical or chromatographic
analyses are required for a precise determination of the substance
responsible for positive reactions with C and critical taxonomic identi-
fications here should not be made without them.

A single Japanese collection containing anziaic acid closely resembles
many of the plants of Cetrelia sanguinea from Java where the species
has been the most extensively collected. It lacks apothecia and has
marginal lobulae somewhat resembling those of C. japonica. A chemical
study of the many specimens referred to C. japonica in herbaria upon
the basis of gross morphology alone might well extend the known
Japanese range of C. sanguinea.

In restricted areas within its range the species may be very common.
For example, there are many specimens in herbaria from Mt. Panger-
ango, the type locality, where collections of the species have repeatedly
been made incidental to other explorations, an apparent testimony to
the abundance and conspicuousness of the lichen there. From labels
giving elevation data, the species in the East Indies seems to be found
between 1,400 and 3,800 m., most of the specimens coming from about
3,000 m. The Japanese specimen is also from a mountain locality.

CHEMICAL ANALYSIS.—-The species is very uniform in the lichen sub-
stances produced. Of the 67 specimens tested microchemically, all
produced atranorin and anziaic acid. Atranorin was usually observed
in the GAo-T solution, and the absence of the colorless needles formed
by olivetoric acid in this solution was noted. Some samples also showed
atranorin in the GE solution. Anziaic acid usually crystallized from
the GAW solution, but less often also from the GE solution. All ex-
tracts were chromatographed to verify the presence of atranorin and
anziaic acid. The crystals of anziaic acid in the GAW solution were
quite variable, but no additional compounds were suspected on the
basis of the chromatographic results. |

The apothecia of this species may contain a yellow pigment (not
usnic acid) which was not identified.

The generic name Aspidelia Stirt., under which Stirton included the
description of a species based upon a specimen referable to C. sanguinea
(see synonymy above), was published without a generic description

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 523

and is thus invalid and cannot be taken up for Cetrelia or for any other
genus (Culberson, 1966a).

ADDITIONAL SPECIMENS EXAMINED:

JAPAN: Honsuv: Mt. Osore-san, Sato, 1935 (H).

JAVA: Mt. Pangerango, 3,000 m., Méller, 1897 (LD, S$); the same, Poch, 1906
(WU); the same, 2,500 m., Schiffner 3007 (M); the same, 3,050 m., Hochreutiner
936 (G); Mt. Gede, Karsten, s.d. (TUR); the same, 2,400 m., Hochreutiner 876 (G) ;
Tjibodas, Mt. Gede, 1,400 m., Palmer & Bryant 822 (US); Mt. Ardjoeno, 3,800 m.,
Groenhart 160 (H); Res. Pasoeroean, Goenoeng Ardjoena, track from Lalidzina to
Ardjoeno-top, Du Rietz 18a:1, 18a:2, 24:4, 51:3 (UPSV); the same, Lalidjina-
Welirang-track, Du Rietz 54:3, 60:3, 60:4, 60:5, (UPSV); the same, Tretes-Lalid-
jina-track, Du Rietz 95:7, 95:17a, 95:17b, 99:1 (UPSV); and the same, Skottsberg
1929 (UPSV, 14 collections).

SUMATRA: Mt. Singalang, 2,500 m., Schiffner 3441 (W, WU).

CHINA: Yunnan: Mekong-Salween-Scheidegebirge, Gebauer, 1914 (W).

PAKISTAN (?): Himalayas, Lingalelah Range, 12,000 ft., Watt 7070 (BM).

14. Cetrelia sinensis Culb. & Culb., sp. nov. PLATE 20

Thallus maximus, 17-26 cm. latus; laciniae 1-2 cm. latae, marginibus
dense instructae vel planta tota saepe cristata lobulis palmatis vel sub-
pal natis, vel angustis et isidioideis, vel expansis et foliosis. Superficies
superior aequaliter pallide cinerea (albida apud plantas vivas?), levis,
pseudocyphellata, pseudocyphello punctato vel nonnihil elongato
parvae (ca. 0.5 mm. latae). Superficies inferior picea, marginibus ad
extremos lacinianum pallida, non punctata; rhizinae nigrae. Cortex
superior 19-28 crassus, subprosoplectenchymatosus, strato corticali
summo ferrungineo-colorato, medulla 140-183. crassa; cortex inferior
16-22u crassus. Apothecia ignota. Pycnidia magna, nigra, in margini-
bus laciniarum; conidia 1.56, recta.

Thallus large to very large, 17-26 em. broad; lobes 1-2 em. broad,
the margins densely fringed with palmately or subpalmately divided
lobulae, narrow and isidioid or more broadly expanded and lobelike, so
dense that the specimens may appear cristate. Upper surface uniformly
light brown (perhaps grayish-white when fresh), smooth, pseudocy-
phellate, the pores punctiform or somewhat elongate, very small (ca.
0.5 mm. broad). Lower surface jet-black, a marginal zone near the
tips of the lobes colored like the upper surface, apparently not punc-
tate; rhizines black. Upper cortex 19-28, thick, subprosoplectenchy-
matous and essentially pseudoparenchymatous, the upper part of the
cortex suffused with a brownish, noncrystalline pigment; medulla
140-183» thick; lower cortex 16-22y thick, prosplectenchymatous.
Apothecia unknown. Pyenidia large, black, restricted to the marginal
lobulae; conidia 1.5 X 6y, rod-shaped.

524 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

‘Typr.—‘ ‘Sur les pierres dans les bois aux sources du Lan-Kien-ho,
prés du col Hee-chan-men,” Yunnan, China, Delavay 30 (PC, holo-
type).

MEDULLARY REACTIONS.—K—, C—, KC—, PD—.

ConsTITUENTS.—I mbricaric acid and atranorin.

DistripuTion (Map, Figure 18)—High mountains of Yunnan,
China, and Formosa; over boulders (and on trees?), apparently very
rare.

Since this species is known from only two collections, it is not pos-
sible to assess its morphological variability. It appears, however, that
it may not be different morphologically from Cetrelia japonica, which
contains microphyllinic acid. Cetrelia sinensis must be identified by the
presence of imbricaric acid. See the discussion for Cetrelia japonica (p.
511) for a further discussion of morphologically similar species and
their distinguishing characteristics.

CHEMICAL ANALYSIS.—The six known specimens (from two collec-
tions) of the species were tested and all contained imbricaric acid and
atranorin. Hydrolysis of the residues from two samples and chromatog-
raphy of the products show a faint spot at a position corresponding to
olivetol-2-carboxylic acid. It may or may not be due to the presence
of perlatolic acid.

ADDITIONAL SPECIMENS EXAMINED:

FORMOSA: Mt. Shin-Kao-San, Mt. Shi-San, 3,300-3,600 m., Kurokawa 449
(TNS).

CHINA: Yunnan: Lan-Kien-ho near Hee-Chan-men, Delavay 30 (PC).

Platismatia Culb. et Culb., gen. nov.

Thallus 5-18(—23) em. latus; laciniae (0.1—)0.3-2.2(-2.5) cm. latae.
Superficies superior pallide cinerea vel citrella, viridiolivacea, vel apud
exemplaria vetera herbarii aequaliter obscure citrella, saepe rugosa,
pseudocyphellata vel isidiata sed raro sorediata. Superficies inferior
nigra, rhizinata, saepe punctata. Cortex superior (15—)18-28(82)u
crassus, prosoplectenchymatosus; medulla (30-)65-210(-280) crassa,
alba; cortex inferior (10—-)13-28(-30) crassus. Apothecia (0.1—)0.4—-
1.5(—4.0) em. lata, marginalia vel submarginalia, vulgo perforata;
hymenium (34-)39-70(-125)u crassum; asci IKI+ caerulescentes;
strata subhymenialia (13—)18-50(-78) crassa, IK1+ caerulescentia,
viridicaerulescentia, lavendulina, vel purpurea, subtus fere sine algis.
Sporae 8-nae, parvae, ellipsvideae vel subglobosae, (3.5-)5-8(-10)
<3-5yu. Pyenidia marginalia, immersa, saepe deficientia; conidia rara,
1<5(-7)p, recta, extremis non inflatis. Typus generis P. glauca est.

Thallus 5-18(-23) em. broad; lobes (0.1—-)0.8-2.2(-2.5) em. broad.
Upper surface ashy-white, tan, or greenish-brown (uniformly dull tan

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 525

in old herbarium specimens), often rugose, pseudocyphellate in some
species, isidiate in some species, rarely sorediate. Lower surface black,
with few rhizines, punctate in some species. Upper cortex (15—) 18-28
(-32)y thick; medulla (80—)65-210(-280)y thick; lower cortex (10-)
13-28 (-30)u thick. Apothecia (0.1—)0.4-1.5(—4.0) em. broad, marginal
or submarginal, commonly perforate; hymenium (34-)39-70(-125)n,
the asci IKI+ blue; subhymenial layers (13-)18-50(—78)y thick,
IKI+ blue, blue-green, lavender, or purple, usually without under-
lying algae. Spores 8, small ellipsoid or subglobose, (3.5)5-8(-10) x
3-5y. Pycnidia marginal, immersed, often absent; conidia rare, 5-(7) X
1u, rod-shaped, the ends not inflated.

ConsTITUENTs.—Atranorin and caperatic acid; fumarprotocetraric
acid in one species.

Certain authors have referred some members of the Cetrarza glauca
group, which constitutes the genus Platismatia, to the genus Platisma
(sometimes written as the orthographic variant “‘Platysma”’). Platisma
as a generic name has probably been associated primarily with the
species Cetraria glauca more than with any other species, at least in the
minds of most lichenologists, and we hoped that it could be used for the
Cetraria glauca group essentially in the sense that Dahl (1952) had
proposed earlier. The genus had not been typified, however, and when
an exhaustive nomenclatural study of the Platisma names was made
(Santesson and Culberson, 1966), it was impossible to typify any of
them in such a way that they could be used for this or any other group
of plants. The new generic name Platismatia is a madeup derivative
from the old name Platisma.

Artificial Key to the Species of Platismatia

1. Thallus isidiate or sorediate.
2.Upper surface or margins of lobes isidiate or sorediose-isidiate.

3. Some lobes becoming narrow, coralloid, fruticose appendages of the nor-
mally foliose thallus; upper surface not pseudocyphellate; lower surface
not punctate.

4. Lobes of the foliose part of the thallus expanded, commonly more than

5 mm. wide; broadly distributed. . . . . 3. P. glauca (some forms)
4, Lobes of the foliose part of the thallus narrow, strap-shaped, less than 5
mm. wide; northwestern North America. . 4. P. herrei (rare forms)

3. All the lobes broadly foliose, none becoming coralloid or fruticose; upper
surface pseudocyphellate; lower surface punctate or not.

5. Upper surface with prominent, vertical, reticulate ridges, the isidia con-
fined primarily to the crests of the ridges; thallus IKI — ; lower surface
not punctate; North America and Europe. . . . 7. P. norvegica

5. Upper surface without prominent reticulate ridges but often with poorly
defined, interrupted ones; thallus IKI— or IKI-+-; lower surface
punctate or not.

6. Upper surface pseudocyphellate, IKI— or IKI+; lower surface punc-
tate or not; isidia rarely on the margins of the lobes; Asia.

526 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

7. Pseudoeyphellae minute, inconspicuous, usually confined to the
crests of the thallus ridges; thallus surface IKI—; lower surface
usually punctate; mountains of southern Asia and Japan.

1. P. erosa

7. Pseudocyphellae large and conspicuous, round to elliptical, seat-
tered; thallus surface IKI-+ ; lower surface not punctate; Japan.

5. P. interrupta
6. Upper surface not pseudocyphellate, [XI-+ ;lower surface not punctate ;
isidia commonly on the margins of the lobes; America or Europe.

8. Lobes broadly foliose, not strap-shaped, commonly more than 5 mm.

wide; widely distributed. . . . . . 3. P. glauca (some forms)
8. Lobes narrow and strap-shaped, less than 5 mm. wide; northwestern
North America... toe eee ee ee 64 PL herrei

2. y pper surface sorediate, not isidiate.
9. Upper surface conspicuously pseudocyphellate; Japan.
5. P. interrupta (some forms)
9. Upper surface not pseudocyphellate; widely distributed but absent from
Japan... ... . . 38 P. glauca (some forms)
Thallus neither isidiate nor sore ‘ediate.
“10. Upper surface with sharply defined, continuous, reticulate ridges, not pseu-
docyphellate; medulla of American plants PD+ orange-red.

11. Medulla PD —; lower surface punctate; Formosa. . 2. P. formosana
11. Medulla PD+ orange-red; lower surface not punctate; northwestern
North America... ... . . 6 P. lacunosa

10. Thallus without sharply de fined reticulate ridges but often with inconspic-
uous, interrupted ridges and folds or thallus pitting, pseudocyphellate
or not; medulla PD—.

12. Upper surface pseudocyphellate; apothecia unknown; Asia.

13. Pseudocyphellae large and conspicuous; upper surface not commonly

blackened; lower surface not punctate; Japan. . 5. P. interrupta

13. Pseudocyphellae small; upper surface or margins commonly darkened

or mottled with black; lower surface punctate; Asia.

14. Thallus surface reticulately lined, the pseudocyphellae associated
with the lines; mountains of southern Asia and Japan.

1. P. erosa (nonisidiate forms)

14. Thallus surface without conspicuous lines; the margins of some lobes

blackening conspicuously; Sabah (Borneo). . . 8 P. regenerans

12. Upper surface not pseudocyphellate; apothecia common; North America.

15. Lobes narrow and strap-shaped, rarely reaching 5 mm. broad; western

North America. . . ... . . 9 P. stenophylla

. Lobes broadly expanded, usually. excee eding 5 5 mm. broad; eastern North
America. . 2... 2... 10 iP. tuckermanii

1. Platismatia erosa Culb. & Culb. sp. nov. PLaTE 20

Cetraria formosana var. isidiata Zahlbr., Repert. Spee. Nov. Fedde 33:60.
1934. Type: Mt. Niitaka (=Mt. Morrison), Formosa, Sasaki (W, holotype).
Thallus mediocris, ad 11-12 em. latus; laciniae 0.5-2 cm. latae. Super-
ficies superior sordide citrella vel pallide grisea, marginibus lacimiarum
olivacea, reticulata, reticulo, saepe nigrescente; isidiata, isidis fere
raris, brevibus, pinquibus, simplicibus et in reticulo positis sed coral-
loideis, in marginibus laciniarum. Superficies inferior punctata, picea,

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 527

marginibus fusca vel pallida; rhizinae paucae, simplices vel fasciculatae,
nigrae, solum in partibus vetustioribus thalli. Cortex superior 22-26
(-28)u crassus; medulla 50-90(-114)u crassa, IKI—; cortex inferior
16-21(—26)u crassus. Apothecia rarissima, ad 7 mm. lata, submar-
ginalia, perforata; hymenium 39-47 crassum; asci IKI+ intense
caerulescentes; strata subhymenialia 21-29y crassa, IKI+ lavendula;
sporae 8-nae, ellipsoideae, 3-4. 5-8. Pyenidia ignota.

Thallus medium, to 11-12 cm. broad, loosely attached; lobes 0.5-2
em. broad, broadly rounded. Upper surface light tan or ashy-gray,
tinged with brown at the margins of the lobes, broadly reticulately
ridged and veined, the crests of the ridges often darkening, and pseudo-
cyphellate with minute pores and often bearing isidial scars resembling
pseudocyphellae; isidiate, the isidia usually infrequent, short, stout, or
simple and usually confined to the ridges or thallus cracks but becom-
ing coralloid and well developed at the margins of the lobes in some
specimens. Lower surface punctate, jet-black, the marginal zone light
brown or tan; rhizines few, simple or fasciculate, black, confined to
the older parts of the thallus. Upper cortex 22—26(-28) uy thick; medulla
50-90(-114)y thick, IKI— ; lower cortex 16-21 (—26) yu thick. Apothecia
very infrequent, to 7 mm. broad, submarginal, perforate; hymenium
39-47 thick, the asci IKI-+ dark blue; subhymenium 21—29u thick,
IKI+ lavender, without underlying algae; spores 8, ellipsoid, 3-4
5-8. Pycnidia unknown.

Typse.— Yatsu-ga-take, Mount Kitayoko-dake, Province of Shinano,
Japan, Kurokawa 58349 (TNS, holotype).

MEDULLARY REACTIONS.—K—, C—, KC—, PD—.

CONSTITUENTS. Caperatic acid, atranorin, and an_ unidentified
yellow pigment.

Distrisution (Map, Figure 30).—Mountains of Honshu, Japan,
Formosa, Philippines, Java, Vietnam, Sikkim, and Nepal; on tree
trunks,

Platesmatia erosa is distinguished by the broadly ‘‘eroded”’ reticula-
tion of its upper surface, the pseudocyphellae and isidial scars asso-
ciated with the thallus ridges, and the puncta on the lower surface.

In Japan some specimens of Platismatia erosa have previously been
referred to Cetraria formosana as var. isidvata, but the plants differ
from P. formosana in a number of ways. The thallus folds and ridges,
sharp and conspicuous in P. formosana, are less distinct in P. erosa.
Platismatia formosana never has isidia, and P. erosa is typically isidiate,
although rarely richly so, and some fragmentary or poorly developed
specimens may lack isidia altogether. The crests of the thallus ridges
in P. erosa always bear at least a few pores. While most of these pores
are doubtless pseudocyphellae, some of them seem to be scars where

528 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

isidia have broken off. Holes in the upper cortex such as these, regard-
less of their origin, are unknown in P. formosana.

Platismatia erosa also superficially resembles P. norvegica although
their ranges do not overlap anywhere. It differs from P. norvegica in
having pseudocyphellae on the upper thallus surface and puncta on the
lower surface while the latter species never produces these pores. Al-
though reticulate on the upper surface, P. erosa lacks the delicate yet
sharp and continuous network that ornaments the surface of 2. norve-
gica. The species also differ in many minor characteristics such as the
morphology of the isidia. The isidia of P. erosa are short and very
coarse and usually simple while those of P. norvegica are slender, deli-
cate, and often coralloid.

Platismatia erosa as it is represented in Japan and in the high moun-
tains of the mainland of southern Asia is a very clearly defined species
of conservative variation. Since most of the specimens seen in this
study were from these regions, a few somewhat different-looking plants
from the Philippines and Java are assigned to this species with reluc-
tance and qualification. The anomalous southern plants differ in their
habit by a number of subtle, hard-to-describe ways from those of the
main part of the distribution to the north. They have larger and more
abundant pseudocyphellae on the upper surface, fewer puncta on the
lower surface, more richly developed isidia, and less well defined thallus
ridges and reticulations. These southern plants (especially the Java-
nese ones) when considered together, however, seem to be quite uni-
form. Perhaps they belong to a distinct taxon.

Some individuals of Platismatia erosa, and especially the atypical
southern specimens provisionally referred to this species, may also
superficially resemble P. regenerans from Sabah. The latter species,
however, has no isidia, and it has pseudocyphellae which are larger and
more conspicuous than those of typical P. erosa. It also lacks the ridg-
ing seen in P. erosa, and its upper surface is entirely smooth.

CHEMICAL ANALYSIS.—Of the 17 samples tested, nine were extracted
directly with acetone. The residues from these extracts all showed
caperatic acid in both the GE solution (two erystal forms) and the
GAW solution, and atranorin in the GE and GAo-T solutions. Resi-
dues from eight samples extracted with cold benzene showed atranorin
in the GE and GAo-T solutions (Plate 2, Figure 7) but no protol-
chesterinic acid in the GE solution. The residue from a subsequent ace-
tone extract gave the characteristic crystal forms of caperatic acid in
the GE solution (Plate 3, Figure 18) and in the GAW solution (Plate
2, Figure 11). In addition to atranorin and caperatic acid, nine samples
showed yellow crystals in the GE solution (Plate 6, Figure 35) which
may be due to an unidentified yellow pigment. Some residues contain-
ing the yellow pigment also showed clusters of fine yellowish-tan nee-

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 529

dles (Plate 6, Figure 36) in the GAo-T solution. The same combination
of yellow crystals in the GE solution and fine needles in the GAo-T
solution was observed in preparations from some samples of Platis-
matia interrupta, P. formosana, and P. regenerans, suggesting that the
same substance or combination of two substances occurs in these
species as well.

ADDITIONAL £PECIMENS EXAMINED:

JAPAN: Honsuu: Prov. Kai: Mt. Kobushi, 2,000 m., Asahina, 1933 (TNS, 3
collections). Prov. Shinano: Mt. Tadeshina, Kurokawa 510038 (TNS); Yatsu-ga-
take, Mt. Tengu, Kurokawa 58240 (TNS, US); Yatsu-ga-take, Takamic-ishi,
Kurokawa 58301 (TNS); Yatsu-ga-take, Mt. Kitayoko-dake, Kurokawa 58349
(TNS); Yatsu-ga-dake, Mt. Naka-no-dake, Togasht & Kurokawa (TNS). Prov.
Musashi: Mt. Kobushi, Pass Karisaka, Kurokawa 521018 (TNS).

FORMOSA: Mt. Niitaka, Sasaki, 1927 (W).

PHILIPPINES: Mowuntain Prov.: Mt. Data, Barrio, Hale 26301, 26751
(US); the same, Heald Logging Area 9, Hale 26082, 26105 (US).

JAVA: Res. Pasoeroean, Goenoeng Ardjoena, Lalidjina-Welirang-track, Du
Rietz 61:2, 61b:2, 62:1, 65:3, 68:1 (UPSV); the same, track from Lalidjina to
Ardjoeno top, Du Rietz 36:1 (UPSY).

VIETNAM: Langbian Mountains near Dalat, 1,800 m., Tizier, 1962 (DUKE).

CHINA: YitLinexone: Gomba La, 3,700 m., Smith 14062 (UPS).

SIKKIM: No locality, Thomson (H-Herb. Nyl., K, M); no locality, Thomson,
289 (G,M). East Himalayas, 8,500 ft., Awasthi 354 (H).

NEPAL: East Nepal, Mewakhola Valley, 11,000-11,500 ft., Awastht 2292
(UPS) ; Khumbu, 3,900-4,000 m., Poelt L36, L37 (M).

2. Platismatia formosana (Zahlbr.) Culb. & Culb., comb. nov. PLATE 25
Cetraria formosana Zahlbr., Repert. Spee. Nov. Fedde 33:59. 19383-1934
(1934). Type: Formosa, Mt. Arisan, Asahina F.89 [not ‘84’ as in the orig-

inal citation]. (W, holotype).

Thallus small to medium, 5-19 ecm. broad; lobes 0.3—2.5 cm. broad.
Upper surface ashy-white or gray (uniformly brown in old herbarium
specimens), abruptly interrupted with broad reticulate ridges up to 1
mm. high at right angles to the surface, not pseudocyphellate. Lower
surface black toward the center, but large zones, especially toward the
margin, chestnut-brown, punctate, the entire lower surface covered
with abundant, tiny pores; rhizines few, black, simple or compound-
fasciculate. Upper cortex 19-31, thick; medulla 29-102 thick between
the ridges, IKI—; lower cortex 10—-25y thick. Apothecia infrequent,
submarginal, perforate, 1.5-4.0 mm. in diameter; hymenium 39-47
thick; subhymenial layers 18—26y., without underlying algae. Spores 8,
small, ellipsoid, 5-7 3-4u. Pycnidia marginal, immersed, only the
ostiole visible; conidia 15-74, rod-shaped, the ends not inflated.

MEDULLARY REACTIONS.—K—, C—, KC—, P—.

ConsTITUENTS.—Caperatic acid, atranorin, and an unidentified
yellow pigment.

Distrinution (Map, Figure 28).—Mountains of Formosa.

530 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

The thallus of Platesmatia formosana has a broad and severe reticula-
tion that is particularly striking. The long, continuous ridges are com-
posed of both cortex and medulla and may be as high as 2 mm. with
perfectly vertical sides and without correspondingly deep depressions
in the lower surface of the thallus. The sharpness of the ridges is strik-
ing in comparison to the worn, more gentle ridges of P. erosa, although
the overall pattern of ridging of the thallus surface is highly similar and
the species are obviously very closely related.

Platismatia formosana might be compared with P. lacunosa, but
their patterns of thallus ridging are quite different. In P. formosana the
spaces between the broadly separated, vertical ridges are flat while in
P. lacunosa the sloping ridges are separated by U-shaped valleys. The
two species can be separated immediately by their medullary reactions
with p-phenylenediamine: P. formosana is negative while P. lacunosa
is brick-red due to the presence of fumarprotocetraric acid.

The species is known only from the mountains of Formosa and
earlier records for it from Japan (Sato, 1943) are doubtless all referable
to Platismatia erosa.

CHEMICAL ANALYSIS.—Two samples were tested and both showed
atranorin and caperatic acid. Both erystal forms of caperatic acid were
observed in the GE recrystallization of the acetone extract. Both sam-
ples also showed a yellow pigment (not usnic acid) soluble in warm
benzene and recrystallizing from the GE solution. The crystals in this
solution resemble those from P. erosa (Plate 6, Figure 35), and one
sample also gave the clusters of fine needles in the GAo-T solution that
seemed to be the same as those associated with the occurrence of the
yellow pigment in P. erosa and P. interrupta (Plate 6, Figure 36). The
same yellow pigment may also occur in P. regenerans.

ADDITIONAL SPECIMENS EXAMINED:

FORMOSA: Mt. Arisan, Numanodaira, above 7,000 ft., Asahina F.89 (W);
Mt. Niitaka (Mt. Morrison), Sasaki, 1927 (W), Suzuki, 1935 (11); the same,
Hattsu-kan, Suzuk7, 1935 (TNS, 3 collections); Mt. Daibu, Yasuda 448 (II).

3. Platismatia glauca (L.) Culb. & Culb., comb. nov. PLATE 21

Lichen glaucus L., Sp.P1.1148. 1753. Type: Europe, Linnean Herbarium Sheet
1273-139 (LINN, lectotype).

Cetraria glauca (L.) Ach., Method. Lich. 296. 1803.

C. glauca var. chilensis Ris., Ann. Bot. Zool.-Bot. Fenn. Vanamo 20(3):4.
1955. Type: Chile, Prov. Chiloé, Petrohué, Hosseus 167 (M, isotype).

Lobaria glauca (L.) Hoffm., Deutsehl. Fl. oder Bot. Taschenb., Theil fiir das
Jahr 1795, Cryptogamie. 149. 1795 (1796).

Parmelia glauca (L.) Hepp, Lichenen-Fl. Wiirzburg. 23. 1824.

Physcta glauca (L.) DC. in Lam. & DC., Fl. Frang., ed. 3, 2:401. 1805.

Platysma glaucum (L.) Frege, Deutschl. Bot. Taschenb., 2. Theil fiir das Jahr
1812. 167. 1812. [Not seen.]

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 53]

Thallus medium, 8-15 em. broad; lobes 0.3-2.0 em. broad, the mar-
gins with soredia, simple or coralloid isidia, or both, or branched and
divided into fruticose lobes, which in turn may be sorediate or isidiate
or both. Upper surface whitish or whitish-green or tinged with yellow
or, especially in South American specimens, very distinctly yellow,
often with darkened areas, smooth to strongly reticulately wrinkled,
not pseudocyphellate, smooth or with coralloid or sorediose isidia in
discrete patches or diffusely scattered. Lower surface jet-black, the
margins brown, colored like the upper surface or white, smooth or
reticulately wrinkled and pitted, not punctate; rhizines few to many,
simple or compound, brown or black. Upper cortex 16-26u thick,
prosoplectenchymatous; medulla white, 62-155(-200)u thick, IKI+
lavender; lower cortex 16-254 thick, prosoplectenchymatous.
Apothecia very rare, 0.5-1.0 cm. broad, marginal, perforate or not;
hymenium 34-56 thick. the asci IKI-+ blue or blue-green: subhy-
menial layer 16-52y thick, IKI+ lavender or bright purple, not
underlain by algae; spores 8, small, ellipsoid to ovoid, 3.5-8.5 X 3-5.
Pyenidia not seen. (See the taxonomic discussion below.)

MEDULLARY REACTIONS.-K—, C—, KC—, PD—.

CONSTITUENTS Japeratic acid and atranorin.

Disrripution (Map, Figure 22)—Southern Greenland, northeastern
and northwestern North America and coastal Alaska, Patagonia,
central Asia (U.S.S.R.) west through all of Europe, the Canary Islands
and the Azores, and in the high mountains of Kenya and Tanzania; on
tree trunks and on rocks and boulders.

No other species treated in this monograph has a broader range of
morphological variation than Platismatia glauca. There is a bewildering
range of forms from plants with lobes bearing soredia to sorediose isidia
to coralloid isidia to isidioid fruticose branches. All these traits, how-
ever, intergrade, and all possible combinations of characters can be
found in the herbarium, although certainly not at every locality in the
field where the species occurs. If only the extremes of the variations
were seen, it would be unbelievable that all the forms constituted a
single species. In spite of the plastic morphology of this species, far
fewer infraspecific taxa of it have been described than one might ex-
pect, considering the taxonomic tradition in Europe. All this reflects,
we believe, a general recognition by lichenologists of the innate con-
tinuity of the variation in this doubtless very ancient species.

When a large series of specimens is examined or when the species is
studied in the field, one is tempted to see distinct local races. For
example, the eastern North American populations seem to have a
somewhat shiny thallus surface as compared to that of the western
North American populations which seem to be much duller. These
apparent differences, however, could not be associated with any con-

932

CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

22

eo be?

aoe

ORR. *
A, a an
a i
.
on Oe: i

Spee

on

Figures 22-25. Distributions.

lacunosa (Ach.) Culb. & Culb.

PP)

“oe.

stenophylla (Tuck.) Culb. & Culb. 24. P. herret (Imsh.) Culb.

Platismatia glauca (L.) Culb, & Culb. 23,
& Culb. 25.

P.

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 5338

stant or demonstrable structural or chemical differences. Similarly,
the South American plants in general look more distinctly yellow than
those from the Northern Hemisphere. No chemical explanation for this
color difference could be found.

In the present study no pycnidia of this species were seen. Hillmann
and Grummann (1957), however, note in the description of this species
in their flora of Brandenburg: ‘‘Pykniden selten, randstindig, braun
oder schwirzlich. Pyknokonidien gerade, an einem Ende verdickt,
6-7 «0.8-1.0n.”

It isnot surprising that the species showing the greatest morpholog-
ical variation of all those treated here should also have the broadest
geographic range. Platismatia glauca is a boreal to temperate species,
but it is conspicuously absent from eastern Asia where it is replaced by
other members of the genus, notably P. interrupta and P. erosa. In
Africa it is known with certainty only from Mt. Aberdare in Kenya
and Mt. Kilimanjaro in Tanzania, although Flagey (1896) cites it
from the Atlas Mountains of North Africa. These localities, moreover,
are the only certain African stations for any species of ‘‘Cetraria” in
the broad sense.

The map of the total range of Platismatia glauca (Figure 22) is only
approximate. Too few Soviet specimens with adequate geographic in-
formation were seen to allow the construction of a meaningful map for
eastern Europe and central Asia. The outlined eastern part of the range
of the species in the Old World is based on a map published by Rassa-
dina (1950) in her careful study of Cetraria in the U.S.S.R. Since there
are no differences in the taxonomy of P. glauca at the species level
between her treatment and ours, we have used directly the information
that she gives. Also, in the range as indicated for western North
America, the real distribution may actually extend a bit farther north
than we have mapped it.

Although Platismatia glauca is widely distributed, it is rare in large
parts of its range, for example in the southern parts of its distribution in
North America. In Europe the species is doubtless by far most common
in the milder western parts and in the British Isles.

CHEMICAL ANALYSIS.—Eleven samples from throughout the range
of the species were tested and all contained caperatic acid but no proto-
lichesterinic acid. The cold benzene extract showed atranorin in the
GE and GAo-T solutions but no recognizable fatty acid in the GE or
GAW solutions. The subsequent acetone extract contained caperatic
acid (GE, GAW, and GAo-T). Tests in the GE solution gave mixed
crystal forms of caperatic acid (Plate 3, Figures 14 and 16). The ace-
tone extracts were also chromatographed in pyridine-ethyl acetate-
water (1:2:1) and sprayed with a stable PD solution, but no PD+
compounds other than atranorin were detected.

534 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

NoMENCLATURE.—The oldest specific epithet for this polymorphic,
widely distributed lichen is Lichen glaucus, from Linnaeus’ Species
Plantarum (1753). There are, however, several other old specific
epithets that also refer to Cetraria glauca (L.) Ach. sens. lat.: Lichen
fallax Web. (Spicilegium Florae Goettingensis. 244. 1778), Lichen
fraudulentus Gmel. (in L., Syst. Nat., ed. 18, 2 [2]:1371. 1791), and
Lichen membranaceus Lam. (Encycl. Méthod., Bot. 3: 492. 1789 [1791-
1792]). Zahlbruckner (1930) lists nomenclatural synonyms of all these
basionyms. None of these old names has been typified in the present
study because none is required for the present taxonomic treatment.
In the event that a future interpretation recognizes more than one
species, it will be necessary to determine the precise reference through
typification of these names before assigning specific epithets to segre-
gates of Platismatia glauca sens. lat.

In addition to the above names, the epithet Lichen ampullaceus L.
(Sp. Pl. 1146. 1753) has traditionally been assumed to refer to Platis-
matia glauca. There is no specimen bearing the epithet ampullaceus in
the Linnean Herbarium. From the original description (‘Lichen foli-
aceus planiusculus lobatus crenatus, peltis globosis inflatis”), however,
the name has often been interpreted to refer to monstrous forms of P.
glauca infected by parasitic fungi (cf. Santesson and Culberson, 1966).
If so, the name can be rejected as based upon a monstrosity.

REPRESENTATIVE SPECIMENS EXAMINED:

GREENLAND: Jutianenaas District: Uperniviarssuk, Dahl, 1937 (US);
Qagssiarssuk, Dahl, 1937 (US).

MIQUELON ISLAND: Locality unknown, Delamare, s.d. (COLO).

CANADA: LaBrapor: Bottle Harbor, Waghorne, 1882 (US).

NEWFOUNDLAND: St, John, Fries, 1871 (S); Codroy Valley, Oregans, Hall 34, 35,
36, 41 (WIS).

QuEBEC: Gaspé-Est: Bonaventure Island, off Pereé, Imshaug 26501 (MSC),
Shushan 823026 (COLO). Montmorency: Laurentides National Park, Lac Jacques
Cartier, Imshaug 26336 (MSC). Matane Co.: Mont Blane, Frere Fabius, 1953
(DUKE, 4 collections). Frontenac Co.: Mont Megantic, Masson 6068 (WIS).

Nova Scorra: Chester, Greenwood 1558 (COLO- ABS).

New Brunswick: Albert Co,: Fundy National Park, Herring Cove, Jmshaug
26532, 26585 (MSC), .

Onrario: Groundhog River, west of Timmins, Wetmore 6247 (COLO, MSC,
US).

SASKATCHEWAN! Cypress Hills, Lamb 6133 (DUKE, WIS).

ALBERTA: Banff National Park, Johnson Creek, Weber 823560 (COLO).

BrivisH COLUMBIA: Queen Charlotte Islands: Graham Island, Persson (COLO).
Vancouver Island: Locality unknown, Lyall, 1858-1859 (US, 2. collections) ;
Comox, Macoun 88 (CAL, MSC, US, WIS).

U.S.A.: Maine: Hancock Co.: Mt. Desert Island, Davis 21, 46 (US). Knox Co. :
Camden, Merrill, Lich. Exs. 139 (US). Piscataquis Co.: Mt. Katahdin, Degelius,
1939 (US). Washington Co.: Cutler, Davis 14g (US); North Lubec, Cummings,
Decades North Amer. Lich. 279 (MSC, US).

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 535

Figures 26-30. Distributions. 26. Platismatia interrupta Culb. & Culb. 27. P.
norvegica (Lynge) Culb. & Culb. 28. P. formosana (Zahlbr.) Culb. & Culb. 29. P.
regenerans Culb. & Culb. 30. P. erosa Culb. & Culb.

New HampsuirE: Coos Co.: Mt. Washington, Llano, 1941 (WIS). Grafton Co.:
Mt. Lafayette, Kazser 64 (COLO-ABS).

VERMONT: Lamoille Co.: Mt. Mansfield, Cain 25549 (COLO), Merrill 1922
(DUKE).

Massacuvusetts: Berkshire Co.: Mt. Greylock, Imshaug 24095 (MSC).

New York: Ulster Co.: Cornell Mountain, Imshaug 24062 (COLO, MSC).

TENNESSEE: Sevier Co.: Mt. Le Conte, Degelius, 1939 (US, 2 collections).

NortH Carouina: Avery Co.: Roan Mountain, Hale 18086 (US).

MicuiGan: Keeweenaw Co.: Isle Royale National Park, Wetmore 2509 (COLO,
DUKE, MSC).

Minnesota: Cook Co.: Susie Islands Thomson 4023 (WIS).

Sout Dakota: Lawrence Co.: Benchmark, Wetmore 8503 (MSC).

Montana: Flathead Co.: Glacier National Park, Fish Lake, Imshaug 7287
(WIS). Missoula Co.: South of Missoula, Stickney PFS-573 (WIS). Sanders Co.:
South of Thompson Falls, Dry Creek, Stickney 1005, 1005a, 1005b (WIS).

Wyomina: Crook Co.: Taylor Divide Road, Wetmore 9824 (MSC).

Cotorapvo: Boulder Co.: Bull Gulch, Erdman, as Weber, Lich. Exs. 4 (COLO,
DUKE, MSC, WIS).

536 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

ALASKA: Prince William Sound, Thum Bay, Knight Island, Eyerdam 254, 282,
284 (CAL). Washington Bay, Kuiu Island, Eyerdam 1080, 1086, 1142 (COLO).
Kenai Peninsula, Seward, Hultén, 1932 (UPS); Admiralty Island, Windfall Har-
bor, Stephens 243 (CAL).

WasuHinaton: Clark Co.: 5 mi. east of Vancouver, Pechanec S4717 (COLO).
Island Co.: Langley, Grant, 1923 (WIS, 2 collections). Jefferson Co.: Olympic
Rain Forest, Howard 4317 (WTU). King Co.: Red Mountain, Eyerdam 1557
(COLO, WTU). Lewis Co.: Tenina, Frye (COLO-ABS). Pierce Co.: Mt. Rainier
National Park, White River, Imshaug 391 (US, WIS). Snohomish Co.: Marysville,
Grant, 1929 (CAL, US). Spokane Co.: North of Spangle, Cooke 23203 (COLO -ABS,
WIS). Stevens Co.: Springdale, Cooke 23237 (WIS). Thurston Co.: Olympia,
Becking 5203462d (WTU). Whatcom Co.: Maple Mead Farm, Herre, 1949 (WIS).
Whiteman Co.: Kamiak Butte, /mshaug 16446 (MSC).

OrxEGON: Benton Co.: Corvallis, FE. J. K., 1921 (WIS). Hood River Co.: Cloud
Gap, Hale 21558 (DUKE). Jackson Co.: Medford, Darrow 233 (WIS). Lane Co.:
Oakridge, Howard 4713 (WTU). Union Co.: Mill Creek, Sheldon 9010 (US).

Ipano: Bonner Co.: Sandpoint, Knor 815842, $15844 (COLO). Idaho Co.: Nez
Perce National Forest, Cooke 20694, 20708 (WIS). Nez Perce Co.: Clearwater
Canyon, Cooke 22910, 22918 (WIS). Lattah Co.: St. Joe, Herre 7457 (COLO,
WIS). Lewis Co.: Winchester, Cooke 25107 (WIS).

CaLiForNIa: Siskiyou Co.: Locality unknown, Herre, 1942 (WIS). Sonoma Co.:
Santa Rosa, Baker, 1897 (UC).

CHILE: Mauueco: Pidir-Nalmel, Sparre 5092 (S); Angol, Dusén, 1896 (UPS).

Cavutin: Voledin Llaima, Sparre 4826 (5).

Vaupivra: Lago Panguipulli, Santesson 3943 (UPS); Lago Riiihue, Enco, San-
tesson 7249, 7273, 3607 (UPS).

Ays&n: Coyhaique, Santesson 4397, 4453, 4484 (UPS); Puerto Aysén, Sarttesson
4323.

CuiLok: Petrohué, Hosseus 167 (M).

MaGatuanes: Isla Riesco, Santesson 2084, 9011a, 8198 (UPS); Lago del Toro
(Maravilla), Santesson 6330, 6505 (UPS); Tierra del Fuego, Canal Whiteside,
Puerto Yartou, Santesson, Lich. Austroamer. 469b (LD, MSC, UPS); Canal Beagle,
Santesson 7445 (UPS); Isla Hoste, Santesson 1333 (UPS); Punta Arenas, Santesson
5116 (UPS).

ARGENTINA: Rfo Necro: Lake Nahuel Huapi, Lamb 5834 (UPS); Lago
Frias, Lamb 5984 (UPS).

TierRA DEL Fusco: Lago Fagnano, Santesson 1583 (UPS); Ushuaia, Dusén 247
(UPS), Dusén, 1896 (GB, 8, UPS), Santesson 433, 1215 (UPS), Skottsberg, 1902 (8,
3 collections), Santesson, Lich. Austroamer. 469a (LD, MSC, UPS); Sierra Alvear,
Santesson 706 (UPS).

NORWAY: Horpauanp: Granvin, Havaas, 1910 (DUKE).

S6RTRONDELAG: Leinstrand, Hoég, as Magn., Lich. Sel. Scand. Exs. 260 (GB, 8).

SWEDEN: Dacarne: Lima, Hasselroé (8).

JAMTLAND: Are, Hasselrot (8).

SMALAND: Bunn, Santesson, 1932 (5).

VasrerGOrLaNp: Koliby, Tridet, Vrang, Krypt. Exs. Vindobon. 2763 (MSC,
UC).

VARMLAND: N. Ny, Hasselrot, 1938 (S).

FINLAND: Ere.a-Hame: Lammi, Ulvinen, 1960 (UC).

Pousors-Kargaua: Keitele, Vuonamo, Huuskonen, 1960 (UC).

U.S.S.R.: Esronta: Eesti, Kedder, Wasmuth, 1910 (H); Distr. Keila, Vaina,
Parn, 1960 (DUKE); Tallin-Piiskula, Pdrn, 1960 (DUKE); Insula Saaremaa,
Siinnaa, 3.d. (DUKE).

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 537

Liravania: Peninsula Kura, Trass, 1959 (DUKE).

S1BERIA: Sajangebirge, Plotnikov, 1927 (DUKE).

TURKEY: Mt. Ida, Cyrén, 1933 (S).

SCOTLAND: INVERNESS-SHIRE: Moidart Peninsula, Shiel Bridge, Culberson &
Culberson 11827 (DUKE).

ARGYLLSHIRE: Ardnamurchan Peninsula, Kilechoan, Culberson & Culberson
11586 (DUKE).

WALES: BrEcoNSHIRE: Hay-on-Wye, Culberson & Culberson 11859 (DUKE).

CAENARVONSHIRE: Gwynant Lake, Culberson & Culberson 11926 (DUKE).

RADNORSHIRE: Penybont, Culberson & Culberson 11851 (DUKE).

DENMARK: Jy.uanp: River Gudenna, Borre Forest, Leth, 1870 (UC).

GERMANY: Scuieswic-HotsteEtn: Schleswig, Sazen, 1957 (DUKE).

BReEIsGAu: Freiburg, Lésch, as Arn., Lich. Exs. 1723 (PC).

HOLLAND: GELDERLAND: Bermekon, Maas Geesteranus 2480 (S).

CZECHOSLOVAKIA: Banska Stiavnica, Finkova, Suza, Lich. Bohemoslov 56
(DUKE).

GREECE: TueEssauta: Mt. Olympos, Szatala, as Kofaragé-Gyeln., Lichenoth.
193 (BM).

PELOPONNESOS: Measina, Degelius, 1955 (GB).

FRANCE: ArRpENNEs: Montherme, Culberson & Culberson 10530 (DUKE).

FInisTéRE: Roc Trévezel, Culberson & Culberson 10503 (DUKE).

HavutE-GaRONNE: Lac d’06, Culberson & Culberson 11895 (DUKE).

Havrtes-PyrénkeEs: Cauterets, Jeanjean 7771 (DUKE); Pont d’Espagne, Cul-
berson & Culberson 11713 (DUKE).

VosGEs: Schiessrothried, Werner, 1959 (MSC, 2 collections).

SPAIN: Ja&n: Sierra de Cazola, Degelius, 1952 (S).

PORTUGAL: Berra ALTA: Serra da Estrela, Tavares, 1946 (S).

CANARY ISLANDS: Palma, Pitard, 1883, 1886, 1906 (DUKE, 3 collections) ;
the same, Bornmuller (W).

AZORES: Pico, Persson, 1937 (UPS, 2 collections).

KENYA: Mt. Aberdare, Fries & Fries, 1922 (UPS).

TANZANIA: Kilimanjaro, above Marangu, 3,225 m., Hedberg 1374e (UPS).

4. Platismatia herrei (Imsh.) Culb. & Culb., comb. nov. PLATE 22
Cetraria herrei Imsh., Bryologist 57:6. 1954. Based on C. glauca b. stenophylla.
Cetraria glauca b. stenophylla Tuck., Synops. No. Amer. Lich. 1:36. 1882.

Type: U.S.A., Oregon, Hall, 1871 (FH, lectotype).
Cetraria tuckermanii Herre, Proc. Washington Acad. Sci. 7:340. 1906. (Not
Oakes in Tuck., 1843). Based on C. glauca b. stenophylla.

Thallus medium, 9-12 cm. broad; lobes linear, narrow, 0.5-4 mm.
broad, sometimes much broader (up to 8 mm.) near the base. Upper
surface whitish or pale, smooth to very minutely pitted or wrinkled,
not pseudocyphellate or sorediate, isidiate, the isidia simple to branched
and coralloid, densely fringing the margins, rarely on the upper sur-
face. Lower surface white or pale with conspicuous, discrete, well de-
limited, jet-black or dark brown zones, especially in the older parts;
rhizines black, rare, a few at the places of attachment only. Upper
cortex 21-26 thick, medulla 91-177, thick, the hyphae oriented parallel
to the axis of the lobe, IKI+ lavender; lower cortex 15-26(-31) x thick,
prosoplectenchymatous. Apothecia not common, 3-8 mm. broad, ter-

538 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

minal, imperforate; hymenium 47-65 thick; subhymenium 13-184
thick. Spores 8, ellipsoid or spheroid, 4-5 & 5-8u. Pyenidia few, black,
minute, marginal; conidia 1 5y, rod-shaped.

MEDULLARY REACTIONS.—K—, C—, KC—, P—, UV-.

ConstiTUENTs.—Caperatic acid and atranorin.

DistripuTion (Map, Figure 24) —Extreme western North America
from southern Alaska south to central California; on the bark of trees.

Platismatia herrei is one of the most beautiful and intricately formed
species in the genus. It can be distinguished from all other members of
the genus by the delicate but conspicuous isidia fringing the edges of
the linear lobes. In some specimens there are fruticose coralloid struc-
tures on the margin identical to (but invariably smaller than) those
seen in some forms of P. glauca. Occasional specimens of P. herrei,
indeed, show the typically narrow, linear lobes arising from much
broader basal ones making one wonder if the plants called P. herrea
might represent only a localized western race of P. glauca. But although
some specimens may superficially give this impression, the distinctively
narrow lobes, the characteristically delicate marginal corolloid isidia,
and the black mottling in conspicuous discrete patches of the under-
surface are so different from the traits of the typical race of Platismatia
glauca inhabiting the Pacific Northwest that the distinctness of the
species is beyond doubt.

Platismatia herrei is an excellent example of a North American endem-
ic species with an extreme Pacific coastal distribution. Its range is
almost identical in extent to the coastal portion of the distribution of
western hemlock, Tsuga heterophylla (Raf.) Sarg.

CHEMICAL ANALYSIS.—Five samples of this species were tested. The
benzene extracts showed atranorin in the GE and GAo-T solutions and
by chromatography. No fatty acids were observed in these extracts,
but the subsequent acetone extracts showed caperatic acid in the GE,
GAW, and GAo-T solutions. Tests for atranorin in the GAo-T solution
showed two types of yellow crystals—one typical of atranorin and the
other smaller and more sharply curved.

ADDITIONAL SPECIMENS EXAMINED:

U.S.A.: Auaska: Hecla Island, Foster, 1913 (COLO). Prince William Sound,
Knight Island, Thum Bay, Eyerdam 305 (WIS). Washington Bay, Kuiu Island,
Eyerdam 428 (COLO), Kodiak Group, Raspberry Island, Port Vita, Eyerdam 390
(WIS).

WasHINGTON: Clallam Co.: Swan Bay, Ozette Lake, Brown & Muenscher 143
(UC, US); West Fork, Twin Rivers, Eyerdam 3162 (COLO); Sequin, Grant 41
(COLO-ABS) ; Olympic Mountains, Grant 3001 (COLO-ABS). Jefferson Co.: Port
Discovery, Brown & Muenscher 142 (UC, US). Pierce Co.: Mt. Rainier National
Park, Van Trump Trail, Howard 4202 (COLO, WTU); the same, White River,
Imshaug 414, 434 (US); the same, Tahoma Creek, Jmshaug 1741 (US); the same,
Eagle Peak Trail, Imshaug 1563 (US). Snohomish Co.: Mt. Pilchuck, Larrison,

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 539

1947 (WIS); Marysville, Grant, s.d. (COLO). Whatcom Co.: Mt. Baker, 2,000 ft.,
Herre 3234 (COLO-ABS).

OreEGon: Locality unknown, Hall, 1871 (FH). Josephine Co.: Between Cave
Junction and O’Brien, Weber 8044 (COLO). Lane Co.: Florence, Marr, 1941
(COLO); Eugene, collector unknown, 1936 (COLO).

Cauirornia: Locality unknown, Bolander 28 (FH). Humboldt Co.: Eureka,
Howe, Lich. Boreali-Amer. 143 (WIS), the same, Howe, as Cummings et al., Decades
of North Amer. Lich. 213 (DUKE, MSC, US, WIS); Pepperwood, Tavares 361 &
Seear (UC). Jackson Co.: Highway 227, Tavares 111d & Chisaki (UC). Marin
Co.: Mt. Tamalpais, Schneider, 1903 (UC); Tomales Bay, Mann & Bolander 31
(UC); Inverness, Tavares 564 (UC). Mendocino Co.: South fork of Eel River
below Elder Creek, 1,000 ft., Tavares & Sharsmith, 1961 (UC). Monterey Co.:
Locality unknown, Booth, 1878 (FH). Santa Cruz Co.: Boulder Creek, 700 ft.,
Hasse 3051 (UC). Sonoma Co.: Gualala, McMillan 110 (UC); 24 mi. south of
Gualala, Seear & Tavares 318¢ (UC).

CANADA: Britisu Cotumsia: Islands, Gulf of Georgia, Hill (COLO-ABS) ;
Mayan Island, Hill, 1904 (COLO-ABS) ; Queen Charlotte Islands, Moresby Island,
head of Bigsley Inlet, Persson 14 (COLO); Thurlow Island, Dawson, 1885 (H);
Victoria, Mt. Douglas, Kverno $1987 (COLO, US); Vancouver Island, Lyall, 1858-
1859 (US, WIS), Macoun 16 (US); the same, Port Renfrew, Herre, s.d. (UC); the
same, Nanaimo, Macoun 97 (UC, MSC, US, WIS), Krajina et al. 3729 (DUKE).

5. Platismatia interrupta Culb. & Culb., sp. nov. PLATE 22

Thallus parvus vel mediocris, 6-12 cm. latus; laciniae 0.3-2.0 cm.
latae. Superficies superior grisea (sordide citrella apud exemplaria
vetera herbarii) vel marginibus pallide fusca, levis vel reticulis inter-
ruptis indistincte rugosa, pseudocyphellata, pseudocyphellis conspi-
cuis, rotundis, elongatis vel confluentibus, granulose-sorediatis vel
subisidiatis. Superficies inferior picea, marginibus castanea vel albida,
non punctata; rhizinae paucae, nigrae. Cortex superior 18—26y crassus,
prosoplectenchymatosus; medulla 65-104 crassa, IKI— vel IKI+
dilute violacea; cortex inferior 13-21, crassus. Apothecia et pycnidia
ignota.

Thallus small to medium, 6-12 em. broad; lobes 0.3-2.0 cm. broad.
Upper surface gray (uniformly light in old herbarium specimens) or
tinged with light brown on the margins, smooth to very obscurely
wrinkled with a discontinuous reticulation, pseudocyphellate, the
pores conspicuous, round to irregular, elongate or confluent, becoming
coarsely sorediate or isidioid in some specimens. Lower surface jet-
black at the center, a marginal zone up to 2 cm. broad chestnut brown
or conspicuously white (or light brown in old specimens), not punctate;
rhizines very few, black. Upper cortex 18-26u thick, prosoplectenchy -
matous, with minute brownish crystals, IKI— or IKI+ very pale
lavender; medulla white, 65-104u thick; lower cortex prosoplecten-
chymatous, 13-21, thick, crystals absent or, if present, only a few in
the light-colored zone. Apothecia and pycnidia unknown.

Type.— Mount Norikura, Shinano Province, Japan, Asahina, 1952
(TNS, holotype).

540 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

MEDULLARY REACTIONS.—K—, C—, KC—, PD—.

ConstirveNntTs.—Caperatic acid, atranorin, and an unidentified
yellow pigment.

Distripution (Map, Figure 26) Sakhalin Island and Japan, espe-
cially in the mountains; very common on tree trunks and over boulders.

Platismatia interrupta is very common in Japan and is, in fact, one
of the most abundant of all the macrolichens there. It is obviously the
Far Eastern equivalent of P. glauca, a species that is exceedingly com-
mon in large areas of the Northern Hemisphere but which is apparen tly
absent from the extreme Orient altogether.

Platismatia interrupta differs from P. glauca in having pseudocy-
phellae on the upper surface of the lobes and an indistinct and discon-
tinuous reticulation of the upper surface that reminds one of a fur-
ther, almost complete breakdown of the reticulate ornamentation of
the upper surface of P. erosa. In addition, the upper surface of P.
interrupta is IKI—, or nearly so, while it is blue in P. glauca.

The only Japanese species for which the common Platismatia vnter-
rupta might be mistaken is the rare P. erosa. The upper surface of P.
interrupta is not pseudocyphellate in the same manner as P. erosa, and
the habit of the reticulation is different. The pseudocyphellae in P.
interrupta are large, conspicuous, and generally distributed, but they
are small, inconspicuous, and confined to the crests of the ridges in
P. erosa.

Some especially isidiate individuals of P. interrupta may resemble P.
norvegica. Platismatia norvegica has much more prominent reticulate
ridging and lacks the conspicuous pseudocyphellae characteristic of P.
interrupta.

One specimen with pseudocyphellae typical of P. interrupta but with
essentially strap-shaped lobes with marginal isidia was seen from
Sakhalin Island (Aikawa, Asahina, 1932, TNS). With the exception of
the pseudocyphellae, its habit much resembled that of the Pacific North
American P. herrei. More material from the North Asiatic Pacific will
be required to determine whether this combination of characters 1s
typical of another distinct, major population there, but for the present
the specimen is referred to P. interrupta. It should be pointed out,
however, that entirely typical specimens of P. interrupta are known
from Sakhalin Island as well.

CHEMICAL ANALYSIS.—Most tests made on this species were per-
formed by extraction of the sample in acetone without prior extraction
in benzene. All samples contained atranorin and caperatic acid, which
was observed in both crystal forms in the GE and GAW solutions.
Fourteen of the 20 samples tested also showed a yellow pigment re-
crystallizing in the GE solution. This substance seems to occur in P.
erosa, P. formosana, and P. regenerans as well. Tt may be a constant

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 54]

component, but samples which did not give a positive test for the pig-
ment were not retested.

The GAo-T solution gave yellow crystals typical of the atranorin
condensation product as well as smaller finer yellow crystals (Plate 2,
Figure 9). In addition, the GAo-T solution sometimes showed clusters
of very fine colorless needles (Plate 6, Figure 36) which were not identi-
fied but which may be due to the substances producing the unidentified
yellow crystals observed in the GE solution.

NoOMENCLATURE.—The many references to ‘“Cetraria braunsiana
(Mill. Arg.) Zahlbr.” in the Japanese botanical literature all relate to
the present species, the result of a consistent misapplication of the
name of a different plant altogether.

ADDITIONAL SPECIMENS EXAMINED:

U.S.S.R.: SaxHaLin: Ho-e, Fujikawa, 1932 (TNS); Kashiho, Ishiyama, 1932
(TNS).

JAPAN: Hoxkarpo: Prov. Hidaka: Mt. Apai, Ikuse, 1956 (TNS). Prov. Ishi-
kari: Horonaizawa, Asahina, 1935 (TNS); Hot Spring Matsuyama, Asahina, 1935
(TNS); Mt. Tomuraushi, Fujikawa, 1935 (TNS). Prov. Kushiro: Lake Masyu,
Togashi, 1953 (TNS).

Honsuu: Prov. Ettyu: Mt. Tate-yama, Waterfall Syo-myo, Nishijima, 1935
(TNS); Makawa, Asahina, s.d. (TNS). Prov. Izu: Mt. Amagi, Asahina, 1927
(TNS). Prov. Kai: Mt. Fuji, Yoshida-guichi, 1-gome, Togashi (TNS); the same,
Hisauchi, 1928 (TNS); Mt. Kinpu-san, Kurokawa 521207 (TNS); Mt. Mizugaki,
Kurokawa 521133 (TNS); Mt.Ko-fuji, Asahina, 1935 (TNS). Prov. Musashi: Pass
Mikuni, Kurokawa 520764 (TNS). Prov. Mutsu: Mt. Osoresan, Kurokawa 550447
(TNS); Montagne d’Aomori, Faurie, 1897 (PC). Prov. Shinano: Mt. Yatsu-ga-
take, Kurokawa 520674 (TNS), Kurokawa 58240 (MSC); the same, Takami-ishi,
Kurokawa 58300 (TNS); the same, Mt. Iwo-dake, Kurokawa 58436 (TNS); Yuma-
tazawa, Kurokawa 51672 (TNS); Ikenotaira, Daimon Pass, Asahina et al., 1959
(TNS) ; Heights Tadeshina, Kurokawa 58323 (TNS); Mt. Norikura, Asahina, 1952
(TNS, 2 collections); Mt. Daitenjo, Kurekawa 520518 (TNS); Mt. Gaki-dake,
Kurokawa 520598, 520599, 520642 (TNS); Mt. Tsupakura, Kurokawa 520550
(TNS); Mt. Kimpu, Asahina et al., 1960 (TNS); Suwa-gun, Koto-mura, Suguri-
daira, Kurokawa 58271 (TNS). Prov. Shimotsuke: Nikko, Karikcmi-ko, Culberson
& Culberson 11052, 10629 (DUKE), Nuno, 1961 (TNS); Nikko, Yumoto, Nuno,
1961 (TNS); Nikko, Yunc-ko, collector inknown (TNS, 2 collections). Prov.
Suruga: Mt. Fuji, Asahina 72 (UPS); Mt. Fuji, between 5- and 3-gome, Culberson
& Culberson 10655, 10662, 10669, 10757, 10806 (DUKE); Mt. Fuji, Subashiri-
guchi, Asahina, 1924 (TNS, 2 collections).

SHikoKu: Prov. Awa: Mt. Kenzan, Fujikawa, 1934 (TNS), Togashi, 1956
(TNS). Prov. Iyo: Mt. Ishizuchi, Fujikawa, 1933 (TNS).

6. Platismatia lacunosa (Ach.) Culb. & Culb., comb. nov. PLATE 23
Cetraria lacunosa Ach., Method. Lich. 295. 1803. Type: North America, Herb.
Swartzii (S, neotype).
C. lacunosa f. cavernosa (Menz.) Merr., Bryologist 13:26. 1910.
Lichen cavernosus Menz. in Ach., Lichenogr. Univ. 508. 1810 (synonym).
Platysma lacunosum (Ach.) Nyl., Bot. Not. 1855:137. 1855 (1856).

277-920 O- 68-7

542 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

Thallus small to large, 5-16 cm. broad; lobes 0.6-1.5 cm. broad.
Upper surface gray (uniformly brown or tan in old herbarium speci-
mens), the margins conspicuously blackening, prominently ridged by
strong reticulations rising at right angles to the surface, not pseudo-
cyphellate. Lower surface black at the center, chestnut brown at the
margins, somewhat reticulately wrinkled, not punctate; rhizines few,
black. Upper cortex 19-25(-29) thick ; medulla 90-280u thick (meas-
urements taken between the major wrinkles), IKI —; lower cortex
13-16 thick. Apothecia uncommon, submarginal, 0.4—2.0 cm. in diam-
eter, expanded and irregular at maturity, apparently not perforate;
hymenium 62-1244; hypothecium and proper exciple 22—50u thick, the
limit between the two zones visible in some preparations by a some-
what differential coloration with IKI; algae in groups but usually not
in a continuous layer under the proper exciple. Spores 8, ellipsoid to
ovoid, 7-10 3-4.5u. Pycnidia marginal to submarginal or even super-
ficial (on the crests of the reticulations), the ostiole round to irregular
at maturity; conidia 1x 5-7y, rod-shaped.

MpULLARY REACTIONS. —K —, C—, KC—, PD+ orange-red.

ConstTituENTS.—Fumarprotocetraric acid, caperatic acid (at least
in some specimens), and atranorin.

DistrisuTtion (Map, Figure 25) —The extreme west coast of North
America and the coastal islands from the Aleutians and southern
Alaska south to northern California; on the bark of trees (alder, hem-
lock, spruce) and on rock.

Although Platismatia lacunosa superficially somewhat resembles P.
formosana and P. erosa, it can be distinguished from these and from all
other species in the genus by its positive, brick-red medullary reaction
with p-phenylenediamine due to the presence of fumarprotocetraric
acid. It should not be assumed, however, that P. /acunosa is a mere
chemical “strain” of another species; its morphological traits are quite
distinct. The reticulate ridging of the upper surface that is so distinc-
tive results from U-shaped depressions between ridges rather than the
flat-bottomed spaces between thallus ridges as in the other species.

CHEMICAL ANALYSIS._-The samples were extracted with benzene at
room temperature to remove atranorin, which was demonstrated in the
GE and GAo-T solutions. The warm acetone extract of all but one of
the eight samples tested contained caperatic acid (GE, GAW, and
GAo-T). When chromatographed in pyridine-ethyl acetate-water, the
acetone extracts showed fumarprotocetraric acid. ‘Traces of other sub-
stances detected on these chromatograms could not be identified. One
very faint spot with an Ry resembling that of protocetraric acid was
found not to be this substance on the basis of a faint yellow fluores-
cence in UV light as compared to the characteristic quenching pro-
duced by protocetraric acid under the same conditions.

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 543

NoMENCLATURE.—There is a problem in the typification of this
species. The Acharian Herbarium in Helsinki contains no specimens
labeled Cetraria lacunosa. All the specimens which Acharius may
have had of it, including the holotype, have apparently been lost.
In Stockholm, however, there is a specimen labeled “ Cetraria lacunosa
Ach.” and, written just below this but in a different hand: ‘‘Acharius
scripsit” and “Herb. Swartzii.”” The exact history of this specimen
is unknown, but there is no reason to doubt the indication that
Acharius annotated it and that it was part of the herbarium of his
colleague Swartz. The specimen is undated, and we do not know
whether Acharius saw it before the publication of Cetraria lacunosa
in 1803, although it is most probable that he did and that his specimen
should, therefore, be considered the lectotype. In the absence of the
critical historical information, however, it cannot be so designated
and must be considered a neotype instead. The proper designation
for type specimens selected under the conditions met with here—a
common predicament in typifying many of the old names of lichens—
has been discussed in detail (W. L. Culberson, 1963).

ADDITIONAL SPECIMENS EXAMINED:

U.S.A.: Auaska: Unalaska, Setchell et al., 1899 (UC), Eyerdam, 1932 (UC),
Hultén 5277, 5278, 5279 (UPS) ; Prince William Sound, Thum Bay, Knight Island,
Eyerdam 238, 252, 264 (UC); the same, Hinchinbrook Island, Norberg 328 (WIS);
Port Vita, Raspberry Island, Kodiak Group, Eyerdam 365 (WIS); Revillagigedo
Island, Ketchikan, Eyerdam 262 (UC); Annette Island, Eyerdam 497 (WIS);
Admiralty Island, Windfall Harbor, Herre 251; Orca, Setchell, 1899 (UC); Attu
Island, Engineers Canyon, Hardy 30 (S, UPS); Adak Island, Chapel Cove, Hultén
5875 (UPS); Atka Island, Eyerdam, 1932 (S), Hultén 5790 (UPS), Kobayashi, 1931
(TNS, 6 collections); Ketchikan, Deer Mountain, 1,500 ft., Baker, 1956 (COLO),
Washington Bay, Kuiu Island, Eyerdam 428 (COLO), Eyerdam 414, 427, 509, 825
(WIS).

WASHINGTON: Grays Harbor Co.: Montesano, Heller 4310 (US). Jefferson Co.:
Hoh River Ranger Station, Becking 530656 (COLO); mouth of Hoh River, Brown
& Muenscher 154 (UC, US).

OrEGoN: Locality unknown, Hall, 1876 (US, 2 collections). Lincoln Co.: Yagu-
ina Bay, Hall, 1871 (FH); Waldport, Howard 4628 (WIS, WTU). Marion Co.:
Silverton, Powers Creek, Foster 1200 (US).

CALIFORNIA: Humboldt Co.: Lawrence Creek, Park 3512 (UC).

CANADA: British Cotumsra: Queen Charlotte Islands, Moresby Island, 650
m., Persson 1957 (COLO); Vancouver Island, Comox, Macoun 28 (US).

7. Platismatia norvegica (Lynge) Culb. & Culb., comb. nov. PLATE 23
Cetraria lacunosa f. norvegica Lynge, Videnskaps; Skrift., Mat.-Naturv.-Kl.
[Kristiania] 1921(7):196. 1921. Type: Norway, Hordaland, Store Kalsg,
Lynge, 1920 (O, lectotype).
C. norvegica (Lynge) Du Rietz, Bot. Not. 1924:59. 1924.
C. norvegica f. sorediata Degel., Acta Phytogeogr. Suec. 7:52. 1935. Type:
Norway, Monte Seljudsen prope Raegefjord, Degelius, 1932 (not seen).

544 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

C. tuckermanii var. sorediata (Degel.) Riis., Kuopion Luonnon Ystévain
Yhdistyksen Julkaisuja, Sarja B. 2(6):46. 1952.

Thallus medium to large, 7-23 cm. broad; lobes 0.5-2.5 cm. broad.
Upper surface gray, yellowish-gray or (especially in American speci-
mens) distinctly yellowish, ridged, the sharp, low ridges at right angles
to the surface of the thallus, pseudocyphellate, the pores generally
confined to the crests of the ridges, minute, often isidiate and indis-
tinct; isidia usually abundant along the margins of the ridges, gener-
ally associated with the pseudocyphellae. Lower surface jet-black, a
marginal zone up to 1 cm. wide brown or colored like the upper surface,
not punctate; rhizines black, few, usually simple. Upper cortex 19-31
thick; medulla white, 31-2174 thick, much thicker in the ridges, IKI +
pale lavender or (apparently) IKI—; lower cortex 16-31, thick.
Apothecia rare (unknown in America), irregular, perforate, 0.7-1.5
em. broad; hymenium 51-93y; thick subhymenial layers measuring
together 43-78, thick, immediately underlain by a dense zone of algae.
Spores 8, ellipsoid, 6-8.5 x 3-44. Pycnidia immersed, marginal; conidia
not seen.

MEDULLARY REACTIONS.—K—, C—, KC—, PD—.

ConsTITUENTS.—Caperatic acid and atranorin.

DistriputTion (Map, Figure 27).—Newfoundland and Miquelon
Island, Canada, the western coast of North America from southern
Alaska to Oregon; Norway, Sweden and southern Finland, Scotland;
on rocks and the trunks of trees.

The reticulate and usually isidiate thallus of Platismatia norvegica is
so distinctive that the species is easy to recognize. In Europe it always
seems to have an adequate development of isidia, immediately distin-
guishing it from P. glauca. In western North America, however, the
extent of isidia production is extremely variable, and many specimens
from densely isidiate to essentially nonisidiate can be seen in herbaria,
but these are marked by the same sharp, low, reticulate ridges that the
isidiate plants have. This reticulation and the distinct pseudocyphellae
usually associated with the ridges make these nonisidiate specimens
appear quite different from P. glauca. Platismatia norvegica and P.
glauca apparently often grow together in the Pacific Northwest as
witnessed by mixtures of the species in herbarium packets. Although
nonisidiate plants seem to be found only among the American popula-
tions of P. norvegica, the isidiate plants in the same population are
indistinguishable from the European ones.

In his original description, Lynge considered this lichen species as a
form of Cetraria (=Platismatia) lacunosa, but it differs sharply from
that species both in morphology and chemistry. Platismatia norvegica
has a pseudocyphellate, usually isidiate, minutely but sharply ridged
thallus, while P. lacunosa has a nonpseudocyphellate, nonisidiate,

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 545

deeply reticulately ridged and pitted thallus. The medulla of P. norveg-
wa is negative with p-phenylenediamine while that of P. lacunosa is
brick-red.

In an occasional rare specimen of P. norvegica there is the develop-
ment of large capitate soralia on the surface of the lobes. This modifica-
tion was named Cetraria norvegica f. sorediata by Degelius. Though
conspicuous, these soralia are so foreign to the scattered, granular type
produced normally in the genus (as in P. glauca and P. interrupta) that
they give every impression of being eruptions of the medulla through
the upper cortex as the result of some external or pathological influence
altogether. The occurrence of these peculiar soralia is so infrequent and
their structure so bizarre that they are given no taxonomic recognition
here.

Platismatia norvegica is an extremely oceanic species. It is not found
beyond the area of very wet climates devoid of temperature extremes
that characterize the region of the Pacific Northwest and the oceanic
province of northwestern Europe. The belated recognition of the
specific distinctness of this species in the well-known flora of Scandi-
navia prompted a number of reports and studies that ended with
Ahlner’s (1948) comprehensive treatment of the species in Scandinavia
with a map of all collections; the reader is referred to this publication
for details of the northern European range. In that paper and in
Hakulinen’s (1963) lichen flora of Finland, the species is also reported
from extreme southern Finland. No Finnish species were seen in the
present study, but we have no reason to doubt the correctness of those
records.

A map of the worldwide distribution of Cetraria norvegica was also
published by Ahlner (1948). The map in the present treatment (Figure
27) does not differ qualitatively from Ahlner’s although we have added
more American localities. The present study and the data collected by
Ahlner (1948) also show that although the species grows on both trees
and on rocks, it is more common on bark in the southern part of its
range and on rocks in the northern part even though both substrate
types are generally present throughout the range.

CHEMICAL ANALYSIS.—The seven samples tested contained caperatic
acid and atranorin. No protolichesterinic acid was detected in the
benzene extract after treatment with the GE or GAW solutions.
Atranorin was proved in this extract in the GE and GAo-T solutions.
The subsequent acetone extract contained more atranorin in addition
to caperatic acid as found by treating the extract with the GE, GAW,
and GAo-T solutions. No other fatty acids were detected. No other
substances reacting with PD were found in the acetone extract after
chromatography in pyridine-ethy] acetate-water (1: 2:1). In the GAo-T
solution, smaller crystals similar to the o-toluidine derivative of atra-

546 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

norin occurred in abundance accompanied by those of the typical
atranorin derivative.

NoMENCLATURE.—Lynge listed about 30 specimens with the original
diagnosis of Cetraria lacunosa f. norvegica. No type specimen was
designated, but when Du Rietz elevated the forma to specific rank, he
indicated as the type a Norwegian specimen collected by himself but
not among those originally cited by Lynge. Since a taxon must be
typified from the original material if possible, we have picked Lynge’s
collection from Store Kalsg in Hordaland as the lectotype. It is a large,
well developed example of the species.

REPRESENTATIVE SPECIMENS EXAMINED:

CANADA: NEWFOUNDLAND: Locality unknown, Despreaux, 1828 (UPS); King
Cove, Dammon 1 (US); St. John, Fries, 1871 (8S, 2 collections).

British CoLumMBia: Queen Charlotte Islands, Graham Island, Persson 26, 54
(COLO); Vancouver Island, Nanaimo River Valley, Szczawinski 558 (US); the
same, Port Renfrew, Schneider, s.d. (UC); the same, Comox, Macoun 122 (WIS).

MIQUELON ISLAND: Locality unknown, Delamare. 1886 (DUKE, POC),
Delamare, as Arn., Lich. Exs. 1366 (PC).

U.S.A.: Ataska: Kodiak Island, Kodiak, Hultén 5053 (UPS), Eyerdam 79
(WIS); Admiralty Island, Windfall Harbor, Herre 251 (UC); Glacier Bay, Beards-
ley Island, Stephens 242 (UC); Prince William Sound, Knight Island, Kyerdam
252, 254 (UC), Norberg, 1940 (8), College Fjord, Viereck 2419-B, Bainbridge
Glacier, Viereck 1992-A (COLO); Kenai Peninsula, Seward, Hultén 7887 (UPS);
Yukutat, Schmuck 19 (WIS); Juneau, Hultén 8558 (UPS).

WASHINGTON: Grays Harbor Co.: Pacific Beach, Foster 1359 (COLO-ABS).
Jefferson Co.: Mouth of the Hoh River, Brown & Muenscher 154 (UC, WTU).
Pacific Co.: Willapa Bay, Toka Point, Foster (COLO-ABS). Pierce Co.: Mt.
Rainier National Park, Pechanec (COLO). Whatcom Co.: Mt. Baker, Howard
1180a (LD).

OrEGON: Marion Co.: Silverton, Powers Creek, Foster 1200 (US).

SWEDEN: JAMTLAND: Are, Ahiner, 1951 (DUKE), Hasselrot, 1937 (LD),
Hdkansson, 1949 (LD), Santesson 48443, 48443b (UPS); Frostviken, Stenholm,
1918 (GB).

DaLARNE: Malung, Hasselrot, 1934 (LD, 4 ecolleetions), Hasselrot, 1937 (8),
Hasselrot, as Magn., Lich. Sel. Scand. Exs. 184 (GB, LD, 8); Lima, Hasselrot,
1934 (LD).

VARMLAND: Munkfors, Sundell, 1952 (LD); Dalby, Persson, 1948 (8).

VASTERGOTLAND: Bredareds, Hasselrot, 1961 (S).

NORWAY: N6GrpiANpD: Elsfjord, Ahlner, 1939 (S); Nord-Rana, Ahlner, 1937
(S),
Norp-TRONDELAG: Egge, Ahiner, 1934 (8); Grong, Osterlind, 1938 (LD); Vaer-
dal, Osterlind, 1932 (LD); So6rli, Ahiner, 1938 (S); Malm, Ahiner, 1938 (8, 2 collec-
tions).

Horpatanp: Herdla, Santesson, 1937 (US); Voss, Ahlner, 1937 (S).

RoGaLanpb: Hjelmeland, Degelius, 1947 (LD); Solendal, Hasselrot, 1939 (S);
Rege Fjord, Hasselrot, 1939 (US).

Soan oG Fyorpane: Hid, Ahlner, 1937 (S).

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 547

SCOTLAND: Loch Rannoch, ex herb. Stirton (BM, 2 collections); Killin, Craig
Chaillech, M’ Andrew, 1885 (BM).

8. Platismatia regenerans Culb. & Culb., sp. nov. PLATE 24

Thallus parvus vel mediocris, 4-8 cm. latus; laciniae 0.3-1.2 cm.
latae, marginibus ascendentes. Superficies superior albida vel pallide
cinerea centro et marginibus nigra vel sepiacea, levis, pseudocyphel-
lata, pseudocyphellis parvissimis et inaequalibus, corporibus parvis,
regenerantibus(?), isidioideis vel complanatis in partibus vetustioribus
praedita. Superficies inferior picea, marginibus castanea, vulgo reticulato-
rugosa, rhizinata, fere punctata. Cortex superior prosoplectenchy-
matosus, 21-31, crassus; medulla (104—) 120-180(—208) » crassa, IKI— ;
cortex inferior 18-26, crassus. Pycnidia et apothecia ignota.

Thallus small to medium, 4-8 cm. broad; lobes 0.3-1.2 cm. broad,
the margins ascending. Upper surface whitish or pale gray, the center
and margins black or dark brown, smooth, pseudocyphellate, the pores
very small and irregular, the older parts of the surface with scattered
regenerative(?) bodies, isidioid or flattened and lobule-like, mostly less
than 0.5 mm. broad. Lower surface black, the margins dark brown,
commonly reticulately rugose, rhizinate, usually punctate. Upper
cortex prosoplectenchymatous, 21-3ly thick; medulla white (104)
120-180(—208) thick, IKI—; lower cortex 18-26 thick. Pycnidia
and apothecia unknown.

Typr.—Tourist Trail, just above Kambarabga radio tower, Kina-
balu National Park, Sabah (Borneo), 2,000 m. alt., Hale 28207 (US,
holotype).

MEDULLARY REACTIONS.—K—, C—, KC—, PD—-.

CONSTITUENTS.—Caperatic acid and atranorin.

DistriputTIon (Map, Figure 29).—The high mountains of Sabah
(Borneo).

The only species with which Platismatia regenerans might be con-
fused is P. erosa. It differs from P. erosa in that it lacks true isidia
although it may have a few flattened, apparently regenerative lobulae
on the older parts of the thallus that might be mistaken for isidia.

CHEMICAL ANALYSIS.—The residue from the benzene extract of four
samples tested showed atranorin in the GE and GAo-T solutions.
These residues did not give positive tests for protolichesterinic acid in
the GE or GAW solutions. The residues from subsequent acetone ex-
tractions contained more atranorin and caperatic acid. Both crystal
forms of caperatic acid were observed in the GE solution, and caperatic
acid was also detected in the GAW and GAo-T solutions. This species
may also contain the unidentified substance (or substances) observed
in P. erosa, P. formosana, and P. interrupta. One specimen showed
yellow crystals in the GE solution, and another gave clusters of long
fine needles in the GAo-T solution.

548 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

SPECIMENS EXAMINED:

SABAH (BORNEO): Kinabalu National Park, Tourist Trail, just above Kam-
barabga radio tower, 2,000 m., Hale 28207, 28932 (US); the same, woods below
second radio tower (above Kambarabga), 2,200 m., Hale 28642 (US); the same,
open ridge between East and West Mesilau Rivers, 1,800 m., Hale 29084 (US).

9. Platismatia stenophylla (Tuck.) Culb. & Culb., comb. nov. PLATE 24
Cetraria lacunosa b. stenophylla Tuck., Synops. No. Amer. Lich. 1:35. 1882.
Type: U.S.A., California, Bolander (FH, holotype).
Cetraria stenophylla (Tuck.) Merr., Bryologist 13:27. 1910.

Thallus medium, 5-10 cm. broad; lobes linear, narrow, (0.5) 1-4 mm.
broad, often becoming trough-shaped as the margins roll inwards, the
margins occasionally with small lobulae oriented at right angles to the
axis of the lobes, often attached to the substrate only at the base.
Upper surface whitish, tan, pale brown or occasionally somewhat
darkening, smooth, not pseudocyphellate, sorediate, or isidiate. Lower
surface whitish, tan, or brown, not punctate, minutely reticulate-
rugose, sometimes appearing veined with the crests of the ridges
darkening; rhizines rare, a few at the place of attachment only. Upper
cortex 16-234 thick; medulla 86-150(-208) thick, IKI+ lavender;
lower cortex (13-)16-2ly thick. Apothecia common, especially in the
southern part of the range, terminal, 0.4—1 cm. broad, perforate or not;
hymenium 39-52u thick, subhymenium 18-34y, thick, often underlain
with algae; spores 8, small, ellipsoid or spheroid 4-55-84. Pycnidia
few, black, marginal; conidia 1 5-8u, rod-shaped.

MEDULLARY REACTIONS.—K—, C—, KC—, PD—.

ConsTITUENTS.—Caperatic acid and atranorin.

DistriputTion (Map, Figure 23).—Western North America from
Vancouver Island south to west-central California; on trees, usually
near the shore.

Platismatia stenophylla is a well defined species with narrow, linear
lobes, a trait shared in this genus by only P. herrei which is easily
distinguished from the present species by its abundant marginal isidia.
In the northern part of the range, P. stenophylla has more darkly
colored thalli with strongly incurved margins and a paucity of apothe-
cia. The southern plants are more robust, stouter, and fruit more
freely. In spite of these tendencies, specimens cannot be assigned to
their geographical origin with high accuracy upon habit alone, and
intermediate forms are common.

Platismatia stenophylla has the most narrowly restricted geographic
range of any of the American species of the genus.

CHEMICAL ANALYSIS.—The five samples tested of this species showed
atranorin in the GE and GAo-T solutions and by chromatography.
The benzene extract seemed to contain only atranorin while the sub-
sequent acetone extract consisted of this depside mixed with caperatic

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 549

acid, which was identified by the crystals in the GE and GAW solu-
tions. No other fatty acids were noted.

ADDITIONAL SPECIMENS EXAMINED:

CANADA: British CoLtumBia: Vancouver Island, Nanaimo Lakes, Krajina et
al. 8-15 (DUKE).

U.S.A.: Wasuineton: Lewis Co.: Deschutes Park, Snoqualmie National Forest,
Buchta & Hedglin, 1958 (WTU). Pierce Co.: Mt. Rainier, Flett, 1919 (US). Spo-
kane Co.: Spangle, Cooke 23198 (WIS). Whitman Co.: Kamiak Butte, /mshaug
16462 (MSC).

OREGON: Benton Co.: Corvallis, Becking 5304666 (COLO).

Ipano: Latah Co.: Palouse Range Ridge, Imshaug 16413 (MSC).

CauirorniA: Humboldt Co.: Locality unknown, Bolander (FH, US). Santa
Cruz Co.: Castle Rock, Herre 1025, 1942, 4715 (WIS), Herre 3235 (COLO), Shus-
han 14737 (DUKE, US); Devils Canyon and Castle Rock, Herre, as Krypt. Ezs.
Vidobon. 1553 (US); Skyline Boulevard, south of Page Mill Road, Wiggins C-68
(DUKE); Ben Lomond, Thiers 9158 (WIS). San Mateo Co.: Hubbard Park,
Thiers 9232 (WIS).

10. Platismatia tuckermanii (Oakes) Culb. & Culb., comb. nov. PLATE 25
Cetraria tuckermanti Oakes in Tuck., Amer. Journ. Sci. & Arts 45:48. 1843.
Type: U.S.A., Massachusetts, Cambridge, Tuckerman, 1838 (FH, neotype).
C. lacunosa 6 atlantica Tuck., Proc. Amer. Acad. 1:208. 1847. Type: Lake
Superior, Agassiz (FH, lectotype).
C. atlantica (Tuck.) Du Rietz, Bot. Not. 1925:10. 1925.

Thallus medium, to 7-12 cm. broad, loosely attached ; lobes rounded »
0.4-2.2 cm. broad. Upper surface ashy-white or gray, irregularly or
reticulately pitted, often conspicuously so, without pseudocyphellae,
soredia, or isidia. Lower surface white or slightly tan with well defined
patches becoming black or dark brown, especially toward the center of
the thallus, minutely and reticulately pitted, not punctate; rhizines
few, black at the center, pale or tipped with white nearer the margin.
Upper cortex 18-26(-29)u thick; medulla 109—-182(-234)y thick, the
hyphae loosely interwoven, IKI —; lower cortex 16—23(-29)y thick.
Apothecia common, marginal or submarginal, 2-10 mm. broad, usu-
ally perforate; hymenium (39-)50-65(-—73)u; subhymenium 18-26
(—34)u, IKI+ lavender, underlain by at least a few algae; spores 8,
ovoid to subglobose, 4-5 6-84. Pycnidia common, blackening, mar-
ginal or submarginal, immersed, the ostiole minute; conidia rod-
shaped, ca. 1 X5—-8y.

MEDULLARY REACTIONS.—K —, C—, KC—, PD-.

ConsTITUENTS.—Caperatic acid and atranorin.

DistrisutTion (Map, Figure 31).—Southern Ontario and Quebec,
Newfoundland to the Lake States and south in the Appalachians to
Alabama and Georgia; on the bark of trees, primarily of conifers, and
occasionally on old wooden fences.

550 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

60 i) 70

25

90 85 80 75

Figure 31. Distribution of Platismatia tuckermanii (Oakes) Culb. & Culb.

Platismatia tuckermanw is quite distinct from all other species. In
the past it has been mistaken for P. lacunosa although the ranges of
the species do not overlap. The thallus of P. tuckermanii is broadly
reticulate, but it lacks the deep, sharp, thallus folds and ridges that
characterize P. lacunosa and differs chemically in lacking fumar-

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 551

protocetraric acid and therefore reacts negatively with p-phenylene-
diamine.

In herbarium collections, members of the Cetraria ciliaris Ach. and
related species have often been mistakenly identified as Platismatia
tuckermanii, doubtless because these plants regularly grow together
throughout their range. The members of the Cetraria ciliaris group all
have marginal stalked pycnidia, usually have marginal cilia, and the
most common species have aromatic constituents (alectoronic, a-col-
latolic, olivetoric, and physodic acids) which give positive reactions
with C or KC. Platismatia tuckermanii lacks all of these traits. There
are also fundamental differences in the anatomy of the upper cortex
between the species.

Platismatia tuckermanii is also sometimes mistaken for P. glauca
although almost invariably it is the more abundant of the two species
in places where they grow together. It can be differentiated at once
from esorediate and nonisidiate specimens of P. glauca by its negative
thallus reaction with IKI which in P. glauca is strongly positive, pro-
ducing a deep blue-black coloration. Although many morphologic
characters separate the two species, the IKI test is quickly used and
definitive.

Platismatia tuckermanii is broadly distributed in northeastern
temperate North America. The species belongs to the northern conifer-
dominated forests and becomes rare abruptly outside of this vegeta-
tional region.

Platismatia tuckermanii shows a strong preference for the bark of
coniferous trees. In northern Wisconsin where the species is common,
its presence at breast height in quadrats on 2,000 randomly selected
trees was analyzed (Culberson, 1955). It was found on 19 percent of
the white pines (Pinus strobus), 7 percent of the jack pines (Pinus
banksiana), 2 percent of the lindens (Tilia americana), and about 1
percent of the red pines (Pinus resinosa), hemlocks (T'suga canaden-
sis), red oaks (Quercus rubra), yellow birches (Betula lutea), and
sugar maples (Acer saccharum). It was absent from the white birches
(Betula papyrifera), red maples (Acer rubrum), beeches (Fagus grandi-
folia), and poplars (Populus spp.). Elsewhere in its range its preference
for coniferous trees is also obvious. It is also sometimes found on old
wooden fences but almost never on rocks.

CHEMICAL ANALYSIS.—Seven samples were tested and all contained
atranorin as demonstrated in the GE and in GAo-T (Plate 2, Figures 8
and 10) solutions. The benzene extract gave no crystals of protoli-
chesterinic acid in the GE or GAW solutions, but the subsequent ace-
tone extract showed caperatic acid. Crystals of caperatic acid were
observed in the GE (in both crystal forms), GAW, and GAo-T solu-

552 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM

tions. The acetone extract chromatographed in pyridine-ethyl acetate-
water showed no components reacting with PD other than atranorin.

NOMENCLATURE.—No specific collections were cited with the de-
scription of Cetraria tuckermanii Oakes and no original collections of
undisputed authenticity exist. The neotype selected here (see above) is
a specimen in Tuckerman’s herbarium (FH) bearing the name ‘‘Cetra-
ria tuckermanw”’ and collected in Massachusetts five years before
Oakes’ description was published. There is no doubt of the reference
intended in the application of Oakes’ name, and the neotype fits the
description well. It is also highly probable that Oakes even examined
this very collection when he described the new species.

In his description of Cetraria lacunosa 6 atlantica, Tuckerman cited
two specimens, one from New York and the other from Pennsylvania.
Today there are no specimens labeled with this name in his herbar-
ium. In the original description the range is given as ‘‘Lake Superior
to New England.” In Tuckerman’s herbarium there is a specimen col-
lected by Agassiz and labeled ‘Lake Superior.”” This collection, from
Louis Agassiz’s famous trip, is undoubtedly the material upon which
the statement concerning the geographic range in the original descrip-
tion referred. We designate this specimen as the lectotype of C.
lacunosa B atlantica.

Du Rietz (1925) cited specimen number 468 in Reliquiae Farlowi-
anae as the type of C. lacunosa B atlantica, but this undated New
Hampshire collection, issued simply as C. lacunosa, cannot in any way
be associated with the original diagnosis of the variety.

REPRESENTATIVE SPECIMENS EXAMINED:

CANADA. Ontario: Algoma District, Mississagi River, Cain 25867, 25870,
25872 (US), Timagami, Gull Lake, Cain 26782 (COLO, US); Emo, Fink 739 (K,
MSC, UC); Ottawa, Couet 1899 (PC).

QuEBEC: Bellechasse Co.: Ste. Camille, Masson 5007, 5008, 6182, (WIS). Terre-
bonne Co.: Pare du Mont Tremblant, Hermann 15759 (US). Woburn Co.: Fron-
tenac, Masson 6031 (WIS).

U.S.A.: Maine: Hancock Co.: Mt. Desert Island, Tuckerman, 1871 (FH).
Lincoln Co.: Hog Island, Wiley, 1953 (WIS). Piscataquis Co.: Mt. Katahdin,
Degelius, 1989 (US). Washington Co.: North Lubee, Cummings, Dec. North Amer.
Lich, 279 (DUKE, WIS).

VERMONT: Addison Co.: Bristol, Faron, 1879 (WIS).

New Hampsnire: Belknap Co.: Gilmanton, Blake 5 (COLO), Carroll Co.:
Chocorua, Farlow, Rel. Farlowianae 468 (UC, US, WIS). Grafton Co.: North
Woodstock, Cummings, Dec. North Amer, Lich, 46 (OUKE, MSC, US, WIS).

Massacuuserts: Bristol Co.: New Bedford, Willey 109 (US). Essex Co.: Man-
chester, Seymour, 1908 (UC). Worcester Co.: Lunenburg, Sargent, IS83 (UC).

Connecricut: New London Co.: Norwich, Setchell, 1883 (UC).

New York: Essex Co.: Chilson Lake, Harrés, 1898 (COLO-ABS, US). Greene
Co.: Maplehurst, Dir, 1929 (DUKE). Suffolk Co.: Riverhead, Bredo 1097 (MSC).

New JersEY: Bergen Co.: Closter, Grout 6185 (DD UKE). Burlington Co.: Chats-

CULBERSON & CULBERSON—CETRELIA AND PLATISMATIA 553

worth, Darrow 1905 (WIS). Ocean Co.: Lakehurst, Thomson, 1940 (WIS). Sussex
Co.: Highview Mountain State Park, Hale 17291 (DUKE, US).

PENNSYLVANIA: Clinton Co,: Westport, Hale 16082 (US). Huntingdon Co.:
Penn State Nature Camp, Thomson, 1937 (WIS). Monroe Co.: Pocono Pines,
Kaiser, 1910 (DUKE). Susquehanna Co.: Starruca Station, Diz, 1940 (DUKE).
Wayne Co.: Scott Center, Diz, 1941, 1943 (DUKE, 2 collections).

Ox10: Ashtabula Co.: Orwell, Bogue 269 (DUKE). Hocking Co.: Old Man's
Cave, Hale 13328 (US).

Kentucky: Lee Co.: Beattyville, Hale 13758 (US). Menifee Co.: 10 mi. south-
west of Mariba, Hale 13423 (US). Powell Co.: Natural Bridge State Park, Braun,
1916 (DUKE). Whitley Co.: Cumberland Falls, Hale 13105 (US).

West Virginia: Cabella Co.: Milton, Hale 10779 (US). Gilmer Co.: Cedarville,
Hale 10500 (US). Hardy Co.: Mathias, Gillespie 15437 (US). Monroe Co.: Peters-
town, Hale 13064 (US). Pendleton Co.: Judy Gap, Hale 10127 (US). Pocahontas
Co.: Michael Mountain, Gray 3254, (COLO-ABS). Raleigh Co.: Clear Creek, Hale
11840 (US). Tucker Co.: Canaan Mountain, Allard 10907A (US).

Vircinia: Appomattox Co.: Appomattox Surrender Grounds, Hale 15610 (US).
Culpeper Co.: Jeffersonton, Hale 18472 (US). Giles Co.: Mountain Lake, Culberson
11251 (DUKE). Grayson Co.: Comers Rock, Hale, Lich. Amer. Exs. 60 (COLO,
DUKE, MSC, UC, WIS). Page Co.: Shenandoah National Park, Jreland 3776
(US). Smyth Co.: Whitetop Mountain, Hale 18629 (US). Wythe Co.: Wytheville,
Hale 12024 (US).

Maryann: Anne Arundel Co.: Locality unknown, Tuckerman, 1852 (FH).
Prince Georges Co.: Cedarville State Forest, Hale 14346 (US).

Norru Carouina: Avery Co.: Grandfather Mountain, Moore 1897 (DUKE).
Gaston Co.: Gastonia, Culberson 5258 (DUKE). Jackson Co.: Glenville, Culberson
5748 (DUKE). McDowell Co.: Buckereek Gap, Culberson 4873 (DUKE).Macon
Co.: Mt. Staula, Moore 1592 (DUKE). Mitchell Co.: Spruce Pine, Culberson 4847
(DUKE). Orange Co.: Hillsborough, Culberson 12069 (DUKE). Yancey Co.: Mt.
Mitchell, Culberson 4814 (DUKE).

TENNESSEE: Hamilton Co.: Lookout Mountain, Calkins 236 (UC). Johnson
Co.: Backbone Rock Recreational Area, Hale 18408 (US). Sevier Co.: Mt. Le
Conte, Degelius, 1939 (US).

Souru Carona: Greenville Co.: Greenville, Hale 7759 (US).

Geroraia: Dade Co.: Trenton, Hale 7383 (US). Murray Co.: Chatsworth, Hale
7423 (US). Towns Co.: Hiwassee, Culberson 7274 (COLO, DUKE, US). Union
Co.: Vogel State Park, Hale 7343 (US).

ALABAMA: Cleburne Co.: Cheaha Park, Hale 7113 (US). De Kalb Co.: De Soto
Park, Hale 7024 (US). Jackson Co.: Section, Hubricht B2182 (WIS). Lee Co.:
Auburn, Becking 5910004 (US).

MicHiGANn: Baraga Co.: Covington, Imshaug 20846 (MSC). Chippewa Co.: Hul-
bert, Thomson 1008 (WIS). Houghton Co.: Rice Lake, Wetmore 1315 (MSC).
Keweenaw Co.: Isle Royale, Wetmore 2522 (COLO, WIS). Marquette Co.: Cliff
Lake, Imshaug 4325 (WIS). Schoolcraft Co.: Big Spring Indian Lake, Imshaug
19993 (MSC).

Wisconsin: Adams Co.: Friendship, Culberson 3120 (WIS). Ashland Co.: Mel-
len, Lake Galilee, Culberson 3629 (WIS). Door Co.: Porte des Morts Strait, Culber-
son 4145 (WIS). Douglas Co.: Brule River Valley, Thomson, Lich. Wiscon. Erxs. 39
(COLO, DUKE, MSC, WIS). Iron Co.: 16 mi. south of Hurley, Culberson 3125
(MSC, WIS). Taylor Co.: Rib Lake, Culberson 4235 (WIS). Oneida Co.: Wood-
ruff, Thomson, Lich. Wiscon, Exs. 18 (COLO, DUKE, MSC, US, WIS). Vilas Co.:
Lac Vieux Desert, Cheney, 1893 (WIS).

277-920 O - 68 - 8

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Index

(To species and infraspecific taxa. Synonyms in italics; page numbers of principal
entries in boldface.)

Acer rubrum, 551
saccharum, 551

Asahinea chrysantha, 492, 493

Aspedelia wattii, 521

Betula lutea, 551
papyrifera, 551

Cetraria alaskana, 492
atlantica, 549
braunsiana, 493, 497, 541
braunsiana f. isidiosa, 493
chicitae, 504
chrysantha, 451
ciliaris, 470, 517, 551
collata, 451, 452, 455, 505
collata f. isidiata, 493, 495, 514
collata f. microphyllina, 511
collata f. nuda, 495, 513, 514
cucullata, 450, 451
fallax, 450
formosana, 529
formosana var. isidiata, 526, 527
glauca, 450, 451, 452, 530
glauca var. chilensis, 530
glauca b. stenophylla, 537
herret, 537
islandica, 450
japonica, 451, 452, 511
juniperina, 450
lacunosa, 450, 541, 544, 552
lacunosa 6 atlantica, 549, 552
lacunosa f. cavernosa, 541
lacunosa f. norvegica, 543, 546
lacunosa b. stenophylla, 548
nivalis, 450, 451
norvegica, 451, 543
norvegica f. sorediata, 543, 545
sanguinea, 451, 452, 521
sanguinea f. isidiata, 510
sepincola, 450
stenophylla, 548
stracheyt, 451
thomsonii, 451
tuckermanii, 537, 549
tuckermanit var. sorediata, 544

Cetrelia alaskana, 454, 459, 460, 462,
463, 492, 494, 500, 502, 508, 509,
514
braunsiana, 454, 458, 459, 462, 463,
491, 493, 504, 511, 514, 520
cetrarioides, 454, 458, 459, 462, 463,
491, 492, 498, 504, 516, 517
chicitae, 454, 458, 459, 462, 463,
491, 495, 496, 499, 500, 501, 504,
510, 514, 516
collata, 454, 458, 459, 461, 463, 488,
492, 505, 509, 514, 520
davidiana, 452, 459, 461, 463, 491,
505, 506, 507, 514, 522
delavayana, 454, 461, 463, 492, 505,
506, 508, 509, 514
isidiata, 454, 459, 462, 463, 491, 494,
510
japonica, 454, 461, 462, 463, 487,
491, 506, 511, 519, 520, 522, 524
nuda, 454, 458, 461, 463, 488, 492,
495, 496, 505, 506, 508, 513
olivetorum, 454, 462, 463, 491, 494,
498, 504, 508, 515, 520
pseudolivetorum, 454, 460, 461, 462,
463, 491, 494, 510, 512, 519, 520
sanguinea, 454, 459, 461, 490, 491,
496, 505, 506, 508, 510, 511, 512,
514, 521
sinensis, 454, 460, 463, 482, 488,
491, 506, 511, 519, 520, 523
Fagus grandifolia, 551
Imbricaria megaleia, 521
sanguinea, 521
Lichen ampullaceus, 534
cavernosus, 541
glaucus, 530, 534
fallax, 534
fraudulentus, 534
islandicus, 450
membranaceus, 534
Lobaria glauca, 530
Parmelia braunsiana, 493, 496
braunsiana f. isidivsa, 493, 496

557

5008 INDEX

Parmelia—Continued Platismatia— Continued
cetrarioides, 452, 459, 498, 502 interrupta, 454, 458, 460, 462, 463,
cetrarioides f. bisoralifera, 498 488, 497, 526, 529, 530, 533, 535,
cetrarioides var. rubescens, 515 539, 547
cryptochlorophaea, 480 lacunosa, 454, 458, 462, 463, 478,
furfuracea, 470 481, 526, 530, 532, 541, 545, 550
glauca, 5380 norvegica, 454, 460, 462, 463, 501,
megaleia, 521 525, 528, 535, 540, 543
monochorum, 498, 502 regenerans, 454, 462, 463, 488, 526,
olivaria f. subvenosa, 515 528, 529, 530, 535, 540, 547
olivetorum, 452, 459, 515 stenophylla, 454, 462, 463, 526, 532,
perlata 6 cetrarioides, 498, 502 548
perlata 8 olivaria, 515, 517 tuckermanii, 454, 462, 463, 526, 549
perlata B olivetorum, 515, 517 Platysma braunsianum, 493
perlata f. rubescens, 515, 518 braunsianum f. isidiosum, 493
pseudolivetorum, 519 collatum, 505, 514
reticulata, 517 collatum f. microphyllinum, 511
rubescens, 515 collatum f. nudum, 513
tinctorum, 468 glaucum, 530
yunnana f. subnuda, 513 lacunosum, 541
Physcia glauca, 530 megaleium, 521
Pinus banksiana, 551 sanguineum, 521
resinosa, 551 subperlatum, 521
strobus, 551 Pseudevernia furfuracea, 470
ar ss, 52 6, 530, 533, 535, sao, Pscudoparmelia aradensis, 459, 515
cyphellata, 459
542, 547 seudofallaz, 459, 502
formosana, 454, 458, 462, 463, 488, P aoe
526, 527, 529, 535, 540, 542, 547 | Quercus rubra, 551
glauca, 454, 455, 456, 460, 461, 462, | Ramalina siliquosa, 489
463, 525, 526, 530, 532, 540, 544, Thuidium cymbifolium, 514
551 Tilia americana, 551
herrei, 454, 460, 461, 462, 463, 525, | Tsuga canadensis, 551
526, 532, 537, 540, 548 heterophylla, 538

U, S. GOVERNMENT PRINTING OFFICE : 1968 O - 277-920