CM

LIBRARY OF THE

UNIVERSITY OF ILLINOIS

AT URBANA-CHAMPAIGN

BIOLOGY

0-5

MONOGRAPH OF

MARGERY C. CARLSON

FIELDIANA: BOTANY
VOLUME 29, NUMBER 4

Published by
CHICAGO NATURAL HISTORY MUSEUM

DECEMBER 6, 1957
THE LIBRARY OF TH$

JAN : V:ES

BIOLOi
«>..VtrtSITY OF ILLINOIS

MONOGRAPH OF

THE GENUS RUSSELIA

(SCROPHULARIACEAE)

MARGERY C. CARLSON

Associate, Department of Botany

Associate Professor of Biological Sciences
Northwestern University

FIELDIANA: BOTANY
VOLUME 29, NUMBER 4

Published by

CHICAGO NATURAL HISTORY MUSEUM
DECEMBER 6, 1957

THE LIBRARY OF THE

JAN ^ 1CFR

Library of Congress Catalog Card Number: 57-1 ^95 k

PRINTED IN THE UNITED STATES OF AMERICA
BY CHICAGO NATURAL HISTORY MUSEUM PRESS

0

Monograph of the Genus Russelia

History of the Genus

The genus Russelia was described by N. J. Jacquin (14) in 1760.
It was named for an English physician and naturalist, Dr. Alexander
Russell, of Aleppo, Syria. The type species, R. sarmentosa, was col-
lected in woods and dense shrubby areas near Havana, Cuba. Jac-
quin distinguished the genus from others of the Scrophulariaceae
by the four didynamous stamens, 5-parted calyx, cylindrical tubular
corolla, one-loculed capsule, and numerous seeds. He gave a more
complete description of the species later (15).

A. L. de Jussieu (16) later described the genus in more detail
and noted that the capsule has two locules rather than one.

Between 1799 and 1832, ten new species were described, and in
1838, George Don (9) studied the family, divided it into tribes,
and placed Russelia in his Tribe 1, Scrophularieae. He described
briefly nine of the eleven species then known.

G. Bentham wrote the account of the genus in A. de Candolle's
Prodromus (1), placed Russelia in the tribe Cheloneae, and listed
five species. He regarded R. alala as a doubtful species and later
(2) placed it in the genus Anetanthus of the Gesneriaceae.

Hemsley (13) studied the Scrophulariaceae and included Rus-
selia as a Mexican and Central American genus with five shrubby
species, reducing all the other then-known taxa to synonymy.

B. L. Robinson (26) studied the genus in 1900 and stated:
"Bentham's extreme reduction of species is ill-advised; at least
a dozen species may be recognized with advantage." He presented
the first key to thirteen species (three of which were described as
new) based on the characters of the stem, namely: winged, sharply
angled, 4-angled, 6-angled, or subterete. He made R. juncea synony-
mous with R. equisetiformis, R. retrorsa with R. polyedra, and R.
paniculata with R. multiflora, and he made R. verticillata a form of
R. multiflora. He did not include R. coccinea, which had been
described by Wettstein (40) in 1891. Robinson continued to de-
scribe new species (27-31) until 1909, but apparently never made
another intensive study of the genus.

231

232 FIELDIANA: BOTANY, VOLUME 29

Hallier (12) did not remove R. alata from the genus but placed
it in the tribe Gratioliae, while all other species remained in the
Cheloneae.

In 1920, F. W. Pennell (23) studied the Scrophulariaceae of
Colombia, and gave a key to the tribes and sub-tribes. He described
Russelia colombiana Penn., the only species known from Colombia,
and placed it in the tribe Antirrhinoideae, sub-tribe Russelieae,
distinguished by having the posterior lobes of the corolla external in
the bud, the corolla red, nearly regular, tubular, the short lobes
nearly equal, the stigma capitate, the capsule dehiscing to the base
septicidally and filled with slender hairs among the seeds, the seeds
reticulate, not winged, the leaves whorled in threes, the stem with
six angles, and the inflorescence in axillary cymes, two pedicels of
each being terminal to the primary peduncle.

Paul C. Standley (36) gave a key to 17 species, with short de-
scriptions of each. Again, the main distinctions were based on the
nature of the stem — whether angled or terete. He reduced several
species to synonymy as follows: syringaefolia to floribunda; juncea
to equisetiformis; paniculala to multiflora; deamii to a form of
verticillata; retrorsa to polyedra and both of these probably to terni-
folia; and furfuracea to tepicensis.

In 1945, C. L. Lundell (20) stated that the genus Russelia had
been neglected by taxonomists and that many undescribed species
had been pigeon-holed. An extensive series of herbarium material
necessary for a revision of the genus was not available to him, but
he described ten new species and one new variety.

Since 1945, the only work on the genus has been the manuscript
of the Flora of Guatemala by Paul C. Standley and Julian A. Steyer-
mark (37) . They give a key to six species known from Guatemala.

A recent paper by J. W. Thieret (39) gives an account of the
tribes and genera of the Scrophulariaceae in Central America and
follows Pennell by placing Russelia alone in the sub-tribe Russelieae,
distinguished by densely packed hairs among the seeds in the capsule.

Several species of Russelia were introduced into Europe from
Mexico in the early 1800's and became popular greenhouse and
garden plants, which were reported as easy of cultivation and
propagation from cuttings. Their almost perpetually blooming,
showy, crimson flowers and their pendent habit of growth made
them especially desirable for hanging baskets in greenhouses. R.
equisetiformis (R. juncea), the rushy Russelia, was introduced into
Berlin and Munich about 1830 by Count Karwinski, and soon

CARLSON: THE GENUS RUSSELIA 233

afterward into England. R. sarmentosa, multiflora, floribunda,
rolundifolia, and the horticultural varieties, Lemoinei (R. juncea
X R. sarmentosa) and elegantissima, are mentioned in Edwards'
Botanical Register, 1836, and in the Journal of the Royal Horti-
cultural Society, 1851. The plants are known by the vernacular
names of "coral plant," "coral blow," "rain of fire" (lluvia de fuego)
and "ear-rings of the cook" (aretes de la cocinera).

Chromosome numbers of n=W have been reported for R.
equisetiformis and R. rotundifolia (10).

ABBREVIATIONS

F Chicago Natural History Museum

GH Gray Herbarium, Harvard University

LUND Lundell Herbarium

M Botanische Staatssammlung, Munich

MEXU National Herbarium, Institute of Biology, National University of

Mexico

MICH University of Michigan Herbarium

MO Missouri Botanical Garden Herbarium

NY New York Botanical Garden Herbarium

PA Academy of Natural Sciences, Philadelphia

SMU Southern Methodist University Herbarium

UC University of California Herbarium

US United States National Herbarium

Acknowledgments

My gratitude is extended to Dr. Paul C. Standley for suggesting
the problem, to Dr. Julian Steyermark for his help and criticism of
the manuscript, and to Dr. Theodor Just for permission to work at
Chicago Natural History Museum. Grants from the American
Philosophical Society, Philadelphia, the Garden Club of Evanston,
Illinois, and the Sarah Gildersleeve Fife Memorial Fund, New York,
made possible two collecting trips to study the genus in Mexico and
parts of Central America. The following institutions lent their
collections of Russelia: Academy of Natural Sciences, Philadelphia;
Gray Herbarium; Lundell Herbarium; Missouri Botanical Garden;
Munich Herbarium; National Herbarium, University of Mexico;
New York Botanical Garden; Southern Methodist University;
University of Michigan; University of California; United States
National Herbarium. To the curators of these organizations I give
sincere thanks.

234 FIELDIANA: BOTANY, VOLUME 29

Morphology

Plants. — The plants are perennial, suffrutescent or fruticose, erect
or with branches bending over at the tips, in some species scandent
or trailing. The stems may be from 0.5 to 5 or 6 meters long and
up to 1 cm. in diameter at the base. Some species branch profusely,
others only slightly.

Roots. — The perennial root system is not extensive. Adventitious
roots are produced at the nodes of the stems when they lie on the
ground.

Stems. — The stems are somewhat woody at the bases, becoming
herbaceous toward the upper parts. They may be terete, at least
at the base, or variously angulate, in most species with thickened
ridges extending lengthwise on the internodes. The ridges on the
angles may be narrow and inconspicuous (R. coccinea) or wide and
prominent (R. obtusata), and in R. tetraptera the ridges are on wing-
like extensions of the angles. The spaces between the angles may
be convex or flat, with (fig. 7, a) or without striations. In some
species the stems are glabrous, in others variously pubescent.

Leaves. — The leaves may be opposite or verticillate, thin and
membranaceous or thick and coriaceous, sessile or petiolate, and are
exstipulate. Those of R. equisetiformis are caducous. They may be
ovate (fig. 7, a), ovate-lanceolate, oblanceolate, or orbicular, with
apices acuminate, acute, or obtuse, and bases cuneate, obtuse, trun-
cate, or cordate. The margins are entire in four species and are
serrate, dentate, crenate or deeply incised in all others. The main
veins may be more or less impressed on the upper side and more or
less prominent on the lower side, and the smaller veins may be
inconspicuous or conspicuously reticulate. The leaves may be
glabrous or variously pubescent, and with or without resinous
dots.

Pubescence. — The hairs are uni- or multicellular (fig. 7, j, m),
sparse or numerous, long (1.6 mm., R. pubescens) or short (0.02-
0.05 mm., R. obtusata), straight (R. rotundifolia) or intertwined
in a wooly mat (R. villosa). Those on the stem may be present
principally on the angles (R. maculosa), principally between the
angles (R. villosa), or on the entire stem (R. retrorsa). The hairs on
the angles of R. retrorsa have swellings at the base which sometimes
become conspicuous callosities. The hairs on the leaves may be on
either or both surfaces; they are principally on the veins of the lower
surface in R. rotundifolia, between the veins on the upper surface of
R. retrorsa, and on the veins and also between them in R. villosa.

S.H. Grove

FIG. 7. a, Stem and leaf, R. sarmentosa; XI. b, Flower, R. sarmentosa; X 2
c, Pistil, R. sarmentosa; X 2. d, Capsule, R. syringaefolia; X 4. e, Corolla
interior, showing didynamous stamens, staminode, hairs along margins of median
ventral lobe and near insertion of stamens, R. sarmentosa; X 2. f-i, Sepals, X 4-
/, R. jahscensis; g, R. sarmentosa; h, R. cuneata; i, R. maculosa. j, Hair, R. relrorsa-
< 500. k, I, Seeds, X 50: k, R. tetraptera; I, R. rugosa; m, Hair, R. pringlei; X 500
n-q, Resinous dots or scales, X 500: n, R. standleyi; o, R. sarmentosa; p, R. jalis-
censis; q, R. retrorsa.

235

236 FIELDIANA: BOTANY, VOLUME 29

Most species have the margins of the leaves and calyx lobes ciliate,
especially the pubescent ones. Gland-tipped hairs are present on
all parts, including the external surface of the corolla in R. rotundi-
folia, maculosa, longisepala, and coccinea forma stipitata.

Resinous dots. — Resinous dots or scales are present on many
species. They may be circular or somewhat irregular in outline
(fig. 7, p), flattened against the surface (fig. 7, o), raised (fig. 7, q)
or sunken into the surface. They may be white, brown, yellow, or
translucent. They are attached at the center. They range in
diameter from 0.1 mm. in R. tepicensis to 0.3 mm. in R. jaliscensis,
and may be scattered, as on the leaf of R. sarmentosa (fig. 7, a), or
crowded, as in R. tepicensis. They may be present on the stems,
petioles, peduncles, pedicels, calyx lobes, and one or both surfaces
of the leaves. Often they are deciduous, leaving pits.

Petioles. — The petioles may be absent (R. rotundifolia), or up
to 8 mm. in length (R. polyedra), glabrous or pubescent, and with or
without resinous dots. The ridges on the angles of the stem extend
up the sides of the petioles for several millimeters. Usually a line
of hairs extends across the node, joining the bases of the petioles.
When the blade falls, the base of the petiole remains appressed to
the stem.

Inflorescences. — The inflorescences are axillary, one to three
from the axil of a leaf, on the main stem, or on floriferous branches.
The leaves (bracts) on the floriferous branches may be gradually
reduced in size toward the tip, or much reduced all along. The
inflorescences may be sessile as in R. rugosa, or with peduncles up to
4 cm. long as in R. equisetiformis, compact or spreading, and with
few or many flowers. The flowers may be arranged in simple or
complex dichasia (cymes). The pedicels are usually slender. Both
peduncles and pedicels may be glabrous or pubescent, with or with-
out resinous dots. The bractlets of the inflorescences are linear or
lanceolate, sessile, usually with hairs and sometimes with resinous
dots as well.

Calyx. — The persistent calyx is 5-parted almost to the base,
with the lobes ovate (fig. 7, h), ovate-lanceolate, or lanceolate
(fig. 7, /), the apex acute, acuminate, subulate (fig. 7, g), cuspidate
(fig. 7, i) or mucronate (fig. 7, k), usually with membranaceous
margins, and glabrous or pubescent on the dorsal side, sometimes
only on the midrib or margin, and with or without resinous dots.

Corolla. — The corollas (fig. 7, 6) are generally red, but may be
cerise, or mottled red, pink and white, as in R. maculosa, and are

CARLSON: THE GENUS RUSSELIA 237

externally glabrous, except in R. rotundifolia, maculosa, longisepala,
and coccinea forma stipitata, which have gland-tipped hairs. The
corollas are tubular, slightly bilabiate, the mouth not spreading,
except in R. polyedra, verticillata, and slaleyae, in which they are
funnelform. The upper (dorsal) lip is emarginate, the lower (ventral)
has three rounded lobes about 2-3 mm. in length. The upper lobes
are usually slightly shorter than the lower. The tube ranges in
length from 6-7 mm. (R. tepicensis) to 25 mm. (R. equisetiformis).
In all species except R. equisetiformis, flat, unicellular, yellow hairs
with rounded tips occur along the inner side of the median lower
lobe for about two-thirds of the length of the tube, these sometimes
being visible in the open throat, and similar shorter, smaller, flat
hairs occur on the inner surface of the tube near the insertion of
the stamens.

Stamens. — The four stamens are didynamous, included, two of
them 1 or 2 mm. longer than the other two (fig. 7, e). The filaments
are attached above the base of the corolla tube, the longer ones
slightly above the shorter. Hairs similar to those on the inner
surface of the tube near the attachment of the stamens are often
present on the bases of the filaments. The anthers are almost
spherical or are oval and divaricate. A short (1-2 mm.) staminode
with no anther is present in all species except R. equisetiformis, in
which it is 8 mm. long and has an abortive anther.

Pistil. — The pistil (fig. 7, c) is shorter than the tube of the
corolla. The stigma is not divided. The ovary is 2-loculed, with
enlarged placentae in each locule. The ovules are numerous and
anatropous.

Fruit. — The capsule (fig. 7, d) is globose or ovate, and glabrous;
the dehiscence is septicidal. The style is persistent, at least for
a time, but if it falls off it leaves a beak on the capsule. The lobes
of the persistent calyx are usually shorter than the capsule (fig. 7, d)
but sometimes almost equal it in length (R. coccinea).

After fertilization, the surface cells of the placentae between
the developing seeds elongate to form hairs which become longer than
the seeds and densely packed in the locules. This character is
distinctive for the genus.

Seeds. — The seeds are small (the largest, 0.5 mm. long, 0.3 mm.
wide), brown or black, elliptical or oval, reticulate, pitted, or with
elongate ridges on the surface (fig. 7, k, I).

Habitat. — The plants grow in thickets along roadsides, along
the margins of rivers and lakes, in fields, in forests, and on rocky

238

FIELDIANA: BOTANY, VOLUME 29

• R. SARMENTOSA

I I

* R. SARMENTOSA vnv NICARAGUENSIS

MAP 1. Distribution of three species and one variety of Russelia.

cliffs. The habitats of the individual species are given with the
descriptions of the species.

Distribution. — Three species are widely distributed, namely:
R. coccinea, extending southeastward from northeastern Mexico to
western Guatemala (Map 1) ; R. retrorsa, extending in a broad belt in
central-western Mexico, from Sinaloa to Morelos and Guerrero
(Map 3) ; and R. sarmenlosa, extending from Cuba and Vera Cruz to
Colombia (Map 1). Several species seem to be endemic to rather
localized areas, for example: R. acuminata, to the departments of
Quezaltenango and San Marcos, Guatemala (Map 1)', R. campechi-
ana, to the Yucatan Peninsula, British Honduras and Pete"n, Guate-
mala (Map 3) ; R. sonorensis, to the states of Chihuahua and Sonora,
Mexico (Map 2); R. sarmentosa var. nicaraguensis to Nicaragua
(Map 1); R. tennis, tepicensis and tetraptera to a strip of Pacific
coastal area between Sinaloa and northern Guerrero (Map 2).

MAP 2. Distribution of five species of Russelia.

» R. PURPUSM _
• R. POLYEDRA

» R. RETRORSA

MAP 3. Distribution of five species of Russelia.
239

240 FIELDIANA: BOTANY, VOLUME 29

KEY TO SPECIES, VARIETIES, AND FORMS

Al. Plants rush-like; branches slender, numerous, verticillate; inflorescences

2-flowered; interior of corolla lacking hairs Subgenus Juncirusselia

Main stems terete, with 8-12 inconspicuous ridges; branches 4-6-angled,

with ridges on the angles; leaves small, caducous; peduncles 2.5-3 cm.

long, ascending 1. R. equisetiformis

A2. Plants not rush-like; branches not numerous at the nodes, usually opposite

or ternate; inflorescences with more than 2 flowers; interior of corollas

with flattened hairs Subgenus Eurusselia

la. Leaves with entire margins, or with an occasional tooth.
2a. Stems appearing terete but with ridges.

3a. Vegetative parts of plant velutinous 2. R. purpusii

3b. Vegetative parts of plant glabrous 3. R. elongata

2b. Stems 4-angled, ridges on the angles.

4a. Petioles 7-9 mm. long; corollas 11-13 mm. long 4. R. campechiana

4b. Petioles 3-5 mm. long; corollas 15-25 mm. long 5. R. syringaefolia

Ib. Leaves with serrate, dentate or crenate margins.

5a. Stems and /or their ridges glabrous, or essentially so (R. obtusata may

appear glabrous, but has microscopic pubescence) (p. 242).
6a. Stems terete or subterete, or, if apparently angulate, not sharply so, with

ridges inconspicuous or poorly developed, the sides between the ridges

convex.
7a. Not resinous-lepidote on any part.

8a. Leaves long-elliptic, bases cuneate 3. R. elongata

8b. Leaves ovate, bases truncate 6. R. teres

7b. Resinous-lepidote on some part.

9a. Resinous-lepidote on both sides of leaves.

lOa. Resinous-lepidote on calyx lobes; resinous dots prominent on younger

leaves, sunken and inconspicuous on older leaves 7. R. acuminata

lOb. Not resinous-lepidote on calyx lobes; resinous dots on leaves prominent.

8. R. lanceifolia

9b. Resinous-lepidote on lower sides of leaves, not on upper sides or on calyx
lobes.

lla. Inflorescences with less than 10 flowers; margins of leaves deeply incised-
dentate 9. R. laciniata

lib. Inflorescences, except those toward top of stems, with more than 10 flowers;
margins of leaves not deeply incised-dentate.

12a. No stipitate glands present 10. R. coccinea

12b. Stipitate glands on peduncles, pedicels, calyx lobes and interior and exterior
of corollas 11. R. coccinea f. stipitata

6b. Stems sharply angled, or, if apparently subterete, with prominent ridges,

the sides between the ridges flat.
13a. Calyx lobes broadly lanceolate, gradually tapering from base to apex,

5-7 mm. long.
14a. Corollas 16-25 mm. long; calyx lobes 5 mm. long; leaves with large (diam.

0.3 mm.) scattered resinous dots on both sides 12. R. jaliscensis

14b. Corollas 9-11 mm. long; calyx lobes 6-7 mm. long; leaves with few small

sunken resinous dots only on the lower sides 13. R. longisepala

CARLSON: THE GENUS RUSSELIA 241

13b. Calyx lobes ovate, not gradually tapering from base to apex, less than
5 mm. long.

15a. Main stems with wing-like extensions of the angles, the wings 1 mm. wide,
ridged on the margins of the wings 14. R. tetraptera

15b. Stems without wing-like extensions of the angles, ridged on the angles, or,
if with wings, these less than 0.5 mm. wide and ridged on the margins of
the wings.

16a. Leaves obovate or rhombic-oblong, the widest part above the middle,
apices obtuse, bases sharply cuneate, blades attenuate to their attach-
ment; calyx lobes mucronate 15. R. cuneata

16b. Leaves and calyx lobes not as above.

17a. Not resinous-lepidote on any part.

18a. Petioles 3-4 mm. long; peduncles more than 5 mm. long.

23. R. sarmentosa f. eglandulata

18b. Petioles 1-2 mm. long; peduncles less than 5 mm. long.

19a. Leaves broadly ovate or rotund, apices acute or obtuse, bases truncate or
cordate 16. R. floribunda

19b. Leaves narrowly ovate, apices acuminate, bases widely cuneate or obtuse.

18. R. chiapensis
Resinous-lepidote on some part.

Calyx lobes not resinous-lepidote.

Both sides of leaves resinous-lepidote 19. R. standleyi

Lower sides of leaves resinous-lepidote, upper sides with no resinous dots

or very few.
22a. Leaves mostly finely reticulate-veined, glabrous; peduncles glabrous, 8-12

mm. long; pedicels glabrous, 7-10 mm. long 20. R. leptopoda

22b. Leaves not prominently reticulate- veined, short-pubescent, particularly
on the upper side; peduncles pubescent, 3-7 mm. long; pedicels pubescent,

2-6 mm. long •. 21. R. tennis

20b. Calyx lobes resinous-lepidote.

23a. Leaves on main stems more than 2 cm. long.

24a. Leaves ovate, apices acute or obtuse, bases truncate or rounded.

25a. Leaves on main stems mostly less than 6 cm. long; calyx lobes to 4 mm.
long, acuminate 22. R. sarmentosa

25b. Leaves on main stems mostly more than 6 cm. long; vegetative parts,
excepting sometimes veins on the lower side of leaves, glabrous; calyx
lobes to 5 mm. long, long-cuspidate .... 27. R. sarmentosa var. oaxacensis

24b. Leaves ovate-elliptic or ovate-lanceolate, apices acuminate, bases cuneate.
26a. Resinous dots numerous on both sides of leaves and on calyx lobes; leaves

deeply serrate 28. R. oxyphylla

26b% Resinous dots few or none; leaves shallowly serrate 29. R. longifolia

23b. Leaves on main stems less than 2 cm. long.

27a. Corollas tubular; ridges on angles of stems narrow; petioles 2.5-3 mm. long.

30. R. sonorensis

27b. Corollas funnelform; ridges on angles of stems wide; petioles 4-8 mm. long.

28a. Corollas 15-20 mm. long: canaliculate between the ridges which extend

downward from sides of petiole 51. R. polyedra

28b. Corollas 7-10 mm. long; not canaliculate on sides of stem below attachment
of petioles 32. R. verticillata

242 FIELDIANA: BOTANY, VOLUME 29

5b. Stems (at least those of the present year's growth) and /or their ridges
pubescent.

29a. Stems terete or subterete, or, if apparently angulate, not sharply so, with
ridges inconspicuous or poorly developed and sides between ridges convex.

30a. Corollas pubescent externally.

31a. Leaves sessile, rotund or subrotund, bases cordate, clasping; inflorescences
many-flowered; peduncles 8-17 mm. long.

32a. Older stems slightly pubescent, younger ones more so ... 33. R. rotundifolia

32b. All stems heavily pubescent 34. R. rotundifolia f . velutina

31b. Leaves petiolate, ovate, bases cuneate; inflorescences few-flowered; pe-
duncles lacking.

33a. Margins of leaves crenate-serrate, surfaces and veins on lower surface pilose
with septate hairs; calyx lobes 5-7 mm. long 35. R. rugosa

33b. Margins of leaves sharply serrate, all parts covered with short, gland-tipped
hairs; calyx lobes 8-10 mm. long 36. R. steyermarkii

30b. Corollas glabrous externally.

34a. Not resinous-lepidote on any part 37. R. pubescens

34b. Resinous-lepidote on leaves and calyx lobes.

35a. Leaves truncate or subcordate at base; corollas 6-8 mm. long.

38. R. tepicensis

35b. Leaves cuneate at base; corollas 9-11 mm. long 39. R. furfuracea

29b. Stems sharply 4-10-angled, or, if apparently subterete, the ridges prominent
and sides between ridges flat or concave.

36a. Stems with thin wing-like extensions of the angles, these 1 mm. wide, at
least on the main stems, and ridges on the margins of the wings; corollas
maculate 40. R. maculosa

36b. Stems with no wing-like extensions of the angles, or if with wing-like ex-
tensions these less than 0.5 mm. wide, and ridges on the angles; corollas
red.

37a. Ridges and striations of stems roughened by swollen bases of hairs, these
often becoming callosities 41. R. retrorsa

37b. Ridges and striations of stems not roughened by swollen bases of hairs or
callosities.

38a. Stems microscopically pubescent, the hairs scarcely visible to the naked
eye (0.02-0.05 mm. long) 43. R. obtusata

38b. Stems pubescent with longer hairs, readily visible to the naked eye.

39a. Pubescence densely woolly and matted, especially between the ridges,
white or cinereous.

40a. Leaves 1.5-3.5 cm. long, 1-3 cm. wide; peduncles 10 mm. long.

44. R. pringlei

40b. Leaves 5-10 cm. long, 4-6 cm. wide; peduncles 1 mm. long or lacking.

45. R. villosa

39b. Pubescence not densely woolly or matted, not white or cinereous.

4 la. Leaves not resinous-lepidote.

42a. Leaves narrowly ovate or obovate, bases cuneate.

43a. Hairs short, dense, velvety; calyx lobes mucronate 46. R. ternifolia

43b. Hairs longer, not dense or velvety; calyx lobes acuminate. . 18. R. chiapensis

42b. Leaves broadly ovate, bases truncate or cordate.

CARLSON: THE GENUS RUSSELIA 243

Inflorescences lax, peduncles 10-20 mm. long, pedicels 8-10 mm. long;

corollas 12-14 mm. long 47. R. conzattii

Inflorescences closely flowered, peduncles and pedicels 4-6 mm. long;

corollas 10-12 mm. long 17. R. floribunda var. pubescens

Leaves, at least the younger ones, resinous-lepidote on one or both sides.
Calyx lobes not resinous-lepidote.

Leaves with teeth 5-7 mm. long 48. R. grandidentata

Leaves with teeth less than 5 mm. long.

Inflorescences lax; petioles of larger leaves slender, 5 mm. long.

49. R. hintoni
Inflorescences closely flowered; petioles less than 5 mm. long.

Stem hairs on flat sides and on ridges 41. R. retrorsa

Stem hairs on flat sides; ridges glabrous.

Ridges of stems 0.2 mm. broad 21. R. tennis

Ridges of stems 0.5 mm. broad 42. R. retrorsa f. nudicostata

Calyx lobes resinous-lepidote.

Leaves resinous-lepidote mostly on upper sides.

26. R. sarmentosa var. nicaraguensis
Leaves resinous-lepidote on both sides.
Leaves on main stems more than 3 cm. long.

Stem hairs chiefly on angles 24. R. sarmentosa L pubescens

Stem hairs on both flat sides and ridges 25. R. sarmentosa f. velutina

Leaves on main stems 2.5 cm. long or less.

Corollas tubular.

Leaves densely resinous-lepidote; corollas 13-16 mm. long.

31. R. sonorensis var. pubescens

Leaves sparsely resinous-lepidote; corollas 11-12 mm. long. .50. R. parvifolia
Corollas funnelform.

Corollas 15-20 mm. long, petioles 6-8 mm. long 51. R. polyedra

Corollas 11-12 mm. long, petioles 1 mm. long 52. R. staleyae

List of Species

Russelia Jacquin, Enum. PI. Carib. 6. 1760.

Plants somewhat woody, shrubby or scandent, with terete or angulate stems,
usually ridged, and striate between the ridges; leaves exstipulate, opposite or
verticillate; inflorescences axillary, cymose, usually pedunculate, and with one,
two or more series of pedicels; calyces deeply 5-parted, persistent; corollas tubular
or funnelform, slightly irregular, the upper lip emarginate, the lower lip 3-lobed;
stamens 4, didynamous, inserted 1-2 mm. above the base of the corolla; staminode
usually short; ovary 2-loculed; fruit a globose capsule, loculicidally dehiscent,
with enlarged placentae bearing numerous hairs among the many, small, wingless
seeds.

Subgenus Juncirusselia Carlson, subgen. nov.

A. Plantae junciformes; ramis tenuibus numerosis verticillatis; inflorescentiis
2-floris; corollis intus glabris.

244 FIELDIANA: BOTANY, VOLUME 29

Type species: Russelia equisetiformis Schl. & Cham.

1. Russelia equisetiformis Schl. & Cham. Linnaea 6: 377.
1831. R. juncea Zucc. Flora 15, Beibl. 2: 99. 1832.

Plants about 1 meter tall, suffruticose, profusely branched, rush-like (re-
sembling Equisetum), with small, caducous leaves; branches verticillate below,
opposite above, spreading; stems green, glabrous, 4-12-angled, with prominent
ridges on angles and striations between them, upper stems very slender; internodes
5-7 cm. long; lower leaves verticillate, 3-6 in a whorl, ovate or elliptical, acute,
larger ones with one or a few large teeth on each side, 8.5-15 mm. long, 6-9 mm.
wide; upper leaves smaller, opposite, linear, entire, both surfaces glabrous, with
few circular resinous dots, veins not prominent; petioles 3-4 mm. long, often
remaining appressed to stem when leaves fall; inflorescences near top of plant,
opposite or verticillate, 1-2-flowered; bractlets small, linear, with few resinous
dots; peduncles 3-4 cm. long, pedicels 1-1.5 cm. long, both slender, glabrous;
calyx lobes broadly ovate, acute or mucronate, 2-3 mm. long, glabrous, no resin-
ous dots, not spreading; corollas red, 1.5-2.5 cm. long, outer surface glabrous,
dorsal lobe deeply notched, 3 ventral lobes about the same length as dorsal;
stamens to 18 and 20 mm. long, extending to mouth of corolla tube, without hairs
on filaments; anthers orbicular, divaricate, or somewhat pendulous; staminodes
very short (0.5-0.7 mm.); pistils 18 mm. long; style apiculate; capsules globose,
3-6 mm. in diameter; persistent calyx covers lower third of capsule; persistent
style to 15 mm. long; seeds small, oval, warty, light brown, among white hairs
which fill the capsule.

Type: From Papantla, Vera Cruz, Mexico, in sunny places,
Schiede & Deppe 1187 (GH).

Blooms throughout the year. Introduced into Berlin and
Munich by Count Karwinski and from there into Britain in 1833.
Usually an ornamental plant cultivated in greenhouses and in
gardens in tropical parts of the world. Known as "coral" or "lluvia
de coral" in Spanish-speaking countries.

Zuccarini says that R. juncea is distinguishable from R. equiseti-
formis by its "graceful, pendulent stems, glandulose-punctate leaves,
calyx lobes short, mucronate, and corolla twice as long," but J. M.
Greenman examined the material in the Berlin Herbarium and found
no satisfactory differences between them. After examining 120
specimens, I agree with this conclusion. The species differs from
other species in having slender, rush-like stems, small caducous
leaves, long corolla tubes and no hairs along the inside of the ventral
lobe or at the base near the attachment of the stamens.

Although the collectors have not always indicated, most of the
collections are cultivated plants. Besides the many specimens from
Mexico and Central America, collections have been made in the
West Indies, Panama, South America, the Pacific Islands, China,

CARLSON: THE GENUS RUSSELIA 245

India, the Malay Peninsula and Borneo. Even in the United States,
the plant has been collected in Michigan, Ohio and Missouri, as
well as in the southern states of Florida and Texas.

Subgenus Eurusselia Carlson, subgen. nov.

AA. Plantae non junciformes; ramis oppositis vel ternatis; inflorescentiis 3-
vel multi-floris; corollis intus pubescentibus.

Type species: Russelia sarmentosa Jacq.

2. Russelia purpusii Brandegee, Univ. Calif. Publ. Bot. 4:
385. 1913.

Plants shrubby, branched; stems appearing terete, but with inconspicuous
ridges and no striations between the ridges, all densely velutinous on and between
the ridges; internodes 3.5-5 cm. long; leaves opposite, broadly ovate, apex acu-
minate, base truncate or subcordate, 6-7 cm. long, 3.5-4.5 cm. broad, margin
entire, recurved, upper surface with few short hairs, more on margin and veins,
lower surface with more hairs, especially on veins, velvety to the touch, hairs
multi cellular, straight, 0.06 mm. long, upper leaves smaller, lanceolate, bract-like;
petioles 5-6 mm. long, pubescent; inflorescences axillary and terminal; primary
peduncles 2-4 cm. long, pubescent, bearing a pair of leaf -like bracts and spreading
cymes, each 3-6-flowered; bractlets small, linear, pubescent; secondary peduncles
7-10 mm. long, pedicels slightly longer, both pubescent; calyx lobes ovate, apices
acute or acuminate, 3 mm. long, with no hairs or glands; corolla red, tubular,
glabrous on the outside, 15-17 mm. long, dorsal lobe 1.5 mm. long, slightly notched,
ventral lobes slightly longer and more narrow, flat unicellular hairs along inside
of median ventral lobe; stamens 8 and 10 mm. long; anthers rounded, divaricate;
staminode 3-4 mm. long, bases of filaments and inside of corolla tube near their
insertion densely covered with flat hairs; pistil 10-11 mm. long; capsule ovate,
acuminate, 6 mm. to base of persistent style, which is 8 mm. long with knob-like
stigma; persistent calyx lobes cover lower half, or less, of capsule; seeds numerous,
small, black.

Type: Rocky forests at Bafios del Carrizal, Vera Cruz, Mexico,
August, 1912, C. A. Purpus 6039 (UC).

The only pubescent species with entire leaves (Map 3).
Mexico, Vera Cruz: Purpus 6039 (F, MO, NY, UC, US).

3. Russelia elongata Carlson, sp. nov. Plate 1.

Planta suffruticosa, 1-1.5 m. alta; ramis teretibus 4-multi-costatis glabris;
foliis verticillatis ramorum inferiorum caducis; laminis lanceolatis vel anguste
ellipticis 1-2.5 cm. longis 2-8 mm. latis, apice acuminatis, basi cuneatis, parce
serratis, glabris; petiolis tenuibus 2-3 mm. longis glabris; inflorescentiis verti-
cillatis axillaribus 3-5-floris; bracteolis lanceolatis glabris; pedunculis 3-8 mm.
longis glabris; pedicellis 2 mm. longis glabris; sepalis 3 mm. longis ovatis longe
acuminatis glabris; corollis 13-15 mm. longis tubulatis, lobis apice patulis; stam-
inibus 4 vel 5 mm. longis; antheris orbicularibus divaricatis; staminodiis 1 mm.
longis; pistillis 7 mm. longis; capsulis 3 mm. longis; seminibus ovalibus.

246 FIELDIANA: BOTANY, VOLUME 29

Plants suffruticose, 1-1.5 meters tall; stems terete, 4-many-ribbed, some of
the ribs wider than others, glabrous; leaves verticillate, caducous from the lower
parts of stems; blades lanceolate or long-elliptic, 1-2.5 cm. long, 2-8 mm. wide,
apices acuminate, bases cuneate, margins of larger leaves with few shallow teeth
each side, smaller ones with none, glabrous both sides, not resinous-lepidote;
petioles slender, 2-3 mm. long, glabrous; inflorescences verticillate, axillary,
3-5-flowered; bractlets linear, glabrous; peduncles 3-8 mm. long, glabrous;
pedicels 2 mm. long, glabrous; calyx lobes 3 mm. long, ovate, long-acuminate,
glabrous; corollas 13-15 mm. long, narrowly tubular, spreading at top, upper lip
scarcely emarginate, lower lobes 2 mm. deep, 2 mm. broad, barbate within along
median ventral lobe and at base near insertion of stamens; stamens 4 and 5 mm.
long; anthers orbicular, divaricate; staminodes 1 mm. long; pistils 7 mm. long;
capsules 3 mm. long; seeds oval, brown.

Type: Rocky cliffs, shady side, Sapopa Canyon, Rio Mayo,
Sonora, February 8, 1935, H. S. Gentry 1287 (F).

Distinguished from R. coccinea by the small, elongate leaves,
often in whorls of 4 or 5, caducous from lower parts of stems; the
few-flowered inflorescences, also verticillate in the axils of the verti-
cillate leaves; and the narrowly tubular corollas spreading at the top.

Mexico, Sonora: Sapopa Canyon, Rio Mayo, Gentry 1287 (F,
GH, MEXU, MICH, MO, NY, UC).

4. Russelia campechiana Standl. Contr. U. S. Nat. Herb.
23: 1309. 1924.

Plants suffrutescent or scandent, branching, 1.5 meters tall; stems 4-angled,
ridges conspicuous, scattered short appressed hairs; internodes 5-7 cm. long;
leaves opposite, ovate-lanceolate, 5-7 cm. long, 2.5-4 cm. broad, coriaceous,
lustrous on upper surface, glabrous, entire, margin reflexed, apex acuminate, base
rounded, minute impressed glandular dots on lower surface; petioles 7-9 mm. long,
older grooved on upper side, pubescent especially at base; inflorescences toward
top of stem, axillary, cymose, compact, many-flowered; bracts petiolate, like
leaves but smaller; bractlets linear, pubescent; peduncles 5-10 mm. long, pedicels
shorter, both pubescent; calyx lobes ovate, subulate-acuminate, the tips some-
times recurved, short hairs on costa and margin, 3.5 mm. long, no glandular dots;
corolla red, tube 11-13 mm. long, dorsal lobe not notched, 3.5 by 2.5 mm., ventral
lobes 3 by 1.5 mm., densely bearded with yellow hairs on inside of median lobe,
especially at throat, and at base near attachment of stamens; stamens 9 and 10
mm. long, hairs dense on bases of filaments; anthers oval, divaricate; staminode
3-4 mm. long, knob on top; pistil 10.5 mm. long; capsule globose, 1 cm. long
including beak; seeds small, black.

Type: Apazote, near Yohaltum, Campeche, Mexico, December
20, 1900, E. A. Goldman 467 (US).

Other collections in flower from December to July. Distribution :
Yucatan Peninsula and neighboring Pete"n, Guatemala, and British
Honduras (Map 3).

CARLSON: THE GENUS RUSSELIA 247

Mexico, Yucatan: Coba, Quintana Roo, Lundell & Lundell 7850
(LUND, PA). Campeche: Apazote, Goldman 1+67 (US); Tuxpena,
Lundell 1112 (F, LUND, MO, NY, PA, US), 1888 (F, GH, LUND,
MICH, NY).

Guatemala, Pet£n: La Libertad, Aquilar 464 (MICH); Yal'och,
BartleU 12852 (F, MICH, PA); Laguna Perdida, Lundell 1644
(MICH, US).

British Honduras: El Cayo, Bartlett 11999 (F, MICH), Chanek
231 (F, MICH, PA); Corozal, Gentle 385 (F, MICH, US), 647
(F, MICH, PA); Maskall Pine Ridge, Gentle 1140 (F, GH, MICH,
MO, NY, PA); El Cayo, Gentle 2175 (GH, MICH, PA); Vaca, 2269
(GH, LUND); Honey Camp, Lundell 576 (F, GH, MICH, PA, US);
Jacinto Hills, Schipp S653 (F, GH, MICH, MO, NY).

5. Russelia syringaefolia Schl. & Cham. Linnaea 6: 376.
1831. R. subcoriacea Robinson & Seaton, Proc. Amer. Acad. 28:
113. 1893.

Plants suffrutescent, scandent, branching, 5 meters tall; stems 4-sided, with
ridges on angles, inconspicuous striations between ridges, glabrous; internodes
5-8 cm. long, generally shorter than the leaves; leaves opposite, ovate, apices
acute or acuminate, bases rounded or cordate, margins entire, with thickened
edges, 5-8 cm. long, 4-6 cm. broad, subcoriaceous, firm, waxy above, glabrous,
minutely pitted on lower sides, main veins deeply impressed on upper side,
prominent on lower side; petioles 3-5 mm. long, thick, glabrous or slightly pubes-
cent, bases connected with a line, sometimes pubescent; inflorescences axillary,
sometimes two from the same axil, cymose, many-flowered; bracts petiolate, like
leaves but smaller, sometimes pubescent; peduncles and pedicels 8-10 mm. long,
pubescent; bractlets linear, pubescent; calyx lobes ovate, apices acute or acumi-
nate, 3-4 mm. long, glabrous; corollas 15-25 mm. long, scarlet, but cerise within
tube, densely bearded with flat, yellow hairs along inside of median ventral lobe,
especially at throat, and at base of tube near attachment of stamens; stamens
didynamous, 14 and 16 mm. long, bearded below; anthers oval, divaricate;
staminode 4 mm. long; pistil 15-18 mm. long; capsule long-ovate, acute, 8-12 mm.
long, enclosed at base by calyx; seeds 0.8 by 0.5 mm., surface rough, hairs among
seeds to 0.07 mm. long.

Type: Papantla, Vera Cruz, Mexico, December, Schiede &
Deppe 1186 (Vienna).

The species blooms throughout the year and grows on rocky
cliffs, up to elevations of 1,000 meters. It is related to R. cam-
pechiana. The specimens described as R. subcoriacea often have
larger leaves and flowers, but seem to agree with R. syringaefolia in
all other respects (Map 3).

Mexico, Tamaulipas: Adelaida Canyon, near Ciudad Victoria,
Banner 37 (F); Jaumave, near Nogales, von Rozynski 729 (F, NY).

248 FIELDIANA: BOTANY, VOLUME 29

Nuevo Leon: Dulces Nombres, Meyer & Rogers 2664 (F, MO, PA).
San Luis Potosi: El Salto, near Antiguo Morelos, Banner 31 (F),
Carlson 2762 (F); above Tamazunchale, km. 355, Carlson 2781 (F),
s.n. (F) ; Tamasopo Canyon, Pringle 5086 (GH) ; Rio de las Gallinas,
Purpus 5327 (GH, NY). Hidalgo: near Chapulhuacan, Lundell &
Lundell 12396, 12398, 12649 (LUND, SMU). Vera Cruz: Purpus
2378 (GH, MO, NY, UC, US), 8063 (UC). Oaxaca: Orcutt 3299
(MO); Barita de San Jose", Tolosa, Deam 41 (GH, US). Location
not given: Liebman 9434 (NY, US), Salasar s.n. (MEXU).

6. Russelia teres Lundell, Field and Lab. 13: 20. 1945.

Plants suffruticose, 1 meter high; stems and branches terete, with extremely
narrow, inconspicuous ridges, minutely puberulent at base of stems and at nodes,
otherwise glabrous; internodes to 12 cm. long; leaves opposite or ternate; petioles
to 2.5 mm. long, hirtellous; blades ovate or ovate-lanceolate, to 6 cm. long, 3.8 cm.
wide, apices acuminate, bases truncate or shallowly cordate, margins obscurely
crenulate-serrate, sparsely short-hairy on the veins on both surfaces, not resinous-
lepidote, short ciliate, finely reticulate-veined, costa and primary veins plane
above, rather prominent below; inflorescences axillary, many-flowered; bractlets
linear, sparsely hirtellous; peduncles 2-4 mm. long; pedicels 3-4 mm. long; calyx
lobes 5 mm. long, ovate, with long subulate tips, somewhat reflexed, glabrous
except for a few hairs along keel and tip, no resinous dots; corollas red, tubular,
slightly spreading at the top, 7-9 mm. long, upper lobe subentire, lower lobes
2 mm. long, glabrous externally, barbate within on median ventral lobe and near
insertion of stamens; stamens 4 and 5 mm. long, barbate at base; anthers oval,
divaricate; staminodes 1 mm. long; pistils 5 mm. long; capsules and seeds not seen.

Type: In woods, Coalcoman, Michoacan, Mexico, November 11,
1941, G. B. Hinton 16127 (SMU).

This species seems to be related to R. pubescens but differs in the
shape of the leaves and in the lack of the long soft hairs which are
characteristic of that species. Also, the calyx lobes of R. teres are
more narrow and have longer subulate tips than those of R. pubescens.
In some respects R. teres resembles R. coccinea, but the ridges on the
stems are less conspicuous and no resinous dots are present on any
part of the former.

Mexico, Michoacan: Hacienda Coahuayula, Emrick 181 (F);
Coalcoman, Hinton 16127 (GH, LUND, SMU, US).

7. Russelia acuminata Carlson, sp. nov. Plate 1.

Planta suffruticosa, 2-5 m. alta; ramis subteretibus vel inconspicue 4-6-
angularibus, glabris; foliis oppositis vel ternatis, petiolatis, petiolis 2-4 mm. longis,
hirtellis, laminis ovato-lanceolatis, 6-9 cm. longis, 3-4.5 cm. latis, apice acuminato-
attenuatis, basi cuneatis vel acutis, crenato-serratis, resinoso-lepidotis, venis prin-

CARLSON: THE GENUS RUSSELIA 249

cipalibus pubescentibus; inflorescentiis cymosis, 15-25-floris; pedunculis 10-15
mm. longis, glabris, parce resinoso-lepidotis; pedicellis 5-7 mm. longis, glabris;
sepalis ovatis, 3.5-4.5 mm. longis, aristatis, resinoso-lepidotis; corollis coccineis,
12-16 mm. longis; filamentis didynamis, 9 vel 11 mm. longis; antheris divaricatis;
pistillis 12 mm. longis; capsulis ovoideis, 4 mm. longis; seminibus numerosis,
minutis.

Plants suffruticose, 2-5 meters long, scrambling or drooping from cliffs;
stems subterete or with 4-6 angles slightly ridged, glabrous, not resinous lepidote;
leaves opposite or ternate, the blades 6-9 cm. long, 3-4.5 cm. broad, ovate-
lanceolate, cuneate to acute at base, acuminate-attenuate at apex, crenate-serrate,
more resinous dots on the lower than on the upper side, somewhat reticulate-veiny,
minutely pubescent on the main veins, on both upper and lower sides; petioles
2-4 mm. long, pubescent, especially on upper side; inflorescences laxly cymose,
with 15-25 flowers; peduncles 10-15 mm. long, glabrous, sometimes sparsely
resinous-dotted; pedicels 5-7 mm. long, glabrous; bractlets linear, pubescent;
sepals 3.5-4.5 mm. long, ovate with long, acuminate tips, densely resinous-dotted
at base, slightly hairy at tips; corollas red, tubular, 12-16 mm. long, externally
glabrous, internally densely bearded with yellow hairs on median lower lobe and
at base near attachment of stamens; ventral lobes 3 mm. long, 2 mm. wide, dorsal
lobe slightly emarginate, slightly shorter than the ventral; stamens 9 and 11 mm.
long; staminode 0.5 mm. long; anthers oval, divaricate; pistils 12 mm. long;
capsules ovoid, 4 mm. long, with beak; seeds numerous, small, black.

Type: Slender drooping stems somewhat woody, attaining length
of 2 meters, pendent from cliffs in the gorge of Rio Samala, near
Santa Maria de Jesus, Dept. Quezaltenango, Guatemala, at an
elevation of 1,670 meters, August 6, 1934, Alex. F. Skutch 951 (PA).

Other collections are from Quezaltenango and San Marcos,
at elevations of 1,200-2,000 meters (Map 1). In flower from
January to October. It is similar to R. coccinea (L.) Wetts., but
differs in having leaves generally longer for their width, with long-
acuminate apices; inflorescences more spreading and with fewer
flowers per inflorescence; and resinous dots on the calyx lobes as well
as on both sides of the leaves. F. W. Pennell gave this species an
herbarium name, but no description was ever published.

Guatemala, Quezaltenango: Rio Samala, near Santa Maria de
Jesus, Skutch 951 (F, PA), Palmer 1448 (PA); Finca Moca, Skutch
2085 (F, NY, PA, US); Calahuache", Standley 67099 (F, PA); El
Muro, below Santa Maria de Jesus, Standley 67165 (F, NY, PA),
67275 (F); moist cliff, Rio Samala, near Santa Maria de Jesus,
Standley 84737 (F), 84762 (F), 84856 (F), 86690 (F), 87111 (F),
87129 (F); Finca Pirineos, lower south-facing slopes of Volcan Santa
Maria, Steyermark 33214 (F). San Marcos: Between Finca El
Porvenir and Loma Corona, Volcan Tajumulco, Steyermark 37727
(F, PA).

250 FIELDIANA: BOTANY, VOLUME 29

8. Russelia lanceifolia Lundell, Field and Lab. 13: 14. 1945.

Plants suffrutescent, 2 meters tall; stems subterete, inconspicuously 6-angled
with narrow ridges on the angles, sparsely pubescent at first, then becoming
glabrous; leaves opposite or ternate, thin, membranaceous, ovate-lanceolate,
light green above, paler below, 7.5 by 3.5 cm., apices acute, bases cuneate, sparsely
pubescent on both surfaces at first, especially on veins on lower side, sparingly
resinous-lepidote on both surfaces with translucent dots, margins coarsely serrate-
dentate, 4-7 teeth on each side, basal one-third entire, ciliate, costa and veins
slightly impressed above, prominent and white beneath; petioles slender, 4-8 mm.
long, densely pubescent on upper side; inflorescences axillary, cymose, 3-9-
flowered; bractlets subulate, ciliate; peduncles 1-6 mm. long; pedicels 4-6 mm.
long, sparsely pubescent; calyx lobes ovate, acuminate, subulate above the middle,
4 mm. long, keeled, ciliate, sparsely hairy, not resinous-lepidote; corollas red,
cylindrical, 13-17 mm. long, lobes spreading, glabrous externally, barbate within
on lower lobe; filaments glabrous; staminodes 0.8 mm. long; capsules ovoid, 4 mm.
long, with persistent style; seeds 0.3 mm. long, light brown.

Type: Mountain-side near km. 157, Taxco, Guerrero, Mexico,
altitude 1,700 meters, August 1, 1943, C. L. & A. A. Lundell 12317

(SMU).

No flowers are present on the type specimen. Lundell says it is
closely allied to R. coccinea.

Mexico, Guerrero: Lundell & Lundell 12317 (LUND, MICH,

SMU).

9. Russelia laciniata Standl. & Steyerm. Field Mus. Bot.
22: 379. 1940.

Plants shrubby, branching, to almost 1 meter tall; stems slender, terete, with
striations, no ridges on older stems, very narrow ridges on younger stems, pale
green, glabrous; internodes elongate, 4-5 cm. long; leaves opposite, thin, mem-
branaceous, light grass-green both sides, 5-8 cm. long, 2-5 cm. wide, ovate or
rhombic-ovate, acuminate or long-acuminate, base wide-cuneate or subrotund,
margin deeply incised-dentate, the teeth triangular, sharply acuminate, the tips
somewhat recurved, veins prominent, especially on the lower side, few small,
resinous dots on the lower side, none on the upper; petioles less than 2 mm. long,
minutely pilose or puberulent, remaining and somewhat spreading after the blades
fall, giving the stem a nodose appearance; inflorescences axillary, few-flowered;
peduncles 8-10 mm. long, pedicels the same or longer, both glabrous; calyx lobes
4-7 mm. long, lanceolate or ovate, cuspidate-acuminate, glabrous; capsule ovoid,
glabrous, brown, 6 mm. long, tip a rigid beak, 3 mm. long; seeds 0.3-0.4 mm. long,
light brown, reticulate-ridged on surface. No information about the corolla is
available.

Type: Above Finca El Porvenir, along Rio Cab us, to within
2 miles of Cueva de las Palomas, south-facing slopes of Volcan
Tajumulco, Dept. San Marcos, Guatemala, altitude 1,300-1,500
meters, March 16, 1940, J. A. Steyermark 37982 (F).

CARLSON: THE GENUS RUSSELIA 251

Standley and Steyermark (38) state, "The material is incomplete,
only 2 flowers in fruiting state being present on the specimens. The
foliage is so unlike that of any other Russelia we have seen, especially
in its deeply laciniate leaf margins with somewhat outcurved
laciniations, that we have no hesitancy in describing the plant as
a new species."

Guatemala, San Marcos: Finca El Porvenir, J. A. Steyermark
37982 (F).

10. Russelia coccinea (L.) Wettstein, Engl. & Prantl IV, 3b:
62. 1897. Scrophularia coccinea L. Sp. PI. 1: 621. 1753. Russelia
multiflora Sims, Linnaea 6: 376. 1831; Curt. Bot. Mag. 37: t. 1528.
1813. R. paniculata Mart. & Gal. Bull. Acad. Brux. 12, II: 19. 1845.
R. serratifolia Lundell, Field and Lab. 13: 17. 1945.

Plants 3 meters tall with many stems not able to support themselves; stems to
8 mm. in diameter at base, subterete, but with narrow ridges and striations
between the ridges, glabrous, internodes 8-10 cm. long; leaves opposite or verti-
cillate, ovate-lanceolate, apices acute or acuminate, bases truncate or subcordate,
serrate, glabrous except for a few multicellular hairs on the upper side and on the
veins on the lower side, many small yellowish depressed resinous dots on the lower
side which make papillae on the upper side; petioles 4-5 mm. long, glabrous;
inflorescences one or two from an axil, near the ends of the branches, cymose and
sometimes appearing racemose, flowers numerous (15 or more); peduncles 5-8
mm. long, slightly pubescent; pedicels 3-5 mm. long, slightly pubescent; bractlets
linear, pubescent, ciliate; calyx lobes ovate-lanceolate, setaceous, membranaceous,
margins becoming purple, a few hairs on the midrib and tip, no resinous dots;
corollas 10-12 mm. long, the upper lip slightly notched, the lower lip 3-lobed, the
lobes 3 mm. long, slightly longer than the upper; flat hairs on the inner side of
the median ventral lobe of the corolla, and shorter flat hairs near the attachment
of the stamens; two stamens 5 mm. long, the other two 6 mm. long; anthers
divaricate; staminodes small (1 mm. long); pistils 7-8 mm. long; capsules 3 mm.
long, excluding beak, globose, no longer than the persistent calyx lobes; seeds 0.3
mm. long, black, embedded in hairs which fill the capsule.

Type: Near Tancanhuitz, San Luis Potosi, Mexico, E. Seler 705
(GH).

Figures are given by Paxton (Mag. Bot. 16: 163. 1849) and Sims
(35). The species grows in damp thickets, woods, and fields at
altitudes of 20-2,100 meters. It blooms throughout the year. It
is distributed from Tamaulipas, Mexico, to Guatemala (Map 1).
It is often cultivated in greenhouses and gardens.

Mexico, Tamaulipas: Barranca de Chavastla, Jaumave, von
Rozynski 759 (F). San Luis Potosi: Tamajundede, Aguirre & Reko
312 (NY); near Tamazunchale, Carlson s.n. (F), 2770 (F), 2782 (F),
2787 (F), Kenoyer 822 (F, MO), Lundell & Lundell 12401, 12402,

252 FIELDIANA: BOTANY, VOLUME 29

12406, 12422, 12650, 12651 (LUND, MICH, SMU); near Xilitla,
Rowell 3171 (SMU); near Tancanhuitz, Seler 705 (GH, US). Vera
Cruz : Banks of Mexican railroad, above Fortin, Barnes & Land 660
(F, PA); near Consequila, Purpus 8567 (US); hillsides, Chavarillo,
Barnes, Chamberlain & Land 47 (F); Orizaba, Botteri 28 (GH);
Cordova, Bourgeau 1983 (GH, US); near Rio Tonto, Tezonapa,
Carlson 1305 (F); Zacuapan, Purpus 6316 (UC), 6242 (UC); laguna,
near city of Vera Cruz, Greenman 73 (F, GH), 82 (F, GH); Cordova,
Greenman 185 (F, GH, NY); Puente de San Miguel, Langman 3433
(MEXU, PA); Orizaba, Mutter 1369 (NY), 4098 (NY, US); Cate-
maco, Nelson 385 (US); Cordova, Orcutt 3139 (F, GH, MO, US);
Perez, Orcutt 6556 (US); Barranca de Metlac, Pringle 6081 (GH,
MO, NY, PA, UC, US); Cordova, Purpus 436 (MO, UC, US);
Corral de Piedras, Purpus 5749 (F, GH, MO, NY, UC, US); San
Andres Tuxtla, near Matarapan, Seler 4965 (GH); Coatzocoalcos
River, Fortune, Williams 8521 (F, PA). Michoacan: Cerro de las
Naglas, More'lia, Arsene s.n. (F, US). Puebla: Near Metlatoyuca,
Goldman 62 (GH, US). Oaxaca: Canton de Huatusco, Conzatti 835
(GH); Wartenberg, near Tantoyuca, Huasteca, Ervendberg 223
(GH, NY, PA), 263 (GH, NY); Chiltepec and vicinity, Tuxtepec,
Martinez-Calderon 391 (F, GH, UC, US), 472 (GH, LUND, UC,
US); north of Tuxtepec, Nelson 319 (US). Chiapas: Barranca
between San Fernando and Chalona, Langman 3825 (PA) ; between
San Fernando and Plan de Ayala, west of Tuxtla Gutierrez, Langman
3916 (PA); Santa Elena, Acapetahua, Matuda 17251 (F); Jilguero,
Escuintla, Matuda 17273 (F); Santa Rosa Tonala, Matuda 17336
(F).

Guatemala, Chimaltenango : Champerin, Johnston 1743 (F).
Retalhuleu: Rio Samala, Shannon 271 (US); along road between
Retalhuleu and Adintal, Standley 87796 (F); on road between San
Sebastian and Santa Cruz Mulua, Standley 88178 (F). Suchi-
tepequez : Along Rio Madre Vieja above Patulul, Standley 62200 (F).
Escuintla: Between Rio Jute and Rio Pantaleon, on road between
Escuintla and Santa Lucia Cotz, Standley 63592 (F, PA); along
Rio Michatoya, southeast of Escuintla, Standley 89070 (F).

Cuba, Cienfuegos: Harvard Tropical Garden, Soledad, culti-
vated, Jack 4074 (US).

West Indies, Trinidad: Woodbrook, cultivated, Broadway 8006
(MO).

China, Canton: cultivated, Chun 6982 (NY).

CARLSON: THE GENUS RUSSELIA 253

11. Russelia coccinea f. stipitata Carlson, f. nov. R. glandu-
lifera Lundell, Field and Lab. 13: 12. 1945.

A var. coccinea recedit pedunculis, pedicellis, sepalis et corollis extus et intus
dense stipitato-glandulosis.

The form resembles the species except that the peduncles, pedicels, calyx
lobes and the exterior and interior of the corollas are densely covered with stipitate
glands.

Type: Tututepe'c, Oaxaca, Mexico, April 12, 1940, L. H. Bailey
604 (SMU).

The form appears in widely separated localities. Pennell recog-
nized this taxon as a species and named it R. glandulosa, but his
name and description were not published.

Mexico, Oaxaca: Tututepe'c, Bailey 604 (LUND, SMU). Chi-
apas: Siltepe'c, Matuda 136 (MICH, PA, US).

Guatemala, Guatemala: Pacaya, Amatitlan, Mart 1961 (GH,
MICH, NY, PA, US).

12. Russelia jaliscensis Robinson, Proc. Amer. Acad. 35:
319. 1900. R. deamii Robinson, Proc. Amer. Acad. 36: 474. 1901.

Plants suffrutescent, copiously branched, 1-1.6 meters tall; stems 4-6-angled,
narrow ridges on angles, striations on flat sides, glabrous; internodes 3-8 cm. long;
leaves thin, membranaceous, opposite, ovate, elliptic or ovate-lanceolate, apex
acute or obtuse, base cuneate, margin sharply serrate, ciliate, 2-3.5 cm. long,
half as broad; usually with sparse appressed multicellular hairs, but sometimes
glabrous, and usually with large (0.3 mm.) resinous dots, more on the upper than
on the lower side, sometimes deciduous; veins not prominent; petioles 2 mm. long,
pubescent, bases remaining after the blades fall; inflorescences axillary, 1-3-
flowered; bractlets linear, pubescent; peduncles 4-8 mm. long, pedicels 5-10 mm.
long, both slender, usually pubescent, not resinous-lepidote; calyx lobes 5 mm.
long, lanceolate, gradually tapering from base to apex, apices acute, ciliate, some-
times with a few hairs on the midrib and with resinous dots; corollas scarlet, 16-25
mm. long, externally glabrous, internally bearded with flat hairs along the median
ventral lobe and at the base near the insertion of the stamens; upper lip emar-
ginate, lower lobes slightly longer than the upper, 2-3 mm. long, rounded, some-
what spreading; stamens didynamous, 14 and 16 mm. long, with a few flat hairs
at the base; anthers ovoid, divaricate; staminodes 2-3 mm. long; pistils 10-17
mm. long; capsules 5-6 mm. long, globose; seeds oval, brown, 0.5 mm. long, 0.3
mm. wide.

Type: From a canyon near Guadalajara, Jalisco, Mexico, June,
1886, E. Palmer 126 (GH).

The species grows at altitudes from 900 to 1,600 meters, and
flowers from January to August. Its distribution is shown on Map 2.
It is recognized by the thin leaves, with large resinous dots on both

254 FIELDIANA: BOTANY, VOLUME 29

sides; the long calyx lobes, gradually tapering from base to apex;
and the long tubular flowers.

Mexico, Durango: San Ramon, Palmer 149 (GH, NY, US).
Jalisco: Guadalajara, Palmer 126 (GH, MEXU, PA), Pringle 2568
(F, GH, MEXU, MO, NY, US), 8657 (F, GH, MO, NY, PA, UC,
US), Russell & Souviron 18 (US). Michoacan: Morelia, Cerro Azul,
Arsene s.n. (F, US) ; Rincon, Arsene s.n. (US) ; Dos Tetecos, Arsene
s.n. (US). Morelos: Cuernavaca, Carlson 3119 (F), Deam s.n.
(F, MICH, MO), Lyonnet 608 (GH, MO, NY, US), Pringle 8934
(F, GH, MEXU, MO, NY, PA, UC, US). Guerrero: Pino Mina,
Hinton 10465 (F, GH, MO, NY, US). Mexico: Limon, Rusby 9
(NY); between Toluca and Mexico City, Rusby 12 (NY, US).

13. Russelia longisepala Carlson, sp. nov. Plate 2.

Planta suffruticosa, 3-5 dm. alta; ramis 4-6-angularibus, angulis incrassatis,
glabris; foliis oppositis, ellipticis, apice acutis, basi cuneatis, serratis, latis supra et
venis infra parce adpresso-pubescentibus, infra parce resinoso-lepidotis; petiolis

1 mm. longis, pubescentibus; inflorescentiis 2-4-floris; pedunculis 8 mm. longis,
glabris; pedicellis 3-5 mm. longis, glabris; sepalis 6-7 mm. longis, lanceolatis, costa
pubescenti; corollis 9-11 mm. longis; filamentis 7 vel 8 mm. longis; staminodiis

2 mm. longis; pistillis 7-8 mm. longis; capsulis 8 mm. longis, elongatis; seminibus
ovalis, numerosis.

Plants suffruticose, 3-5 dm. tall; stems 4-6-angled, narrow ridges on the
angles, glabrous except at ends of branches; internodes 2-5 cm. long; leaves
opposite, elliptic, apices acute, bases cuneate, serrate; upper sides and veins of
lower sides of younger leaves sparsely short-appressed-pubescent, lower sides
sparsely resinous-lepidote; smaller veins not evident; petioles 1 mm. long, pubes-
cent; inflorescences 2-4-flowered; bractlets long-oval; peduncles 8 mm. long,
glabrous; pedicels 3-5 mm. long, glabrous; calyx lobes 6-7 mm. long, lanceolate,
gradually tapering from base to apex, margins red-purple, pubescent in narrow
line along midrib; corollas 9-11 mm. long, lobes 4 mm. long, spreading, barbate
on inner side of ventral lobe near mouth, not at base; stamens 7 and 8 mm. long;
staminode 2 mm. long; pistils 7-8 mm. long; capsules elongate, 8 mm. long; seeds
oval, brown, numerous.

Type: Tamahu, Alta Vera Paz, Guatemala, altitude 1,300 meters,
April, 1908, H. von Tuerckheim 2249 (NY).

Distinguished by the long, gradually tapering calyx lobes and the
spreading 2-lipped corolla, with lobes longer than in most species.

Guatemala, Alta Vera Paz: Tamahu, von Tuerckheim 2249
(GH, NY, US).

14. Russelia tetraptera Blake, Proc. Biol. Soc. Wash. 33:
120. 1920.

Plants fruticose; stems 4-5 mm. thick at base, with opposite branches, gray-
green, sharply 4-angled, flat or concave with striations between the angles; angles

CARLSON: THE GENUS RUSSELIA 255

with corky-margined wings 1 mm. wide on the main stems, reduced above to mere
ridges, glabrate or puberulent and sometimes hispidulous on the flat sides; inter-
nodes 5-14 cm. long; leaves thin, opposite, 2.5-6.5 cm. long, 2-5 cm. wide, ovate,
apices obtuse or acutish, bases truncate-rounded or slightly cordate, crenate-
serrate or dentate with rounded or acute teeth; upper surface deep green, sparsely
incurved hispidulous or glabrescent, few glandular punctations; lower surface
scarcely paler, sparsely incurved hispidulous along 3-4 pairs of prominent veins,
profusely dotted with large (0.3 mm.) saucer-shaped glandular scales; petioles
3-7 mm. long, densely puberulous; inflorescences axillary, often 2 or 3 from the
same axil, cymose, spreading, with numerous flowers; bractlets linear, pubescent;
peduncles 2-5 mm. long, pedicels 1.5-5 mm. long, both hispidulous with slightly
curved hairs; calyx lobes 3-4.5 mm. long, ovate, green, pale at margins, 1-5-ribbed,
sparsely hispidulous along costa, narrowed into short filiform-subulate tips,
usually with no glandular dots; corollas 10-11 mm. long, externally glabrous,
barbate within with 1-celled flat hairs along ventral side of tube and at base near
insertion of stamens; upper lip emarginate, lower slightly longer, 3-lobed, lobes
equal, oblong-ovate, rounded, 2 mm. long; stamens 6 and 8 mm. long; anthers
oval, divaricate; staminodes 0.8 mm. long; pistils 9 mm. long; capsules subglobose,
glabrous, 3.5 mm. long; seeds small, ovoid, brown, surface with reticulate ridges.

Type: Tepic, Nayarit, Mexico, altitude 300-900 meters, January
5-February 6, 1892, E. Palmer s.n. (US).

Flowers from October to February; readily distinguished by its
4-winged stem and the compact inflorescences with numerous
flowers (Map 2).

Mexico, Sinaloa: Cerro Colorado, Gentry 5121 (GH, LUND,
MO, NY); Sierra Tacuichamona, Gentry 5657 (LUND); Hacienda
Chele, Rosario, Lamb 479 (GH, NY); Santa Maria, Culiacan,
Ortega 6605 (GH, PA, US). Nayarit: Cerro del Cruz, Collins &
Kempton 21 (US); San Bias, Ferris 5416 (US); Tepic, Barclay s.n.
(GH), Gregg 997 (MO), Jones 23217 (MO, UC); San Bias, Maltby 2
(US) ; Tepic, Palmer s.n. (US) ; Acaponeta, Rose, Standley & Russell
14313 (PA, US). Jalisco: 2 miles north of La Resolana, about 20
miles southwest of Autlan, McVaugh 10180 (MICH); Sierra du
Nayarit, Diquet s.n. (NY).

15. Russelia cuneata Robinson, Proc. Amer. Acad. 44:
613. 1909.

Plants fruticose, 1 meter tall, sparsely branched; stems mostly 4-angled,
sometimes 6-angled, with prominent ridges on the angles, flat, with fine striations
between the ridges, sometimes with sparse hairs, and brown or black spotted;
internodes 5-10 cm. long; leaves usually opposite, 6-10 cm. long, 4-5 cm. broad,
rhombic-oblong or obovate, apices obtuse, bases cuneate, crenate-serrate on upper
one-half or two-thirds; blades with a few multicellular hairs, especially on the
veins on the lower side, and with minute glandular scales, more numerous on the
lower than on the upper side, these early deciduous, leaving brown or black spots,
smaller veins prominently reticulate; blades sharply attenuate almost to point of

256 FIELDIANA: BOTANY, VOLUME 29

attachment and the ridges of the stem extending upward on the sides of the very
short petiole; inflorescences one or two from an axil, cymose, spreading, 10-
numerous flowers except at the upper nodes; bractlets small, linear, pubescent;
peduncles 5-20 mm. long, pedicels 4-10 mm. long, both with hairs and glandular
dots; calyx lobes ovate, short-cuspidate or mucronate, 3-4 mm. long, glabrous,
ridged, with or without glandular dots; corollas 10-12 mm. long, lobes of lower lip
2 mm. long, lobes of upper lip shorter, slightly emarginate, inside of corolla with
large (1.05 mm. long) flat hairs in upper part of tube, shorter ones near attachment
of stamens; stamens 5 and 6 mm. long; anthers rounded, divaricate; staminodes
0.3 mm. long; capsules ovoid, light brown, 3 mm. long; styles persistent, 6-7 mm.
long; persistent calyx lobes one-half length of capsules.

Type: From El Ocote, Michoacan, Mexico, altitude 300 meters,
December, 1898, E. Langlasse 723 (GH).

Mexico, Colima: Alzada, Orcutt 4674 (F). Michoacan: El Ocote,
Langlasse 723 (GH); Huizontla, Coalcoman, Hinton 16057 (GH,
LUND, US); Aquila, Coalcoman, Hinton 16129 (GH, UC, US);
Hacienda Coahuayula, Emrich 177 (F). Guerrero: Placeres Cam-
eron, Mina, Hinton 9188 (GH, US); Vallecitos, Monte de Oca,
Hinton 11393 (GH, US), 11585 (GH, US).

16. Russelia floribunda H.B.K. Nov. Gen. et Sp. 2: 359. 1817.
R. ovatifolia Lundell, Field and Lab. 13: 16. 1945.

Plants suffrutescent, 1-3 meters tall; stems 4-6-angled, with slightly thickened
ridges on the angles, striations on the flat sides, glabrous except for a line of hairs
at the nodes; internodes of the main stems 8-10 cm. long; leaves rigid-mem-
branaceous, subsessile, 5-7 cm. long, 4-6 cm. wide, opposite or occasionally
ternate, broadly ovate, apices usually obtuse but often acute, bases usually cordate
but sometimes truncate or rounded, serrate or crenate, with the primary veins
somewhat impressed on the upper side, somewhat prominent and sometimes
sparsely pubescent on the lower side, the surfaces scurfy; petioles 1-2 mm. long,
sometimes slightly pubescent; inflorescences axillary, profusely branched, many-
flowered; peduncles and pedicels 4-5 mm. long, glabrous; bractlets linear, ciliate;
calyx lobes 3 mm. long, ovate, acuminate-subulate, glabrous or with few hairs on
midrib, no resinous dots; corollas red, tubular, or slightly spreading at the mouth,
10-12 mm. long, externally glabrous, upper lip emarginate, lower lip 3-partite,
lobes 2 mm. long, rounded, tube bearded with flat hairs on inside of lower lobe
and at base near attachment of stamens; stamens 6 and 8 mm. long, inserted 1 mm.
above base of tube; anthers orbicular, divaricate; staminodes 0.3 mm. long; pistils
8-9 mm. long; ovary conical, glabrous; style filiform, glabrous, included; capsules
3-5 mm. long, subglobose, style persistent, at least for a time; seeds many, very
small, oblong, subangulate, dark, wrinkled.

Type: Between Papagallo and Venta de Tierra Colorada rivers,
Guerrero, Mexico, Humboldt & Bonpland s.n. (Herb. Mus. Paris).
Only a photograph of the type specimen was seen.

Related to R. rotundifolia, but differs in having less coriaceous
leaves, less prominent reticulations of the smaller veins, shorter
peduncles and no stipitate glands on the exterior of the corollas.

CARLSON: THE GENUS RUSSELIA 257

Mexico, Guerrero: Tierra Colorada, Humboldt & Bonpland s.n.
(Herb. Mus. Paris); 63 miles northeast of Acapulco, Carlson 3108
(F); Acahuizotla, Lundell & Lundell 12597 (LUND, SMU).

17. Russelia floribunda var. pubescens Carlson, var. nov.

A var. floribunda recedit ramis, pedunculis, pedicellis, petiolis et laminis
praesertim venis infra pubescentibus.

Similar to the species, except that the stems, both angles and flat sides, the
petioles, the peduncles, the pedicels and the leaves, especially the veins on the
lower sides, have a short, sometimes dense pubescence.

Type: Between Pie" de la Cuesta and Coyuca, 14.6 miles northwest
of Acapulco, Guerrero, Mexico, about sea level, January 21, 1955,
M. C. Carlson 3081 (F).

Mexico, Guerrero: Acapulco, Carlson 3049 (F), 3079 (F), 3081
(F), Langman 3360 (MEXU, PA), Palmer 252 (F, GH, MO, NY,
PA, UC, US); km. 417-422, Lundell & Lundell 12580, 12581, 12582
(LUND); km. 373, Lundell & Lundell 12595 (LUND).

18. Russelia chiapensis Lundell, Field and Lab. 13: 12. 1945.

Plants suffruticose; stems 4-6-angled, ridged on the angles, glabrous or
pubescent; internodes 3-4.5 cm. long; leaves opposite or ternate; blades charta-
ceous, narrowly ovate, up to 7 cm. long, 4 cm. wide, apices acute or acuminate,
bases broadly cuneate or obtuse, serrate, costa and primary veins impressed
above, prominent below, sparsely or densely pubescent; secondary veins reticulate,
with or without resinous dots; petioles 3 mm. long or less, glabrous; calyx lobes
ovate, 3-3.2 mm. long, acuminate, glabrous except for a few hairs along the keel;
corollas 12-13 mm. long, barbate in tube on lower side and at base near attach-
ment of stamens; stamens 6 and 8 mm. long; pistils 10 mm. long; capsules sub-
globose, 5.5 mm. long, apiculate; seeds minute, numerous.

Type: Mt. Ovando, Chiapas, Mexico, December, 1937, E.

Matuda2088 (SMU).

Similar to R. floribunda except for the longer and narrower leaf,
not at all cordate.

Mexico, Chiapas: Mt. Ovando, Matuda 2088 (MICH, SMU),
2088A (MICH, SMU), 2563 (MICH, SMU); Fraylesca, near
Siltepec, Matuda 5188 (LUND); Mt. Pasitar, Matuda 1643 (MICH).

19. Russelia standleyi Carlson, sp. nov. Plate 2.

Planta suffruticosa, 1 m. alta; ramis principalibus 6-angularibus, ramis
secundariis 4-angularibus, angulis incrassatis, glabris, flavoviridibus; foliis op-
positis vel ternatis; laminis 1-2.5 cm. longis, 8-12 mm. latis, apice acutis vel
obtusis, basi cuneatis vel truncatis, serratis, ciliatis, venis non prominentibus,

258 FIELDIANA: BOTANY, VOLUME 29

venis infra parce pubescentibus, utrinque parce resinoso-lepidotis; petiolis 2 mm.
longis, supra pubescentibus; inflorescentiis 3-5-floris; pedunculis 5-18 mm. longis,
glabris; pedicellis 5-7 mm. longis, glabris; sepalis ovatis, longo-acuminatis, glabris,
eglandulosis, marginibus purpureis; corollis 10-12 mm. longis, tubulosis; filamentis
9 vel 10 mm. longis; pistillis 10 mm. longis; capsulis globosis, 4 mm. longis;
seminibus reticulatis.

Plants suffruticose, 1 meter tall; main stems 6-angled, branches 4-angled,
angles thickened, glabrous, yellow-green; leaves opposite or ternate; blades 1-2.5
cm. long, 8-12 mm. wide, apices acute or obtuse, bases cuneate or truncate, serrate
with few large teeth, ciliate, both surfaces glabrous with few large (0.3 mm. diam.)
resinous scales; veins not prominent, somewhat pubescent on lower side; petioles
2 mm. long, pubescent on upper side; inflorescences 3-5-flowered; peduncles 5-18
mm. long, glabrous; pedicels 5-7 mm. long, glabrous; calyx lobes 3 mm. long,
broadly ovate at base, long-acuminate at tip, glabrous, without glands, the
margins purple; corollas 10-12 mm. long, barbate within on ventral lobe; stamens
9 and 10 mm. long; staminodes 1.3 mm. long, with knob at top; pistils 10 mm.
long; capsules globose, 4 mm. long; seeds reticulate.

Type: Mountains along road between Jalapa and San Pedro
Pinula, Dept. Jalapa, Guatemala, altitude 1400-1800 meters,
November 12, 1940, P. C. Standley 77093 (F).

Guatemala, Jalapa: Standley 77067 (F), 77093 (F), 76941 (F).

20. Russelia leptopoda Lundell, Field and Lab. 13: 15. 1945.

Plants shrub-like, 0.3 meters high; stems and branches rather slender, 6-
angled, glabrous; leaves opposite; petioles slender, 2.5 mm. long, white-villous
on upper side, at least when young; blades very thin, 1-3 cm. long, 0.6-1.5 cm.
wide, ovate or ovate-elliptic, apices obtuse, bases rounded or obtuse, dentate with
3-4 teeth on each side, glabrous except for a few hairs along the midvein on upper
surface, resinous dots on lower side, an occasional one on upper, finely reticulate-
veined; inflorescences cymose, usually 3-flowered; peduncles 8-12 mm. long,
pedicels 7-10 mm. long, both slender, glabrous; bractlets linear-lanceolate, ciliate;
calyx lobes 4 mm. long, ovate, subulate, glabrous, not resinous-lepidote; corollas
13-15 mm. long, tubular, stipitate glands and flat hairs within on lower lobe and
near base, upper lip emarginate, 2 mm. long, lower lobes slightly longer; stamens
7 and 8 mm. long; anthers oval, divaricate; staminodes 1 mm. long; pistils 8 mm.
long; capsules globose, 3 mm. long; seeds small, numerous.

Type: Ocorla, Coalcoman, Michoacan, Mexico, in woods, altitude
900 meters, June 25, 1939, G. B. Hinton 13845 (SMU).

Mexico, Michoacan: Ocorla, Coalcoman, Hinton 13845 (F, GH,
LUND, MO, PA, US).

21. Russelia tenuis Lundell, Field and Lab. 13: 19. 1945.

Plants suffrutescent, 1-2 meters tall; stems sharply 4-6-angled, with narrow
(0.2 mm. broad) ridges on the angles and striations on the flat sides, the ultimate
branches very slender, almost filiform, sparsely hirtellous at first, especially on
the flat sides and near the nodes; bases of petioles connected by a pubescent line;

CARLSON: THE GENUS RUSSELIA 259

leaves opposite or ternate; petioles 2-5 mm. long, pubescent; blades thin, ovate
or ovate-elliptic, up to 4 cm. long, 2.5 cm. wide, the apex acute or obtuse, the base
obtuse or truncate, entire, the rest of the margin rather coarsely serrate-dentate,
4-6 teeth on each side, sparsely pubescent with short hairs, especially on the upper
surface, small, scattered, resinous dots, sunken on lower surface, often with pro-
tuberances on upper surface opposite depressions on lower; costa and principal
veins slightly impressed above, somewhat prominent below, usually 3 on each side;
inflorescences cymose, axillary, usually 4-10-flowered; peduncles 3-7 mm. long,
pedicels slender, 2-6 mm. long, both rather densely pubescent with short, curved
hairs; bractlets linear, pubescent; calyx lobes 3 mm. long, ovate, acuminate, few
short hairs along keel, occasionally with few resinous dots; corollas red, tubular,
slightly wider at mouth, 10-15 mm. long; upper lip emarginate, lower lobes 2 mm.
long; corolla externally glabrous, internally barbate on lower side and at base
near attachment of stamens; stamens 7 and 9 mm. long; anthers oval, divaricate;
staminodes 0.3 mm. long; pistils 8.5 mm. long; capsules 3 mm. long; seeds small,
ovoid, brown.

Type: From Aquila, Coalcoman, Michoacan, Mexico, hanging
from cliffs, March 24, 1941, G. B. Hinton 15841 (SMU).

Other collections are from the states to the north along the
Pacific (Map 2), on rocky hills above the ocean, at elevations of
25-200 meters. In bloom from October to March. According to
J. G. Ortega, called "arete" in Mazatlan, Sinaloa.

Mexico, Sinaloa: northwest base of Cerro Colorado, Gentry 5189
(GH, MO, NY); La Nevada, north of Mazatlan, Mexia 98 (MO,
UC); La Noria, Mexia 225 (MO, UC), 225% (UC); La Nevada,
north of Mazatlan, Mexia 1097a (GH, UC, US); Mazatlan, Gregg
1185 (MO), Ortega 4637 (US), 6491 (PA, US), 6760 (F, PA), Rose,
Standley & Russell 13701 (PA, US), Wright 1203 (F, MO, UC, US);
Capule, Ortega 6062 (US). Nayarit: Maria Madre Island, Ferris
5614 (MO, US); Punta Mita, Howell 10397 (MICH, US); Maria
Madre Island, Maltby 114 (US), Nelson 4289 (GH, US). Jalisco:
Jayamita, Jones 394 (US); south of Puerto Vallarta, Mexia 1122
(F, MICH, MO, NY, US). Michoacan: Aquila, Coalcoman, Hinton
15841 (LUND, SMU, US). Colima: Manzanillo, Palmer 954
(GH, NY, PA, US). Durango: Tamazula, Gentry 5235 (F, GH,
MO, NY, UC).

22. Russelia sarmentosa Jacquin emend. Carlson. R.
sarmentosa Jacquin, Enum. PL Carib. 6, 25. 1760; Selec. Stirp.
Amer. Hist. 178. 1763. R. colombiana Pennell, Proc. Acad. Sci.
Phil. 72: 186. 1920. R. flavoviridis Blake, Contr. U. S. Nat. Herb.
24: 22. 1922. R. tabacensis Lundell, Contr. Univ. Mich. Herb.
6: 59. 1941.

260 FIELDIANA: BOTANY, VOLUME 29

Plants suffruticose, branching, 1-2 meters tall, erect or later reclining; stems
all 4-angled or main stems 6-angled, branches 4-angled, the angles often extended
as wings 0.1-0.5 mm. wide, with thickened ridges on the angles or on the margins
of the wings, the flat sides striate, usually glabrous but sometimes slightly pubes-
cent, and occasionally sparsely resinous-lepidote; internodes 5-13 cm. long; leaves
rigid-membranaceous, 1.5-8 cm. long, 1-5 cm. wide, opposite or ternate, ovate or
broadly ovate, apices acute or obtuse, bases truncate, subcordate or rounded,
margins serrate, dentate, or crenate, 5-9 teeth on each side, upper surface glabrous
and somewhat lustrous or sparsely pubescent with multicellular hairs, lower
surface glabrous or with hairs on the main veins, both surfaces with yellowish or
brownish resinous dots, these sometimes numerous, sometimes few, secondary
veins prominently reticulate; petioles 2-3 mm. long, hairs on each side and in
a band across the node between their bases; inflorescences cymose, 1-3 from each
axil, 3- to many-flowered; peduncles 4-18 mm. long, glabrous or slightly pubescent,
sometimes with a few resinous dots; pedicels 5-8 mm. long, glabrous or slightly
pubescent, often with resinous dots, especially at the top; bractlets linear or
lanceolate, ciliate; calyx lobes 4 mm. long, ovate, long-acuminate, sometimes with
hairs along the midrib and with few or numerous resinous dots; corollas 10-15 mm.
long, tubular, mouth slightly spreading, upper lip emarginate, lower lip 3-partite,
lobes 3 mm. long, 2 mm. wide, rounded at apex, flat hairs on interior of median
lower lobe and smaller similar hairs near insertion of stamens; stamens 6 and 8
mm. long; anthers ovoid, divaricate; staminodes 1-3 mm. long, with knob on top;
pistils 9-11 mm. long; ovary ovate-oblong, attenuate to style; capsules globose,
3-4 mm. long; calyx persistent, lobes little more than half as long as capsules;
seeds numerous, small, surface rough, embedded in hairs which fill the capsule.

Type : From woods and dense shrubby areas near Havana, Cuba,
Jacquin s.n., date unknown.

The type specimen was not seen, but a number of collections
from near Havana have been examined. Some of these agree with
Jacquin's description, which states that R. sarmentosa has 4-angled
stems and 3-flowered inflorescences. However, other specimens
from the same locality have 6-angled stems and inflorescences with
many flowers, although they agree with Jacquin's description in
other respects. This discrepancy has led to confusion.

I have studied numerous specimens from Mexico, Central
America, and Panama which are similar to those from Cuba. All
have 4- or 6-angled stems, with ridges of similar thickness on the
angles, opposite or ternate leaves of about the same shape, short
petioles, similar resinous dots on both sides of the leaves and on the
calyx lobes, and similar flowers. The specimens differ in the size of
the leaves, the amount of pubescence, the number of flowers on an
inflorescence, and the length of the peduncles. However, no con-
sistent differences in these characters could be found and attempts
to separate the plants into species or even into varieties were
fruitless.

CARLSON: THE GENUS RUSSELIA 261

F. W. Pennell separated R. colombiana from R. sarmentosa on the
basis of its 6-angled stems, larger, ternate leaves and longer pe-
duncles. However, even the type collection is variable in these
respects, as some of the stems are 4-angled and some of the in-
florescences have 3 flowers, and it agrees with Jacquin's description
of R. sarmentosa in other respects. Therefore, I cannot agree with
Pennell's separation of R. colombiana.

Pennell also thought that the specimens from Panama should be
placed in a species different from R. colombiana because of their
geographical separation, and that certain specimens with broadly
ovate leaves and large teeth, from Yucatan, Mexico, should be given
specific status, although he did not publish descriptions of these.
Here, too, I disagree with Pennell.

R. flavoviridis, described by S. F. Blake from a specimen from
eastern Guatemala, and R. tabacensis, described by C. L. Lundell
from a specimen from Tabasco, Mexico, fit the description of R.
sarmentosa as emended by Carlson. They were separated chiefly
because of their 6-angled stems and many-flowered inflorescences.

I have concluded that R. sarmentosa is a variable and widely
distributed species, ranging from Cuba, Tabasco and Yucatan,
Mexico, to Panama and Colombia (Map 1), and it is, therefore,
emended to include all the specimens listed below.

Cuba: Rocky banks near sea, near Havana, Curtis 753 (F, GH,
MO, NY, PA, US); Cojimar, Hermann 5067 (NY), Hitchcock s.n.
(F), Kittip 13816 (US), Leon 10656 (NY); Playa de Marianao,
Palmer & Riley 854 (US). Location not given, Auber 13 (GH),
Robbins s.n. (GH).

Mexico, Yucatan: Izamal, Gaumer 403 (F, MO, NY, PA, US);
Calotmul, Gaumer 1746 (GH); Cozum^l, Gamier s.n. (PA); near
Dzitas, Lundell & Lundell 7354 (F, GH, PA, US); Peto, Steere 2287
(F, MICH, MO). Campeche: Tuxpena, Lundell 1331 (F, US).
Quintana Roo: Dzitnup trail, Coba, Lundell & Lundell 7849 (PA);
Lake Chichancanab, Gaumer & sons 23660 (F, MO, NY, PA, US),
Steere 2484 (MICH). Vera Cruz: Mirador, Liebmann 9426 (NY,
US); Zacuapan, Purpus 2378 (F, GH, MO, NY, UC, US); Totutla,
Purpus 10833 (US). Tabasco: Estapilla, Tenosique, Matuda 3524
(F, MICH). Oaxaca: Monte Alban, Smith 1100 (GH); Mena,
Mell s.n. (NY) ; Chivela, Mell 38 (US), Orcutt 3299 (F, GH, MO, US).

British Honduras, El Cayo: Bartlett 11516 (F), Lundell 6131
(MICH); Pine Ridge, Duck Run, Bartlett 11531 (MICH, NY);
Vaca, Gentle 2248 (GH, MICH, PA); San Augustin, Lundell 6728

262 FIELDIANA: BOTANY, VOLUME 29

(MICH), 6925 (MICH), 6928 (MICH). Maskall: Pine Ridge,
Gentle 952 (GH, MICH, NY), O'Neil 8739 (GH, MICH, NY);
Honey Camp, Lundell 526 (F, GH, NY, PA, US), 526B (MO),
Meyer 117 (F), Record s.n. (US). Fresh Water Creek Reserve:
Pelly 15 (F). Location not given: Forestry Dept. 22 (F), Peck s.n.
(GH).

Guatemala, Izabal: Los Amates, Kellerman 5164 (US), 7609
(F, NY); Quirigua, Standley 23891 (US), 24508 (GH, NY, US),
72264 (F), 72282 (F); Izabal, Steyermark 38497 (F). Chiquimula:
El Rincon, Standley 74660 (F), 7-475-4 (F); East Vera Paz and
Chiquimula, Watson 474a (GH). Escuintla: Rio Guacalate,
Standley 58262 (F, PA) ; Escuintla, Standley 63944 (F, PA) ; Finca
Monterrey, Standley 64578 (F, PA) ; Rio Guacalate, Standley 89384
(F, PA). Jutiapa: Jutiapa, Standley 75484 (F), 75705 (F, PA),
75953 (F, PA) ; La Pava, Standley 77591 (F) ; Ovejero, Standley 77615
(F). Santa Rosa: Los Verdes, Standley 60401 (F, PA); Oratorio,
Standley 60668 (F) ; Cuilapan, Standley 77716 (F) ; Laguna Los Pinos,
Steyermark 52174 (F). Sacatepequez : Barranca Hondo, Standley
60235 (F), 60277 (F); Duenas, Standley 63289 (F, PA); Barranca
Hondo, Standley 64947 (F), 88923 (F). Alta Vera Paz: Trece Aguas,
Cook & Doyle 21 (US); Finca Mocca, Johnson 144 (NY, US);
Conception, Johnston 550 (F, PA), Steyermark 45281 (F, PA);
Parramos, Johnston 1561 (F); Tortuga, Steyermark 44626 (F).
Location not given: Heyde 525 (US).

Honduras, Morazan: Rio Yeguare, Glassman 1559 (F); El
Jicarito, Miller s.n. (F); Zamorano, Rodriguez 150 (F), Standley 1666
(F, PA); Tanque, Rodriguez 862 (F); Chauite, Rodriguez 924 (F);
Hoya Grande, Williams & Molina 12252 (F, GH); Tatumbla,
Williams & Molina 13290 (F). Santa Barbara: Los Dragos, Standley
& Lindelie 7416a (F).

El Salvador, Ahuachapan: Ahuachapan, Standley 20037 (US),
20099 (US), Standley & Padilla 2478 (F), 2495 (F, PA), 2840 (F).
Santa Ana: near Metapan, Carlson 789 (F). La Libertad: west of
Santa Tecla, Carlson 170 (F). San Salvador: San Salvador, Calderon
184 (GH, US), 827 (US); Los Planes, Carlson 24 (F); San Salvador,
Renson 172 (NY, US), Standley 19288 (GH, US), 22722 (GH, US),
28312 (GH, NY, US), 23565 (GH, NY, US), Velasco 9007 (US).
San Vicente: San Vicente, Standley & Padilla 3454 (F). Location
not given: Choussy 13 (US).

Costa Rica, San Jos£: San Ramon, Brenes 232 (F), 532 (F), 14408
(GH, US), 17004 (F); San Jos£ Highway, Rowlee & Rowlee 134 (US);

CARLSON: THE GENUS RUSSELIA 263

El General, Skuich 4134 (MO, NY, US); Camino del Hatillo,
Standley 32188 (US); Alajuelita, Tonduz 8775 (US). Cartago: Las
Concavas, Cooper 39 (F, NY) ; Livingston, Rowlee & Stork 686 (NY,
US) ; Turrialba, Skutch 4670 (F, GH, MO, NY, US) ; Dulce Nombre,
Standley 35904 (US). Guanacaste: Liberia, Dodge, Alfaro & Thomas
6245 (GH, MO, PA) ; Potrero, Dodge & Thomas 6434 (MO) ; Nicoya,
Pittier 13650 (US). Alajuela: El Coyolar, Standley 40070 (US);
Alajuela, Smith P2371 (GH). Puntarenas: Delta Rio Esquinas,
Allen 5624 (F) ; Puntarenas to Desmonte, Pittier 498 (US) ; Golfito
de Osa, Brenes 803 (F). Heredia: Heredia, Williams 16039 (F).
Not located by provinces: Vertiente del Pacifico, Biolley 17331 (GH,
US); Salinas Bay, Elmore Ell (MICH); Barbe, Leon 459 (F); San
Lucas, Orozco 236 (F) ; Tuccurique, Rowlee & Stork 846 (NY, US) ;
Cerro de Protti, Solis 328 (F); Jesus Maria, Stork 3328 (F); Rio
Tiliri, Tonduz 7326 (US).

Panama, Canal Zone: Tranquilla, Dodge & Allen 17324 (MO,
UC); Salamanca, Dodge, Steyermark & Allen 16967 (MO); Balboa
Heights, Greenman & Greenman 5024 (GH, MO) ; Paraiso, Hayes 36
(NY); Cerro Ancon, Heriberto 130 (US), Macbride & Featherstone
2771 (PA), Paul 149 (US); vicinity of Ancon, Piper 6021 (PA, US);
Rio Grande, near Culebra, Pittier 2125 (NY, US); Ancon Hill,
Standley 26325 (US); Balboa, Standley 26478 (US), 27000 (US),
29313 (US) ; Pueblo Nuevo, White 297 (MO) ; Penonome and vicinity,
Williams 185 (US). Taboga Island: Celestino 53 (US), Killip 3192
(US), Macbride 2771 (F, PA, US), Standley 27904 (US), Woodson,
Allen & Seibert 1477 (GH, MICH, MO, NY). Code": El Valle,
Allen 92 (GH, MO), 1156 (MO, PA), 2270 (US). Veraguas: San-
tiago, Allen 1081 (GH, MO, PA, US). Chiriqui: Cerro Vaca, Pittier
5384 (US); Rio Chiriqui Viejo Valley, White 77 (GH, MO, PA).
Panama: Juan Diaz, Cornman 2025 (US); Old Las Cruces Trail,
Standley 29141 (US).

Colombia, Santa Marta: Smith 1361 (F, GH, NY, PA, US).

23. Russelia sarmentosa forma eglandulata Carlson, forma
nov.

A var. sarmentosa recedit laminis et sepalis eglandulosis.

Similar to the species except that the leaves and calyx lobes are eglandulate,
or with only an occasional resinous dot.

Type: In thicket on river bank and plains, vicinity of Sigua-
tepeque, Comayagua, Honduras, altitude 1050 meters, July 1, 1936,
T. G. Yuncker, R. F. Dawson & H. R. Youse 5572 (F).

264 FIELDIANA: BOTANY, VOLUME 29

Honduras, Comayagua: Siguatepeque, Rodriguez 2704. (F),
Standley 55919 (F, US), 56230 (F, US), Standley & Chacdn 6244 (F),
6489 (F), Yuncker, Dawson & Youse 5572 (F, GH, MICH, MO, US);
Comayagua, Standley & Chacdn 5241 (F), 5525 (F), 6221 (F).

El Salvador, La Union: Standley 20883 (GH, MO, NY, US).

Panama, Chiriqui: Paso Ancho to Monte Lirio, Allen 1582 (GH,
MO, NY, PA, US) ; Bajo Mono, Boquete, Davidson 472 (F, GH, MO,
US).

Guatemala, Zacapa: Santa Rosalia, Steyermark 29308 (F, PA).

24. Russelia sarmentosa forma pubescens Carlson, forma
nov.

A var. sarmentosa recedit caulium costis pubescentibus, petiolis pedunculis
pedicellis laminis sepalis pubescentibus.

Similar to the species, but with hairs on the angles of the stems, on the petioles,
peduncles and pedicels, more dense on the leaves, especially on the veins, and on
the calyx lobes.

Type: El Cayo District, Mt. Pine Ridge, Vaquero, British Hon-
duras, July-August, 1936, C. L. Lundell 6869 (PA).

Mexico, Vera Cruz: Jalapa, MacDaniels 818 (F).

British Honduras, El Cayo: Mt. Pine Ridge, Vaquero, Lundell
6868 (MICH, NY, PA), 6869 (GH, LUND, NY, PA, US), Bartlett
11716 (MICH), Lundell 6706 (MICH). Toledo: Cow Pen, Gentle
4060 (GH, MO, NY).

Guatemala, Chiquimula: Lehmann 1722 (US). Alta Vera Paz:
Secanquim, Cook & Griggs 305 (US), Goll 99 (US), 143 (US); Serra-
quitche", Donnell-Smith 1642 (GH, US); Finca Volcan, Wilson 251
(F). Pete"n: La Libertad, Aguilar Ml 94 (F, LUND); Uaxactum,
Bartlett 12263 (PA, US), 12665 (MICH). Quiche": Nebaj, Skutch
1770 (F, PA, US). Huehuetenango: Finca Soledad, Steyermark
49555 (F).

Honduras, Morazan: Zamorano, Rodriguez 3150 (F), 3151 (F),
Standley 1349 (F), Williams & Molina 10051 (F, UC, US) ; Las Mesas,
Molina 964 (F), Williams & Molina 10118 (F, UC, US); Las Artillas,
Williams & Molina 14330 (F, US); Santa Clara Creek, Williams &
Molina 15843 (F); San Antonio de Occidente, Standky 487 (F);
Tegucigalpa, Von Hagen 1210 (F, NY). Olancho: Guaimaca, Molina
3078 (F). El Paraiso: Guinope, Molina 3350 (F, US), Rodriguez 1698
(F). Santa Barbara: San Pedro Sula, Thieme 5384 (GH, NY, PA,
US) . Amapala : Isla Tigre, Standley 20722 (US) . Province unknown :
Cuyamel, Carleton 462 (US).

CARLSON: THE GENUS RUSSELIA 265

El Salvador, San Vicente: San Vicente, Standley 21630 (GH, NY,
US), Standley & Padilla 3353 (F).

Costa Rica, San Jose": La Urura, Biolley 71 (F); Hatillo, Solis 82
(F); Las Pavas, Standley 36033 (GH, US); Santa Ana, Ledn 871 (F);
San Lucas, Orozco 333 (F); Las Concavas, Lankester K6 (F). Guana-
caste: Tilaran, Brenes 12696 (F); Hacienda Santa Maria, Dodge &
Thomas 6223 (GH, PA), 6223a (MO). Province unknown: Tacares,
Valeria 283 (F); Escosu, Solis 366 (F).

25. Russelia sarmentosa forma velutina Carlson, forma nov.

A var. sarmentosa recedit ramis petiolis pedunculis pedicellis laminis dense
velutinis.

Similar to the species, but with all vegetative parts densely velutinous.

Type: Seine Bight, British Honduras, November 25, 1930, altitude
1 meter, along sea beaches, W. A. Schipp 671 (GH).

British Honduras: Seine Bight, Schipp 671 (GH, MICH, MO,
NY, US).

Guatemala, Guatemala: Aguilar 68 (F). Santa Rosa: Santa
Rosa, Heyde & Lux 2943 (GH, NY, US) ; La Joya de Limon, Standley
78254 (F). Alta Vera Paz: Lanquin, Steyermark 44056 (F, PA).
Pete"n: La Libertad, Lundell 2457 (F, MICH), 3507 (F, MICH).

El Salvador, Santa Ana: Santa Ana, Standley 20412 (GH, US).

26. Russelia sarmentosa var. nicaraguensis Carlson, var.
nov.

A var. sarmentosa diff ert praecipue foliis subtus non resinoso-lepidotis vel parce
resinoso-lepidotis, nervis subtus dense pubescentibus, caulis angulis incrassatis
pubescentibus, atque foliis juvenilibus subtus punctis negris minutis instructis.

Similar to the species except that the thickened angles of the stem are usually
pubescent, the veins on the lower sides of the leaves are densely pubescent,
no resinous dots, or only an occasional one, are found on the lower sides of the
leaves, and minute black pits are sometimes conspicuous on the lower sides of
the leaves, especially on the younger ones.

Type: southwest slopes of Santiago Volcano, near Masaya,
Nicaragua, altitude 300-480 meters, July 5, 1923, W. R. Maxon
7434 (US).

The absence of resinous dots on the lower sides of the leaves and
dense pubescence on the angles of the stems and on the leaves,
especially on the veins of the lower sides, seem to be characteristic
of most of the specimens from Nicaragua. F. W. Pennell also recog-
nized this difference. The distribution is shown on Map 1.

266 FIELDIANA: BOTANY, VOLUME 29

Nicaragua, Esteli: Somoto, Morley 753 (GH, US). Chinandega:
Volcan El Viejo, Baker 101 (GH, MO), 880 (US), 2168 (GH). Carazo:
Vicinity of Jintepe, Standley 84-43 (F). Masaya: Masaya, Baker 197
(GH, MO, NY), 580 (US), 2215 (GH); southwest slopes of Santiago
Volcano, Maxon 7434 (US) , 7654 (PA, US) , 7665 (PA, US) . Managua :
Managua, Chaves 63 (US), Gamier 58 (PA), 267 (US), 4297 (GH), Rene
12 (US), 69 (US); Casa Colorada, south of Managua, Maxon, Harvey
& Valentine 7366 (PA, US), 7451 (US); Sierra de Managua, near El
Crucero, Standley 8184 (PA). Grenada: Mt. Mombacho, Grant 761
(GH). Chontales: San Miguelito, Shank & Molina 4583 (US);
vicinity of Juigalpa, Standley 9437 (F), 9449 (F), 9486 (F). Lake
Nicaragua: Ometepec Island, Shimek & Smith 20 (MEXU, MO, US),
Smith s.n. (MO). Jinotega: southwest of Jinotega, Standley 10109
(F); Cerro Sialci, Standley 10595 (F). No location given: Herndon
s.n. (US), Wright s.n. (GH, US).

Costa Rica, Guanacaste: Tilaran, Brenes 168 (F); Catalina, Stork
2788 (F, MICH).

Panama: Location not given, Bigelow s.n. (NY).

27. Russelia sarmentosa var. oaxacensis Carlson, var. nov.
Plate 3.

Planta suffruticosa, 3-4 m. alta, ramis striatis 4-6-angularibus, angulis parce
incrassatis, glabris, parce resinoso-lepidotis; foliis oppositis; petiolis 2-3 mm.
longis, glabris; laminis ovatis, 5-9 cm. longis, 3-5.5 cm. latis, apice acutis, basi
truncatis, crenatis, glabris, infra punctis minutis resinosis impressis, supra resin-
osis et papillosis, supra venis impressis, infra prominentibus, glabris; inflorescentiis
multifloris; pedunculis 3-8 mm. longis, glabris; pedicellis 4-6 mm. longis, glabris;
sepalis 5 mm. longis, ovatis, longe acuminatis, dense resinoso-lepidotis; corollis
10-12 mm. longis; filamentis didynamis, 7 vel 8 mm. longis; pistillis 10 mm.
longis; capsulis globosis, 4 mm. longis; seminibus parvis numerosis.

Plants suffruticose, 3-4 meters high, main stems 6-angled, the angles some-
times extended as wings, ridges on the angles or on the margins of the wings;
branches 4-angled with narrow ridges on the angles, striations between the ridges,
glabrous, with scattered resinous dots; leaves somewhat coriaceous, lustrous,
opposite; petioles 2-3 mm. long, glabrous; blades ovate, 5-9 cm. long, 3-5.5 cm.
wide, apices acute, bases truncate or sometimes broadly cuneate, crenate or
shallowly serrate, glabrous, both surfaces with small resinous dots depressed on
lower side and with papillae on upper side opposite the depressions; main veins
impressed on the upper side, prominent on the lower; smaller veins prominently
reticulate; inflorescences one or two from an axil, cymose or pseudo-racemose,
with numerous flowers; peduncles 3-8 mm. long, glabrous; pedicels 4-6 mm. long,
glabrous; calyx lobes 5 mm. long, ovate, long-acuminate, densely resinous-lepidote;
corollas 10-12 mm. long, upper lip slightly emarginate, lower lobes somewhat
longer, 1.8 mm. long, bearded with flat hairs along the ventral lobe and at the base
near the attachment of the stamens; stamens 7 and 8 mm. long, with hairs at the

CARLSON: THE GENUS RUSSELIA 267

base; anthers ovoid, divaricate; staminodes 1 mm. long; pistils 10 mm. long;
capsules globose, 4 mm. long; seeds small, numerous, black.

Type: Trail Yaveo to Rio Yaveo, District of Choapam, Oaxaca,
Mexico, altitude 450 meters, March 17, 1938, Mexia 9164 (US).

This variety is distinguished by the large crenate leaves with
numerous resinous dots, especially on the lower sides, and papillae
on the upper sides; the long- tipped calyx lobes, densely resinous-
lepidote; and the numerous flowers per inflorescence. According to
Mexia, it has long whip-like branches which are eaten by stock.
Pennell recognized the taxon, but did not publish a description.

Mexico, Oaxaca: Choapam, Nelson 857 (US); Yaveo, Mexia
9164 (F, GH, LUND, MO, NY, US); Ubero, Williams 9251 (F),
9254 (F, PA), 9321 (F, PA); Almoloya, Williams 9848 (F); Tolosa,
Beam s.n. (GH, MICH, US). Vera Cruz: Minatitlan, Mett s.n.
(NY), Smith 418 (MO); Fortune, Williams 9056 (F, PA). Tabasco:
Teapa, Rovirosa 893 (PA); Lomas de San Sebastian, Rovirosa 418
(PA, US), 151 (PA). Location not given: Johnson 115 (NY).

28. Russelia oxyphylla Lundell, Contr. Univ. Mich. Herb.
7: 51. 1942.

Plants herbaceous; stems 6-angled, with thickened ridges on the angles,
densely hairy at the nodes but sparsely pubescent or glabrous otherwise, the basal
internodes of the ultimate branches flattened on the inner side; leaves ternate,
membranaceous, ovate to ovate-lanceolate, 2.5-5 cm. long, 1.6-2.8 cm. wide, apex
acuminate, base sharply but broadly cuneate, margin above cuneate base deeply
dentate-serrate, teeth 3-4 mm. long, sparsely pubescent with short hairs along
main veins on both surfaces and on margin, many resinous dots on under surface,
fewer on upper; costa and primary veins slightly impressed above, conspicuous
beneath, 5 or 6 on each side; veinlets prominently reticulate; petioles 1 mm. long
or lacking; inflorescences axillary, few-flowered (2-6); bractlets linear, hirtellous;
peduncles 4-ribbed, 3-6 mm. long; pedicels slender, to 8 mm. long, with few
resinous dots; calyx lobes resinous-punctate, with few hairs on midrib, lobes ovate,
3.4-4.2 mm. long, apex subulate, ciliate, base subcordate or rounded; corollas red,
tubular, 12 mm. long, externally glabrous, sparingly short-barbate within on
apical two-thirds of lower side of tube and at base near insertion of stamens, the
emarginate upper lobe 2.3 mm. long, the lower lobes oblong, elliptic, 3 mm. long,
rounded at apex; stamens didynamous, 6 and 7 mm. long; anthers ovoid, divari-
cate; staminodes 4 mm. long; no fruits available.

Type: From Forest Reserve, near crossing of Cruces Trail and
Madden Dam road, Canal Zone, Panama, flowering, June 29- July
12, 1940, H. H. Bartlett & T. Lasser 16349 (MICH).

The species is related to R. sarmentosa Jacq. and may be a
variety of it, but differs in having more lance-elliptic leaves, broadly

. 268 FIELDIANA: BOTANY, VOLUME 29

cuneate at the base, shorter peduncles, and fewer flowers per in-
florescence.

Panama: Canal Zone, Forest Reserve, Bartlett & Lasser 16349
(MICH).

29. Russelia longifolia Carlson, sp. nov. Plate 3.

Planta suffruticosa, 1-1.5 m. alta; ramis 4-6-angularibus, glabris, striatis,
angulis alatis, incrassatis; foliis oppositis vel ternatis; petiolis 2-3 mm. longis,
glabris vel parce pubescentibus; laminis 7-10 cm. longis, 3-4 cm. latis, ovato-
lanceolatis vel ellipticis, apice longe acuminatis, basi cuneatis, serratis, glabris,
laminis novellis parce resinoso-lepidotis; venis secundariis tenuiter reticulatis;
inflorescentiis cymosis, 5-7-floris; pedunculis 10-12 mm. longis, glabris; pedicellis
5-8 mm. longis, glabris; sepalis 3-4 mm. longis, ovatis, glabris, basi parce resinoso-
lepidotis; corollis 12-14 mm. longis; filamentis didynamis 8 vel 10 mm. longis;
antheris divaricatis; pistillis 10-11 mm. longis; capsulis 3-4 mm. longis, globosis;
seminibus parvis, numerosis.

Plants suffruticose, 1-1.5 meters tall; stems 4-6-angled, wings on the angles,
ridges on the margins of the wings; leaves thin, membranaceous, opposite or
ternate; petioles 2-3 mm. long, glabrous or slightly pubescent; blades 7-10 cm.
long, 3-4 cm. wide, ovate-lanceolate or elliptic, apices long-acuminate, bases
cuneate, sharply serrate, glabrous; young leaves with resinous dots on both sides,
deciduous from older leaves; primary veins prominent on lower side, sometimes
purple red, secondary veins finely reticulate; inflorescences cymose, 5-7-flowered;
peduncles 10-12 mm. long, glabrous; pedicels 5-8 mm. long, glabrous; sepals 3-4
mm. long, ovate, apex long-acuminate, glabrous, few resinous dots at the base,
ridges along the keel; corollas 12-14 mm. long, ventral lobes 2.5 mm. long, 1.5
mm. wide, dorsal lip slightly shorter, slightly emarginate, barbate along median
ventral lobe and slightly so near base; stamens 8 and 10 mm. long; staminodes
3 mm. long; anthers divaricate; pistils 10-11 mm. long; capsules 3-4 mm. long,
globose; seeds small, numerous.

Type: On a rocky cliff along the Danto River, slopes of Mt.
Cangrejal, vicinity of La Ceiba, Dept. Atlantida, Honduras, July 30,
1938, T. G. Yuncker, J. M. Koepper & K. A. Wagner 8719 (F).

Honduras, Atlantida: La Ceiba, Yuncker, Koepper & Wagner
8719 (F, GH, MO, NY, US).

Guatemala, Izabal: Rio Dulce, near Livingston, Steyermark
39398 (F, PA).

30. Russelia sonorensis Carlson, sp. nov. Plate 4.

Planta suffruticosa, 1 m. alta; ramis gracilibus 4-6-angularibus, angulis in-
crassatis striatis glabris; foliis oppositis vel ternatis, ramorum inferiorum caducis;
laminis parvis (4-15 mm. longis) ovatis, apice acutis, basi truncatis vel obtusatis
glabris utrinque dense resinoso-lepidotis; petiolis 2.5-3 mm. longis, glabris; in-
florescentiis 3-5-floris; bracteolis linearibus pubescentibus; pedunculis 2-15 mm.
longis, glabris; pedicellis 4-6 mm. longis, parce pubescentibus; sepalis 2.5-3 mm.

CARLSON: THE GENUS RUSSELIA 269

longis, ovatis, apice acutis parce pubescentibus dense resinoso-lepidotis; corollis
13-16 mm. longis; staminibus 9 vel 10 mm. longis; staminodiis 2 mm. longis;
pistillis 11 mm. longis; capsulis 3 mm. longis; seminibus ovalibus.

Plants suffruticose, 1 meter tall; stems slender, 4-6-angled, the angles thick-
ened, striate on the flat sides, glabrous; leaves opposite or ternate, those on lower
parts of stems caducous; blades small (4-15 mm. long), ovate, apices acute, bases
truncate or obtuse, glabrous, both sides densely resinous-lepidote; petioles 2.5-3
mm. long, glabrous; inflorescences 3-5-flowered; bractlets linear, pubescent;
peduncles 2-15 mm. long, glabrous; pedicels 4-6 mm. long, slightly pubescent;
sepals 2.5-3 mm. long, ovate, acute, slightly pubescent along costa, densely
resinous-lepidote; corollas 13-16 mm. long, ventral lobes 2 mm. long, 2 mm. wide,
dorsal lip emarginate, slightly shorter, interior barbate along median ventral lobe
and at base near insertion of stamens; stamens 9 and 10 mm. long; staminodes
2 mm. long; pistils 11 mm. long; capsules 3 mm. long; seeds oval.

Type: From rock crevices, hill slope, Guasaremos, Rio Mayo,
Chihuahua, Mexico, August 10, 1936, H. S. Gentry 2374 (GH).

Distinguished by the leafless lower stems and the small leaves
on the ultimate branches at the time of flowering. According to
Gentry, the plant, called "pa-ah," is cooked in water and drunk as
a tea for fevers (Map 2) .

Mexico, Chihuahua: Guasaremos, Rio Mayo, Gentry 2374 (GH,
MO, UC); Barranca de Cobre, Hewitt 14 (GH); Mojarachic,
Knoblock 5623 (F); Rio Aros, Le Sueur 1381 (F, GH); location not
given, Palmer 249 (GH, US). Sonora: Los Conejos, Rio Mayo,
Gentry 1128 (F, MO); Caramechi, Rio Mayo, Gentry 1193 (F, GH,
MO); San Pedro de las Cuevas, Wiggins 7511 (MICH, UC, US).
Sinaloa: San Bias, Jones 23169 (MO, NY, UC). Hidalgo: Jacala,
Edwards 816 (F), Fisher 45184 (SMU).

31. Russelia sonorensis var. pubescens Carlson, var. nov.
A var. sonorensis recedit ramis foliisque pubescentibus.

Differs from the species in that the stems and leaves are pubescent.

Type: Alamos, Sonora, Mexico, March 13, 1910, J. N. Rose, P. C.
Standley & P. G. Russell s.n. (US).

Mexico, Sonora: Alamos, Drouet & Richards 3943 (F), Gentry
851M (MICH), Rose, Standley & Russell s.n. (US).

32. Russelia verticillata H.B.K. Nov. Gen. et Sp. 2: 360. 1817.
R. ephedroides Benth. PL Hartw. 69. 1840.

Plants suffrutescent; stems with the sides below the attachment of the leaves
extending downward from the base of the petiole to the next node as broad ridges

270 FIELDIANA: BOTANY, VOLUME 29

with narrow depressions between, these ridges usually flat but sometimes grooved
or concave, in which case the stem appears 6-8-angulate, with thickenings on the
angles; stems striate or not striate, usually glabrous but sometimes pubescent,
often yellow-green; branches frequently verticillate, slender; internodes 4-6 cm.
long; leaves verticillate, caducous, especially from the main stems; blades 0.3-0.6
mm. long, subrotund, apices obtuse, bases sharply cuneate, margins with 1 or 2
teeth, glabrous, resinous-lepidote on both sides; petioles 4-6 mm. long, glabrous
or hirtellous; inflorescences axillary, verticillate, cymose, with 3-10 flowers;
peduncles 2-4 mm. long or lacking, glabrous or hirtellous; pedicels 3-5 mm. long,
hirtellous; calyx lobes 4 mm. long, ovate-lanceolate, acuminate, hirtellous; corollas
red, 7-10 mm. long, gradually but slightly widening from base to mouth, upper lip
rotund, emarginate, lower lip 3-partite, lobes suborbicular, bearded along inside
of ventral lobe and at base; stamens didynamous, 6 and 8 mm. long, inserted at
base of corollas, bases of filaments dilated, glabrous; anthers oblong, divaricate;
staminodes 0.5 mm. long; ovary conical, glabrous, attenuated into the style;
styles filiform, glabrous, as long as the corollas; stigmas slightly thickened; cap-
sules globose or conical, glabrous; seeds small, numerous.

Type: Near Speluncam, Puente de la Madre de Dios, Hidalgo,
Mexico, no date given, Kunth s.n. (Herb. Mus. Paris).

The type of this taxon has not been examined, but two photo-
graphs, one of a specimen cited as the type in the herbarium of the
Museum of Paris, the other of a specimen in the Berlin Herbarium,
are at hand. Both of these specimens are poor and it has been
difficult to determine which collections are referable to this species.
It is evident that the species has not been understood and, as a
result, it has been a "catch-all" for hundreds of specimens. Most
of the specimens that have been identified as verticillata belong to
other species.

I have concluded, at least tentatively, until the opportunity
arises to study the type specimen, that R. verticillata and R. ephe-
droides are the same. An isotype of R. ephedroides and other col-
lections which are similar are at hand. These fit the rather in-
complete description of R. verticillata.

R. verticillata, with its verticillate branches and very small, often
caducous leaves, gives the general appearance of R. equisetiformis
but differs in its uniquely ridged stems and its much smaller flowers,
more than two per inflorescence, which are barbate within the
corolla.

Mexico, Vera Cruz: Orizaba, Balls 4272 (GH), Bolter i 123 (GH),
293 (GH), Bourgeau 2163 (GH), Le Roy 153 (NY), Mohr 307 (US),
Purpus 1276 (F, NY, UC); Yavijea, Galeottii 1109S (F); Nogales,
Matuda 1144 (MICH, PA). Oaxaca: Hacienda de Santa Ana,
Hartwig 501 (NY); Rio Zia, Liebman s.n. (NY, US).

CARLSON: THE GENUS RUSSELIA 271

33. Russelia rotundifolia Cav. Icones 5: 9. pi. 415. 1799.

Plants fruticose, erect, branching, 1.5 meters tall; stems subterete or obscurely
4-sided, ridges on the angles very narrow, tomentose or glabrescent; internodes
4-6 cm. long; leaves coriaceous, opposite, 5-7 cm. long, 4-6 cm. broad, sessile,
clasping, subrotund, apex obtuse, base cordate, coarsely dentate-crenate at base,
serrate above, no hairs on upper surface, numerous short hairs on margin and on
veins on lower surface, smaller veins forming conspicuous reticulum, especially on
lower surface; inflorescences axillary, spreading, 5-7 cm. long, pseudo-racemose,
the flowers alternating in two's, many-flowered; bractlets 2 mm. long, linear;
peduncles 8-17 mm. long; pedicels 3 mm. long; bractlets, peduncles, pedicels,
calyx lobes and outer surface of corolla tubes densely covered with hairs, some
multicellular and gland-tipped, others unicellular; calyx lobes 4-5 mm. long,
lanceolate, acuminate, often with the tip recurved; corollas red, tubular, 11 mm.
long, somewhat spreading at the mouth, upper lip deeply notched, lower lip
3-lobed, the lobes 2 mm. long, with flat hairs along inner surface of median lobe
and shorter hairs at base near attachment of stamens; stamens didynamous, 5 and
6 mm. long, hairs at base of filaments; anthers ovoid, divaricate; staminodes
1 mm. long; pistils 6-8 mm. long; capsules glabrous; seeds small (0.3 mm. long),
brown, pitted.

Type: Acapulco, Guerrero, Mexico, near sea level, October, 1894,
to March, 1895, E. Palmer 509 (F).

Mexico, Guerrero: Acapulco, Palmer 509 (F, GH, MO, NY, UC,
US); Montes de Oca, Vallecitos, Hinton 11773 (GH, NY).

34. Russelia rotundifolia forma velutina Carlson, forma nov.

A var. rotundifolia differt partibus omnibus dense velutinis.
Similar to the species, but all parts much more densely velutinous.

Type: Location not given, 1791, Haenke 1066 (F).
Mexico, ?: Haenke 1066 (F).

35. Russelia rugosa Robinson, Proc. Amer. Acad. 43: 58. 1907.

Plants fruticose; stems terete or subterete with 4-6 very narrow inconspicuous
ridges and fine striations between the ridges, pubescent with multicellular hairs
of unequal lengths, at least on the stems of the current year's growth; branches
slender; internodes 5-8 cm. long; leaves thin, opposite or ternate, generally
rugose, 5-7 cm. long, 3-4.5 cm. wide, elliptic, apices acute or obtuse, bases cuneate,
margins crenate-serrate except at the base, both surfaces pilose with few scattered,
long, septate hairs, especially beneath on the veins, no resinous dots, veins slightly
impressed on the upper surface, prominent and lighter colored on the lower;
petioles 4-5 mm. long, pilose; inflorescences axillary, sessile, 2-3-flowered, much
shorter than the leaves; peduncles lacking; pedicels slender, 4-6 mm. long, pilose;
calyx lobes linear-lanceolate, 5-7 mm. long, pilose; corollas tubular, red, 11-15 mm.
long, deeply cleft, lobes spreading-undulate, exterior with hairs, some gland-tipped,
upper lobe cleft 1.5 mm. deep, lower lobes 3 mm. long, 2.5 mm. broad, hairs near

272 FIELDIANA: BOTANY, VOLUME 29

throat on inside of ventral lobe of tube and at base near attachment of stamens,
the hairs much finer and more matted than in other species; stamens 3 and 6 mm.
long, with fine hairs at base; staminodes 2 mm. or more, with aborted anther;
anthers oval, divaricate; pistils 7 mm. long; capsules ovoid, 4 mm. long; seeds
large (0.8 mm. long), with parallel ridges running lengthwise.

Type: Gualan, Zacapa, Guatemala, altitude 128 meters, January
18, 1905, C. C. Beam 183 (GH). Known only from this locality.

Guatemala, Zacapa: Gualan, Deam 183 (F, GH, MICH, MO,
NY, PA, US), 6335 (GH, UC, US).

36. Russelia steyermarkii Carlson, sp. nov. Plate 4.

Planta suffruticosa, 1-2 m. alta; ramis teretibus, costatis, striatis; ramis secun-
dariis pubescentibus; foliis oppositis vel ternatis, ellipticis, apice acutis, basi cune-
atis, serratis, glanduloso-pubescentibus; petiolis 4-5 mm. longis, glanduloso-
pubescentibus; inflorescentiis sessilibus, 2-3-floris; pedicellis 5-10 mm. longis,
glandulosis; sepalis lanceolatis, 8-10 mm. longis, glanduloso-pubescentibus;
corollis 12-15 mm. longis, extus glanduloso-pubescentibus, intus tubo supra
basin et lobo ventrali prope faucem lanuloso-barbatis; filamentis 4 vel 7 mm.
longis, basi lanuloso-barbatis; pistillis 10-11 mm. longis; capsulis 4 mm. longis.

Plants suffruticose, 1-2 meters tall; stems terete with narrow ridges and
striate between the ridges; the slender lateral branches densely pubescent, the
main stems less so; leaves of the lateral branches opposite or ternate, elliptic,
apices usually acute, bases cuneate, margins sharply serrate except at the base,
both surfaces densely covered with short gland-tipped hairs, upper surfaces,
margins and veins of lower surfaces also with long appressed multicellular hairs;
petioles 4-5 mm. long, with gland-tipped hairs; inflorescences axillary, sessile,
2-3-flowered; peduncles lacking; pedicels 5-10 mm. long, densely covered with
gland-tipped hairs; calyx lobes lanceolate, 8-10 mm. long, densely covered
with gland- tipped hairs and also with longer hairs on margins, midrib and tip;
corollas red, 12-15 mm. long, usually with hairs, some gland-tipped on exterior of
tube, lobes somewhat spreading; few fine, matted hairs on inside of ventral lobe
and more at base near attachment of stamens; stamens 4 and 7 mm. long, the
shorter attached at a higher level than the longer, fine hairs at base; staminodes
0.2 mm. long; pistils 10-11 mm. long; capsules 4 mm. long, longer than broad.

Type: In ravine along Rio Santa Clara, valley of Vegona, between
Vegas and Calera, Sierra de las Minas, Zacapa, Guatemala, altitude
1,200 meters, January 20, 1942, J. A. Steyermark 42980 (F).

This species is similar to R. rugosa but differs in having smaller,
non-rugose leaves with more sharply serrate leaf margins, longer
calyx lobes, more elongated capsules and all parts densely covered
with short gland-tipped hairs in addition to long pointed multi-
cellular hairs. According to Steyermark, its leaves have a fetid
odor.

Guatemala, Zacapa: Valley of Vegona, Steyermark 42980 (F, PA).

CARLSON: THE GENUS RUSSELIA 273

37. Russelia pubescens Lundell, Amer. Midi. Nat. 29: 491.
1943.

Plants suffrutescent, erect, 1.5-2 meters tall; stems terete, but with 4-6
inconspicuous ridges and striations between the ridges, pilose, with long, fine,
straight, multicellular hairs of irregular lengths; internodes 4-10 cm. long; leaves
opposite or ternate, 4-10 cm. long, 3-6.5 cm. broad, acute or obtuse, base rounded
or subcordate, crenate-dentate, velvety, pilose on upper side and principally on
veins on lower side, ciliate on margins, membranaceous or thin-chartaceous, not
resinous-lepidote, veins 5-7 each side of midrib; petioles 2 mm. or less, pilose;
inflorescences axillary, 10-12-flowered; bractlets linear-subulate; peduncles 5 mm.
long, pilose; pedicels 3.5 mm. long, pilose; calyx lobes 3-4 mm. long, ovate,
abruptly long-cuspidate for one-half or three-fourths of their length, tips some-
times reflexed, sparse long hairs on margins, midrib and chiefly on cuspidate tip;
corollas crimson, 9-12 mm. long, tubular, slightly constricted one-third above
base, lobes undulate, dorsal lobes 3 mm. long, shallowly emarginate, ventral lobes
almost as long, rounded, barbate within on median ventral lobe and at base near
attachment of stamens; stamens 3 and 4 mm. long, filaments barbate at base;
anthers ovoid, divaricate; staminodes 0.7 mm. long; pistils 4-5 mm. long; capsules
ovoid, glabrous, 6 mm. long, with beak; seeds small, brown.

Type: From woods at Hacienda del Ototal, east of San Se-
bastian, Jalisco, Mexico, altitude 1500 meters, abundant, March 5,
1927, Ynes Mexia 1815 (MICH).

Lundell says: "Distributed as R. iepicensis Rob., a species with
subappressed sparse pubescence, resinous-punctate leaf blades,
calyx lobes scarcely half as long, and corolla only 6-7 mm. long.
R. pubescens is allied to R. rotundifolia Cav., but the small, crowded
cymes immediately distinguish it from that species." It is also
related to R. floribunda H.B.K., but the terete stems and the long
hairs distinguish it from this species.

Mexico, Jalisco: Quimixto, trail from San Pedro el Tuito, Mexia
1238 (UC, US); east of San Sebastian, Mexia 1815 (F, GH, MICH,
MO, NY, PA, UC, US). Colima: Manzanillo, Ferris 6082 (US).

38. Russelia tepicensis Robinson, Proc. Amer. Acad. 35:
321. 1900.

Plants suffrutescent, less than 1 meter tall, branching; branches opposite or
verti ciliate; stems terete or subterete, with 4-8 thin (narrow) ridges, striations
between them, densely hirsute, sometimes with minute resinous dots; internodes
2-6 mm. long; leaves opposite, ternate, or occasionally in whorls of 4, membran-
aceous, broadly ovate, apices obtuse, bases truncate or subcordate, crenate-
serrate, 3-4 cm. long, 2-3 cm. broad, strigose with a few long, appressed hairs
along the main veins of both surfaces and numerous, minute (diam. 0.1 mm.),
translucent resinous dots crowded on the lower surface, sometimes much less
abundant on the upper, main veins impressed on the upper side, prominent on the
lower, secondary veins reticulate on both surfaces; petioles 2-4 mm. long, ap-

274 FIELDIANA: BOTANY, VOLUME 29

pressed, pubescent; inflorescences short, cymose, 5-8-flowered, in axils of leafy
bracts; bractlets linear; peduncles and pedicels 2-5 mm. long, all with hairs and
minute resinous dots; calyx lobes 2-2.5 mm. long, ovate, cuspidate, the tips
sometimes recurved, margins thin, few hairs along midrib, numerous minute dots;
corollas red, 6-8 mm. long, tubular, spreading at the top, externally glabrous,
ventral lobes 2.5-3 mm. long, 2 mm. broad, rounded, dorsal lip emarginate, bearded
on inside of ventral lobe and near the insertion of the stamens; stamens 3 and 4
mm. long; anthers oval, divaricate; staminodes 0.5 mm. long; pistils 3-4 mm. long;
capsules ovoid, 3-4 mm. long, equal to or slightly longer than the calyx lobes;
styles persistent; seeds rough, 0.5 mm. long, 2.5 mm. wide.

Type: From Zopelote, Nayarit, Mexico, altitude 600-900 meters,
in flower and fruit, February, 1895, F. H. Lamb 578 (GH).
The distribution is shown on Map 2.

Mexico, Nayarit: Acaponeta, Tiger Mine, Jones 23170 (MO, UC) ;
Zopelote, Lamb 578 (GH, MO, NY, PA, UC, US) ; Acaponeta, Rose,
Standley & Russell 14354(PA, US).

39. Russelia furfuracea Brandegee, Zoe 5: 219-220. 1905.

Plants suffrutescent, many stems from the base, 2-3 dm. high; stems terete,
with very thin ridges and striations, densely hirsute, many minute resinous dots;
internodes 2-5 cm. long, often shorter than the leaves; leaves opposite or ternate,
ovate, apices acute or obtuse, bases cuneate, coarsely but not deeply crenate-
serrate, slightly pubescent on upper side and on veins on lower side, both sides
with numerous, crowded minute circular dots, larger leaves 3-4 cm. long, 2-2.5
cm. broad, main veins impressed on upper side, prominent on lower; petioles short
(1 mm. long); inflorescences near top of numerous branches, in axils of leafy
bracts, cymose, few-flowered; bractlets linear; peduncles and pedicels 3-5 mm.
long, with hairs and minute resinous dots; calyx lobes 3 mm. long, ovate, long-
acuminate, few hairs, numerous resinous dots; corollas scarlet, 9-11 mm. long,
spreading at the top, ventral lobes 2.5-3 mm. long, 2 mm. broad, rounded, dorsal
lip emarginate, the whole about the size of the ventral lobe, few flat yellow hairs
inside of tube along median ventral lobe and many at base near attachment of
stamens; stamens 3 and 4 mm. long; anthers divaricate; staminodes 1 mm. long;
pistils 5 mm. long; capsules ovate, acuminate, about as long as the calyx lobes;
seeds oval, brown, finely reticulate on surface.

Type: From cliffs in sunny exposures of Cerro Colorado, Culia-
can, Sinaloa, Mexico, T. S. Brandegee s.n. (UC).

This species is similar to R. tepicensis, but differs in that the
leaves are cuneate at the base while those of lepicensis are truncate,
the bracts which subtend the inflorescences are larger, and the
corollas are longer. Standley (36) states that it is either synony-
mous with, or a form of R. tepicensis.

Mexico, Sinaloa: Culiacan, Brandegee s.n. (GH, PA, UC, US);
oak forest, Cerro Colorado, Gentry 5076 (F, GH, LUND, MO, NY,

CARLSON: THE GENUS RUSSELIA 275

UC) ; Caperadero, Sierra Tacuichamina, Gentry 5563 (GH, LUND,
MEXU, MO, NY).

40. Russelia maculosa Lundell, Contr. Univ. Mich. Herb.
6: 57. 1941.

Plants herbaceous, clambering, 2 meters long, with slender branches; stems
sharply angled, usually 4- sometimes 6-angled, the angles with wing-like extensions
about 1 mm. wide and pubescent ridges on the margins of the wings, flat sides
striate, usually glabrous; internodes 3-6 cm. long; leaves chartaceous, opposite,
1.5-3.5 cm. long, 1-2.5 cm. wide, ovate, apices acute, bases truncate or rounded,
conspicuously serrate above base, with long, scattered, multicellular hairs on
upper side, these only on veins on lower side, resinous-lepidote, with numerous
scales on lower side, fewer on upper side, primary veins 3-5 on each side of midrib,
impressed on upper, elevated on lower side; petioles 3-5 mm. long, densely pu-
bescent with same type of hairs as on the leaves; inflorescences axillary, solitary,
2-5-flowered; bractlets narrowly linear or subulate, up to 4 mm. long; peduncles
4-angled, 2-4 mm. long; pedicels 2-4 mm. long; bractlets with sparse long hairs
especially on the margins, and peduncles and pedicels sparsely pubescent and
densely resinous-lepidote; corollas variegated maroon, pink and white, 13 mm.
long, distinctly bilabiate, essentially 4-lobed, the 3 lower lobes 4 mm. long, the
upper shorter, slightly emarginate; corollas barbate with narrow flat hairs on inner
surface of lower lobe- and with shorter flat hairs and gland-tipped hairs near
attachment of stamens; stamens 6 and 7 mm. long, hairs at base; anthers small,
slightly oval; staminodes 1 mm. long; pistils 8 mm. long; capsules 3-5 mm. long,
globose, glabrous; seeds brown, oval, 0.5 mm. long.

Type: On ledges of limestone cliff along highway above Tama-
zunchale, San Luis Potosi, Mexico, altitude 175 meters, in flower and
fruit, July, 1937, C. L. & A. A. Lundell 7231 (MICH). Known only
from this locality.

This species is well marked by the winged stems, pubescent on
the margins of the wings, the long aristate calyx lobes, covered with
resinous scales, and the small, distinctly bilabiate, maculose corollas.

Mexico, San Luis Potosi: Tamazunchale, Kenoyer A247 (F),
Lundell & Lundell 7231 (LUND, MICH, PA, SMU), 12166 (LUND,
SMU), 12257 (LUND, SMU), 12491 (LUND, US), 12652 (LUND,
SMU).

41. Russelia retrorsa E. L. Greene, Pittonia 1 : 176. 1887-1889.
R. trachypleura Robinson, Proc. Amer. Acad. 36: 474. 1901.

Plants suffrutescent, branching, 1.2-1.6 meters tall; stems 4-8-angled, angles
prominent, narrow, with ridges usually roughened by swollen bases of hairs which
often become callosities, the hairs 3-9-celled, 0.6-0.7 mm. long, areas between the
ridges flat, with striations and usually with hairs, the hairs usually numerous but
sometimes few and scattered, usually straight but sometimes retro rse; internodes
3-8 cm. long; leaves opposite or ternate, ovate or elliptic-ovate, 2-6 cm. long,

276 FIELDIANA: BOTANY, VOLUME 29

1-3.5 cm. broad, apices acute or obtuse, bases rounded or cuneate and entire,
serrate or crenate above base, both surfaces densely pilose, upper and sometimes
lower with large resinous scales, veins 3-4 on each side of midrib, not strikingly
prominent on lower side, densely pilose; petioles 1-2 mm. long, pilose; inflo-
rescences axillary, cymose, short, with 3-7 flowers; bracts similar to leaves but
smaller; bractlets linear, pubescent; peduncles and pedicels 3-7 mm. long, pu-
bescent; calyx lobes 2.5-3 mm. long, ovate, acuminate, margins membranaceous,
few hairs along the midrib or sometimes lacking; corollas bright scarlet, tubular,
10-12 mm. long, upper lip deeply notched, lobes of lower lip rounded, 2 mm. long,
barbate on inner surface of median lobe near throat and near attachment of
stamens; stamens 6 and 7 mm. long, small hairs at base of filaments, anthers ovoid,
1 mm. long, divaricate; staminodes 1 mm. long; pistils 9-10 mm. long; capsules
ovoid, 3-5 mm. long, about as long as the persistent calyx lobes; seeds 0.3 mm.
long, ovoid, brown.

Type: Rio Blanco, near Guadalajara, Jalisco, Mexico, September,
1886, E. Palmer 540 (US).

The species grows from 1200-2400 meters altitude, and blooms
throughout the year. It is found in the west-central part of Mexico
(Map 3).

Mexico, Guanajuato: Near Guanajuato, Duges s.n. (GH), 28
(GH), 376 (GH). Mexico: Mexico, Valle de Bravo, Carlson 3012
(F), Lundell 12646 (LUND); Los Hornos, Temascaltepec, Hinton
2111 (GH, LUND, NY, PA), 2112 (F, GH, PA, US); near Ozumba,
Pringle 8712 (F, GH, NY, PA, UC, US), 13152 (F, GH, US).
Morelos: south of El Parque, Morelos, Langman 2783 (PA); Sierra
Chalchi, southwest of Tepoztlan, Langman 3693 (PA); Cuautla,
Lyonnet 369 (GH, MO, NY, US) ; Sierra de Tepoztlan, Pringle 8445
(F, GH, MEXU, MO, NY, PA, UC, US); Barranca of Cuernavaca,
Pringle 11069 (F, GH, MO, NY, US); Casa del Tezoztero, Seler 43B
(F, GH). Jalisco: Etzatlan, along road to San Domingo Mine,
Barnes & Land 302 (F, PA) ; km. 41, Guadalajara to Autlan, Lang-
man 3127 (MEXU, PA); Rio Blanco, near Guadalajara, Palmer 540
(GH, NY, PA, US); near Guadalajara, Pringle 2149 (GH), 9540
(F, GH, MEXU, MO, US); Tuxpan, Pringle 13400 (GH, US),
Purpus 514 (UC). Michoacan: Rincon, Morelia, Arsene s.n. (US);
Coalcoman, Hinton 12892 (F, GH, LUND, MO, PA, US); Zitacuaro,
Hinton 13197 (GH, LUND, US); Tancitaro, Uruapan, Hinton 15620
(GH, LUND, NY, US) ; between Uruapan and Jumpimito, Langman
3298 (PA) . Guerrero : environs of Taxco, Abbott 1 21 (GH) ; Teotepec,
Galeana, Hinton 11105 (GH, US); between Ayusinapa and Petatlan,
Nelson 2143 (US); Chilpancingo, Sharp 441460 (PA). Durango:
Sianori, Ortega 5340 (US); Sierra Madre, Rose 3506 (US). Sinaloa:
Mazatlan, Brandegee s.n. (UC); Quebrado de Mansana, Sierra

CARLSON: THE GENUS RUSSELIA 277

Surotato, Gentry 6484 (GH, LUND, MICH, MO, NY) ; Mazatlan,
Ortega 5658 (MEXU). Colima: Manzanillo, West 3505 (GH, MO,
US), Orcutt 4477 (F, GH, MO). Baja California: Sierra San Lazaro,
Brandegee s.n. (GH, UC, US) ; Sierra de La Laguna, Hammerly 278
(GH); Laguna Mts., Jones 24429 (MO, NY), 24430 (F, UC), 27409
(MO). Chiapas: Comitan, Langman 3740 (PA). Location not
given: Keating s.n. (PA); Pringle s.n. (MO); Ortega 88 (MEXU).

42. Russelia retrorsa f. nudicostata Carlson, forma nov.

A var. retrorsa recedit caulium costis glabris 0.5 mm. latis, laminis glabris vel
minute pubescentibus.

Similar to the species except that the angles of the stems are broad (0.5 mm.)
and glabrous, and the leaves are glabrous or minutely pubescent.

Type: Sierra de La Laguna, Lower California, Mexico, January
26, 1890, T. S. Brandegee 428 (US).

Mexico, Lower California: Sierra de La Laguna, Brandegee 428
(GH, PA, UC, US); Sierra San Lazaro, Brandegee s.n. (GH); San
Jos4 del Cabo, Purpus 205 (MO, UC, US).

43. Russelia obtusata Blake, Proc. Biol. Soc. Wash. 33:
119. 1920.

Plants suffrutescent below, creeping base, erect stems, simple or with opposite
branches, 5 meters tall, stout, 5 mm. thick at base; stems pale green, 6-8-angled,
with broad flattened ridges on the angles, densely and microscopically pubescent;
internodes 6-8 cm. long; leaves in three's or four's, papery or pergamentaceous,
1-3 cm. long, 1-3 cm. broad, ovate, obovate or suborbicular, apex acute or obtuse,
base broadly rounded to cuneate, crenate-serrate, 4-8 pairs of teeth, acute or
obtuse, upper surface deep green, glabrous or sparsely incurved tomentose, lower
paler green, sparsely incurved tomentose along the costa and 3-4 pairs of promi-
nent veins, pitted, no resinous dots either side; petioles slender, 2-8 mm. long, with
densely spreading or incurved very short hairs; inflorescences axillary, 3-5-
flowered; bractlets linear, tomentose; peduncles 4 mm. long to obsolete; pedicels
1.5-6 mm. long, densely and finely tomentose; calyx lobes 2.5 mm. long, ovate or
oval-ovate, abruptly short-pointed or acuminate, not striate, glabrous or slightly
hairy along the midrib, thick, with rosy margins; corollas red, 13-16 mm. long,
glabrous outside, upper lip emarginate, lower lip with 3 equal lobes, crenulate,
3 mm. long, 2 mm. wide, pilose with one-celled hairs along inside of median lower
lobe and near the attachment of stamens; stamens didynamous, 8 and 9 mm.
long, pilose at base; anthers ovoid, divaricate; staminodes 1.3 mm. long; pistils 10
mm. long; capsules subglobose, 4 mm. broad, tipped with persistent style, 7 mm.
long; seeds ovoid, light brown, 0.5-0.7 mm. long, surface rough.

Type: Near Tehuacan, Puebla, Mexico, altitude 1500-2000
meters, August 30-September 8, 1920, J. N. Rose, J. H. Painter &
J. S. Rose 10026 (US).

278 FIELDIANA: BOTANY, VOLUME 29

This taxon is readily recognized by the broad, somewhat flattened
ridges on the angles and the dense covering of microscopic hairs on
all parts except the calyx and corolla. It is related to R. polyedra
and was so named by Karwinski (1094; see p. 283).

Mexico, Puebla: near Tehuacan, Pringle 9621 (GH); Barranca de
Thalcuilosto, Purpus s.n. (US); vicinity of San Luis Tultitlanapa,
Purpus 3270 (F, GH, MO, NY, UC, US); Chiltepin, near San Luis
Tultitlanapa, Purpus 3270a (MO, US) ; Tlacuisloltepec, Purpus 3952
(UC); near El Riego, Tehuacan, Rose, Painter & Rose 10026 (NY,
US). Oaxaca: San Felipe, near Oaxaca, Conzatti s.n. (NY); Monte
Alban, Conzatti & Gonzalez 1068 (GH), Smith 154 (MO), 400 (MO);
Barranca de San Felipe, Conzatti 5275 (MICH, PA); Natividad
Road, Kenoyer 1578 (GH); Monte Alban, Lundell 12285 (LUND,
SMU) ; San Miguel Alborrados, Nelson 538 (US) ; Valley of Oaxaca,
Nelson 1202 (US), 1293 (US); Jazacutlon, Rusby s.n. (NY); Totala-
pan, Seler 1747 (GH).

44. Russelia pringlei Robinson, Proc. Amer. Acad. 43 : 26. 1907.

Plants suffrutescent, 1-2 meters tall, branching; stems appearing terete but
with 6-8 broad flat ridges, 8 mm. in diameter at base, gray, canescent, villous,
with matted, white, unicellular hairs (0.09 mm. long); internodes 5-6 cm. long;
leaves opposite, ternate or occasionally in whorls of 4, 1.5-3.5 cm. long, 1-3 cm.
wide, ovate or ovate-lanceolate, apices acute, bases truncate or subcordate, upper
one-fourth or one- third serrate-dentate, with 2-4 widely spaced teeth; upper
leaves often entire; upper surfaces with sparse long hairs between the veins, lower
surfaces with more hairs, especially on the veins, both surfaces with scattered
glandular dots (0.2 mm. in diameter), more on the lower than on the upper surface;
main veins impressed on upper, prominent on lower side; petioles 1-2 mm. long,
villous; inflorescences axillary, spreading, cymose, 1-3 cm. long, with 4-12 flowers;
bracts like the leaves except smaller and with fewer or no teeth; bractlets linear,
densely hairy; peduncles slender, 4-10 mm. long, villous, without glandular dots;
pedicels 4-5 mm. long, villous, without glandular dots; calyx lobes 5 mm. long,
ovate-lanceolate, caudate, with a few hairs and numerous crowded glandular dots
along the midrib; corollas red, 14-17 mm. long, glabrous externally, upper lip
deeply notched (0.8 mm. deep), lower lobes 2 mm. long, 1 mm. wide, bearded
with numerous flat hairs on median lower lobe and at base near attachment of
stamens; stamens 10 and 12 mm. long, extending to throat of corolla tube; anthers
oval, 0.6-0.7 mm. long, divaricate; staminodes 0.8 mm. long; pistils 12 mm. long;
capsules ovoid, 4-6 mm. long; style persistent, 10 mm. long; persistent calyx
extending almost to top of capsule; seeds black, 0.54 mm. long, 0.3 mm. wide,
reticulate on surface.

Type: Iguala Canyon, Iguala, Guerrero, Mexico, altitude 760
meters, December 28, 1906, C. G. Pringle 10367 (GH).

Known from only three collections on vertical limestone walls of
Iguala Canyon (Canyon de la Mano), 6.5 miles north of Iguala. In
flower and fruit September to January.

CARLSON: THE GENUS RUSSELIA 279

Mexico, Guerrero: Canyon de la Mano, Carlson 3132 (F), Piingle
10148 (GH, LUND, US), 10367 (F, GH, MO, NY, PA, UC, US).

45. Russelia villosa Lundell, Field and Lab. 13: 21. 1945.

Plants suffrutescent, 1.5-2 meters tall, branching; stems thick, bases up to
1.2 cm. in diameter, 4-8-angled, angles with prominent ridges, copiously white-
villous between the ridges and somewhat on the ridges, without glandular dots;
internodes 10-12 mm. long; leaves opposite or ternate, 6-10 cm. long, 4-5 cm.
wide, broadly ovate, apices acute but not sharply so, or obtuse, bases cordate or
truncate, often broadly cuneate, margins serrate-dentate, persistently villous on
both surfaces, paler beneath, many irregular resinous dots on the lower surface,
fewer on the upper; basal leaves reduced, sometimes only 1 cm. long, suborbicular;
veins 5-7 on each side of the midrib, prominent and lighter colored on the lower
side; petioles short (1-2 mm. long), woolly, no glandular dots; inflorescences near
tops of stems, axillary, compact, with 5-10 flowers; bractlets linear, woolly;
peduncles very short or lacking, woolly; pedicels 2.5-5 mm. long, woolly; calyx
lobes 3 mm. long, ovate, with membranaceous margins, acuminate, keeled, short
ciliate, with a few hairs on the keel, no glandular dots; corollas rose-red, 7-10 mm.
long, externally glabrous, lobes slightly spreading, lower lobes up to 2 mm. long,
upper emarginate, hairs at throat and along inside of lower median lobe, and near
attachment of stamens; stamens 4 and 5 mm. long, hairs at base; anthers oval,
divaricate; staminodes minute; pistils 7 mm. long; capsules globose, 4-5 mm. long,
basal half covered by persistent calyx; seeds small, black.

Type: Mountain side near Taxco, Guerrero, Mexico, October 19,
1943, C.L.&A.A. Lundell 12568 (SMU).

This species is very similar to R. ternifolia H.B.K. and may be
a variety of it. The specimens from near Taxco have long, matted
hairs between, but not so much on, the ridges; short peduncles, so
short that the inflorescences appear umbelliferous; resinous dots on
both sides of the leaves; few hairs on the calyx lobes.

Mexico, Guerrero: near Taxco, Abbott 63 (GH), Carlson 3111 (F),
3117 (F), Lundell & Lundell 12568 (GH, LUND, MEXU, MICH,
SMU, US); Suriana, Mexia 8827 (F, GH, MO, NY, US). Michoa-
can: Apatzingan, Leavenworth & Hoogslraal s.n. (F); Jorullo, Nelson
6947 (US). Mexico: Tamascaltepec, Hinton 3191 (GH, US), 7297
(GH).

46. Russelia ternifolia H.B.K. Nov. Gen. et Sp. 2: 359. 1817.

Plants with stems and branches 4-6-angled, the angles slightly thickened,
densely hirsute-pubescent; internodes 5-10 cm. long; leaves opposite or ternate,
5-6.2 cm. long, 3.3-5.2 cm. wide, either obovate and rounded at the tip or ovate
and obtuse at the tip, bases truncate or cuneate and decurrent on the petiole,
margins coarsely serrate or serrate-crenate except at the base, rigid-membran-
aceous, hirsute on both surfaces, especially on the veins, not resinous-lepidote,
changing to purple, veins reticulate, with the midrib and primary veins prominent

280 FIELDIANA: BOTANY, VOLUME 29

on the lower side; petioles 4-5 mm. long, hirsute; inflorescences cymose, axillary,
solitary, many-flowered; bractlets linear, hirsute- tomentose; peduncles 5-7 mm.
long, hirsute- tomentose; pedicels 4 mm. long, filiform, tomentose; calyx lobes
2.5-3.5 mm. long, ovate, mucronate, margins incumbent, puberulous; corollas
8-10 mm. long, slightly wider at the top, externally glabrous, limb 4-lobed, the
upper a little wider than the lower and slightly notched, the lobes rounded; inner
surface of the lower lip barbate, stamens included, inserted at the base of the
corolla, with hairs on the inner surface; filaments filiform, dilated at the base;
anthers orbicular; ovaries oblong, glabrous; styles glabrous, scarcely equaling the
stamens; capsules globose, 3 mm. long; seeds 0.4-0.5 mm. long, black.

Type: From declivity of Volcano Jorullo, Michoacan, Mexico,
altitude 1100-1300 meters, flowering in September, Kunth s.n. (Herb.
Mus. Paris).

Only a photograph of the type specimen was seen.

Mexico, Oaxaca: Ixtepec, Fisher 35323 (F, MO, NY, US);
Cerro de Picacho, Purpus 7288 (GH, MO, NY, UC, US); Salina
Cruz, Deam s.n. (GH). Chiapas: Sierra de Tonala, Purpus 6882
(F, GH, MO, NY, UC, US). Mexico: Ixtapan, Hinton 1167 (F,
GH, MO, PA, US).

47. Russelia conzattii Carlson, sp. nov. Plate 5.

Planta suffruticosa; ramis 4-6-angularibus, angulis incrassatis striatis pubes-
centibus; foliis oppositis vel ternatis; laminis 8 cm. longis, 5 cm. latis, ovato-
lanceolatis, apice acutis, basi truncatis vel cordatis, sessilibus vel subsessilibus
pubescentibus; petiolis 1 mm. longis vel nullis; inflorescentiis multifloris; bracteolis
linearibus pubescentibus; pedunculis 15-20 mm. longis pubescentibus; pedicellis
8-10 mm. longis pubescentibus; sepalis 2.5-3 mm. longis ovatis acuminatis parce
pubescentibus; corollis 12-14 mm. longis; staminibus 8 vel 9 mm. longis; stamin-
odiis 0.5 mm. longis; pistillis 10 mm. longis.

Plants suffruticose; stems 4-6-angled, angles slightly thickened, flat sides
striate, both angles and flat sides pubescent; leaves opposite or ternate, sessile or
subsessile; blades 8 cm. long, 5 cm. wide, ovate-lanceolate, apices acute, bases
truncate or cordate, pubescent, especially on the veins; petioles 1 mm. long or
lacking, pubescent; inflorescences many-flowered; bractlets linear, pubescent;
peduncles 15-20 mm. long, pubescent; pedicels 8-10 mm. long, pubescent; calyx
lobes 2.5-3 mm. long, ovate, acuminate, slightly pubescent along costa; corollas
12-14 mm. long, ventral lobes 2 mm. long, 1.5 mm. wide, dorsal lip slightly
emarginate, internally barbate along median ventral lobe and at base near in-
sertion of stamens; stamens 8 and 9 mm. long; staminodes 0.5 mm. long; pistils
10 mm. long; no mature capsules present on specimen.

Type: Near Panixllahuaca, Dist. Tuquila, Oaxaca, Mexico,
altitude 850 meters, December 25, 1921, C. Conzatti 4525 (US).

Mexico, Oaxaca: Panixllahuaca, Conzatti 4525 (US); Cerro de
Chacalina, Conzatti 4478 (US).

CARLSON: THE GENUS RUSSELIA 281

48. Russelia grandidentata Carlson, sp. nov. Plate 5.

Planta suffruticosa; ramis 4-6-angularibus pilosulis striatis, angulis in-
crassatis 0.35 mm. latis; foliis oppositis vel ternatis; laminis inferioribus ovatis
vel deltoideis, 3.5 cm. longis, 3 cm. latis, apice acutis, basi truncatis, parce breviter
pubescentibus, grosse dentatis, dentibus 5-7 mm. longis; petiolis 4-5 mm. longis
breviter pubescentibus; inflorescentiis 3-5-floris; pedunculis 3 mm. longis dense
pilosulis; pedicellis 3-4 mm. longis, dense pilosulis; sepalis ovatis acuminatis,

3 mm. longis, costis breviter pubescentibus; corollis 15-17 mm. longis; capsulis
globosis, 4 mm. longis; seminibus ovatis.

Plants suffruticose; stems 4-6-angled, ridges on the angles 0.35 mm. wide,
striations between the angles, both ridges and flat sides with soft hairs of irregular
lengths (0.05-0.4 mm. long, unicellular); leaves opposite or ternate; blades of
lower leaves ovate or deltoid, 3.5 cm. long, 3 cm. wide, slightly short-pubescent,
often caducous, lobed or large-dentate, teeth 5-7 mm. long, 3-4 on each side, upper
leaves much smaller, resinous-lepidote on both sides; petioles 4-5 mm. long,
short-pubescent; inflorescences 3-5-flowered; peduncles 3 mm. long, densely
short-pubescent; pedicels 3-4 mm. long, densely short-pubescent; calyx lobes
ovate, acuminate, 3 mm. long, pubescent along costa; corollas 15-17 mm. long,
upper lip emarginate, lower lobes 2 mm. deep, 1 mm. wide, barbate within along
median ventral lobe and at base near insertion of stamens; stamens 9 and 11 mm.
long; anthers divaricate; staminodes 1.5-2 mm. long; pistils 10 mm. long; capsules

4 mm. long, beaked; seeds brown, oval.

Type: Vinorama, east of La Paz, Baja California, Mexico, April
1, 1949, Annetta Carter 2621 (UC).

Mexico, Lower California: Vinorama, Carter 2621 (PA, UC),
2687 (PA).

49. Russelia hintoni Lundell, Field and Lab. 13: 14. 1945.

Plants suffruticose, 2 meters tall; stems and branches 4-angled, persistently
hirtellous; leaves thin, membranaceous, opposite, 2.5-6 cm. long, 2-4.5 cm. wide,
ovate, ovate-elliptic or elliptic, apices acute or obtuse, bases truncate or rounded,
margins coarsely serrate-dentate, basal third subentire, sparsely hairy on upper
surface, more densely hairy on lower, especially on the veins, resinous-lepidote on
both surfaces, sometimes sparingly so, costa and principal veins slightly impressed
above, elevated beneath, 4-5 pairs on each side; petioles slender, 2-5 mm. long,
hirtellous; inflorescences laxly cymose, axillary, up to 4 cm. long, few- to many-
flowered; bractlets linear, hirtellous; peduncles 4-6 mm. long, densely hirtellous;
pedicels slender, 4-8 mm. long, hirtellous; calyx lobes ovate at base, abruptly
cuspidate or long-acuminate, tip sometimes recurved, 3 mm. long, sparsely
pubescent along costa, with an occasional resinous dot; corollas tubular, slightly
narrowed below, 10-12 mm. long, externally glabrous, barbate within on lower
side and at base near attachment of stamens; stamens 7 and 9 mm. long, barbate
at base on inner side; anthers ovoid, divaricate; staminodes 0.3 mm. long; pistils
8 mm. long; capsules 3 mm. long, globose; seeds numerous, small.

Type: Near Galeana, Guerrero, Mexico, altitude 400 meters,
November 14, 1937, G. B. Hinton 10921 (SMU).

282 FIELDIANA: BOTANY, VOLUME 29

Lundell says: "From R. cuneata Rob., to which it is related, R.
hintoni differs in being densely pubescent and in having leaves obtuse
or rounded at the base, not cuneate-attenuate."

Mexico, Guerrero: Atoyac, Galeana, Hinton 10921 (GH, LUND,

NY, SMU).

50. Russelia parvifolia Carlson, sp. nov. Plate 6.

Planta suffruticosa, 1-1.5 m. alta; ramis dense hirsutis, 4-6-angularibus,
angulis incrassatis; foliis oppositis vel ternatis, petiolatis; petiolis 2-4 mm. longis,
dense hirsutis; laminis 1-2 cm. longis, 0.8-1.5 cm. latis, ovatis, apice acutis vel
obtusis, basi truncatis, serratis vel serrato-crenatis, utrinque dense hirsutis,
minute resinoso-lepidotis; inflorescentiis cymosis, 3-8-floris; pedunculis 4-8 mm.
longis, dense hirsutis; pedicellis 3-4 mm. longis, dense hirsutis; sepalis 3 mm.
longis, ovatis, subulatis, pubescentibus, parce resinoso-lepidotis; corollis coccineis,
11-12 mm. longis, tubulosis; filamentis 7 vel 9 mm. longis; antheris ovalibus,
divaricatis; pistillis 8 mm. longis; capsulis globosis, 3 mm. longis; seminibus
numerosis, parvis.

Plants suffruticose, 1-1.5 meters tall, branching; stems 4-6-angled, ridges on
angles thickened, broad and flat, densely hirsute with straight multicellular hairs
of unequal lengths; internodes 3-8 cm. long; leaves opposite or ternate, 1-2 cm.
long, 0.8-1.5 cm. wide, broadly ovate or elliptic, apices acute or obtuse, bases
truncate or widely cuneate, margins serrate or serrate-crenate, densely hirsute on
both surfaces and especially on the veins on the lower side, scattered resinous-
lepidote on both sides; petioles 2-4 mm. long, hirsute; inflorescences spreading,
with 3-8 flowers; peduncles 4-8 mm. long, pedicels 3-4 mm. long, both densely
hirsute and glandless; calyx lobes 3 mm. long, ovate, pubescent medianly and on
the subulate tip, densely or sparingly resinous-lepidote; corollas tubular, slightly
spreading at the mouth, 11-12 mm. long, ventral lobes 3 mm. long, 1 mm. wide,
dorsal lip shorter, emarginate; corollas glabrous externally, bearded inside along
median ventral lobe and at base near attachment of stamens; stamens 7 and 9 mm.
long, with hairs at base of filaments; anthers oval, divaricate; staminodes 1 mm.
long; pistils 8 mm. long; capsules 3 mm. long, globose; seeds numerous, minute.

Type: Sides of ravine, near Fiscal, Guatemala, Guatemala,
altitude 110 meters, May 31, 1909, C. C. Beam 6069 (MICH).

R. parvifolia differs from polyedra in having shorter corollas,
not funnelform, but merely wider at the top; longer stamens and
pistils; and more dense pubescence on the vegetative parts. The
taxon was recognized but not described by F. W. Pennell.

Guatemala, Guatemala: Fiscal, Beam 6069 (GH, MICH, PA),
Standley 59584 (F, PA), 80371 (F, PA, US), 80390 (F), 80427 (F);
Agua Caliente, Greenman & Greenman 5899 (MO). Zacapa: Rio
Hondo, Standley 73994 (PA), Steyermark 29464 (F), 42917 (F, PA).
Baja Vera Paz: Santa Rosa, Von Tuerckheim 1198 (GH, PA, NY,
US).

CARLSON: THE GENUS RUSSELIA 283

51. Russelia polyedra Zuccarini, Abh. Akad. Wiss. Muenchen
2: 328. 1831-36. R. pennelliana Lundell, Contr. Univ. Mich. Herb.
6: 58. 1941. R. pennelliana var. pilosa Lundell, Field and Lab. 13:
17. 1945.

Plants fruticose, 1-1.5 meters tall; stems 6-10-angled, with broad, flattened
ridges on the angles, the faces between the angles flattened or concave, striate;
thinly or densely velutinous, especially on the ridges, or glabrous, i.e., some
branches may be very hairy while others on the same plant may be glabrous,
internodes 5-7.5 cm. long; leaves opposite or verticillate, 1.2-2.5 cm. long, 1.2-2
cm. wide, suborbicular or broadly ovate, apices obtuse, bases rounded and entire,
margins deeply and irregularly serrate-dentate, teeth obtuse, softly pubescent or
glabrous, resinous-lepidote on both sides; petioles 6-8 mm. long, slender, widened
at base, softly pubescent; inflorescences in axils of upper leaves, 2-5-flowered;
bractlets linear, pubescent; peduncles and pedicels 6-8 mm. long, slender, pubes-
cent, resinous-lepidote; calyx lobes ovate, 4 mm. long, acute, pubescent, resinous-
lepidote, margins membranaceous at base; corollas red, funnelform, 15-20 mm.
long, lobes decidedly spreading, somewhat undulate, ventral lobes 4.5 mm. long,
5 mm. wide, dorsal slightly shorter, emarginate, cleft 1 mm. deep; corollas ex-
ternally glabrous, internally barbate with erect yellow hairs along lower lip and
at base near attachment of stamens; stamens 5 and 7 mm. long; anthers oval,
divaricate; staminodes 1 mm. long, filiform; pistils 7 mm. long; capsules 5 mm.
long; seeds oval, 0.8 mm. long.

Type: No locality in Mexico is given for type specimen, Kar-
winski 1093 (Munich).

Two specimens, Karwinski 1093 and 1094, were sent to me from
Munich as the type of this taxon. These two are combined in
a photograph (8657, F) which was distributed as the type. They
represent two collections and, although similar, they are not alike.
One (1094} is densely tomentose, with microscopic hairs which are
barely visible to the naked eye, and has no resinous dots on any
part. This is the readily recognized taxon which was described by
Blake as R. obtusala and is treated as a valid species in the present
work. The other specimen (1093} fits the description and is desig-
nated as the type of R. polyedra.

This species has apparently not been correctly defined in the
past, as many specimens which have been identified as R. polyedra
are definitely not polyedra. The type specimen has the striking,
large, funnelform corollas which are characteristic of the specimens
named by Lundell as R. pennelliana and R. pennelliana var. pilosa.
Therefore, these are reduced to synonymy. The type specimen is
definitely pilose, but the specimens referred to this species may
have both pilose and perfectly glabrous branches on the same plant
or on plants of the same collection. It seems impossible, therefore,

284 FIELDIANA: BOTANY, VOLUME 29

to separate one as a variety of the other. However, those specimens
from the vicinity of Ciudad Victoria are generally pubescent, while
those from farther east (Jaumave) are generally glabrous (Map 3).

As noted by Lundell, this species is worthy of cultivation.
Calyces and corollas with 6 lobes and 5 or 6 stamens occur.

Mexico, Tamaulipas: Cerro de la Tamaulipeca, near San Migue"!,
Bartlett 10542 (F, US); La Morita, Marmolejo, Bartlett 10776 (F);
13 miles south of Ciudad Victoria, Hitchcock & Stanford 6886 (GH,
MO, NY, UC); Chamal, Kenoyer & Crum 3647 (MICH); Mesa de
Llera, Lundell & Lundell 7288 (LUND, MICH); Cuesta de Llera,
Lundell & Lundell 12170, 12487, 12487 A, 12489 (LUND); Rio San
Marcos, 5 miles west of Ciudad Victoria, Meyer & Rogers 2490
(MO, PA); Ciudad Victoria, Foster 900 (LUND), Palmer 259 (US);
San Vicente, Jaumave, von Rozynski 41a (MICH, UC); Jaumave,
von Rozynski 41 (F, GH) ; Nogales, Jaumave, von Rozynski 485 (F,
PA) ; 30 km. southwest of Victoria, 3 km. north of Huisachal, Stan-
ford, Lauber & Taylor 2036 (PA) ; 19 km. southeast of Miquihuana
on road to Palmillas, Stanford, Retherford & Norther aft 826 (GH,
MO, NY, UC) ; Jaumave, Viereck 482 (US) ; Yerba del Suena, near
San Vicente, Woolen s.n. (US). Guanajuato: Xichu mine, Kenoyer
2261 (GH). Hidalgo: Jacala, Kenoyer s.n. (MO); west slopes of
mountain near km. 278, on highway north of Jacala, Moore 1791
(GH). No location given, Purpus s.n. (UC).

52. Russelia staleyae Carlson, sp. nov. Plate 6.

Planta suffruticosa; ramis 4-6-angularibus, angulis incrassatis striatis tomen-
tosis; foliis oppositis vel ternatis; laminis 12 mm. longis, 10 mm. latis, orbiculatis,
apice obtusis, basi obtusis, crenatis, dense tomentosis utrinque resinoso-lepidotis;
petiolis 1 mm. longis, tomentosis; inflorescentiis multifloris; pedunculis 3-5 mm.
longis, tomentosis; pedicellis 3-4 mm. longis, tomentosis; sepalis 3 mm. longis,
ovatis acuminatis parce pubescentibus; corollis 11-12 mm. longis, cornutis lobis
grandibus; filamentis 3 vel 4 mm. longis, basi barbatis; staminodiis 1.5 mm. lon-
gis; pistillis 4 mm. longis; capsulis 3 mm. longis, globosis; seminibus minutis.

Plants suffruticose; stems 4-6-angled, angles thickened, striate on the flat sides,
both angles and flat sides densely tomentose; leaves opposite or ternate; blades
12 mm. long, 10 mm. wide, orbicular, apices obtuse, bases obtuse, crenate,
densely tomentose, both sides resinous-lepidote; petioles 1 mm. long, tomen-
tose; inflorescences many-flowered; peduncles 3-5 mm. long, tomentose; pedicels
3-4 mm. long, tomentose; calyx lobes 3 mm. long, ovate, acuminate, sparsely
pubescent; corollas 11-12 mm. long, funnelform, the lobes large (3 mm. long, 3
mm. wide) and spreading, the dorsal lip slightly emarginate, slightly barbate
along the median ventral lobe, densely barbate at base; stamens 3 and 4 mm.
long, with a few large hairs at the base; staminodes 1.5 mm. long; pistils 4 mm.
long; capsules 3 mm. long, globose; seeds minute.

CARLSON: THE GENUS RUSSELIA 285

Type: Cliff near Pan-American Highway, just beyond the road
to Juchitan, Oaxaca, Mexico, March 22, 1949, M. C. Carlson 1494
(F).

This taxon is similar to R. parvifolia, but the flowers are smaller
and more funnelform. I take pleasure in naming it for Miss Kate
Staley, my companion on several collecting trips in Mexico and
Central America, who discovered it.

Mexico, Oaxaca: Near Juchitan, Carlson 1494 (F); Tehuantepe"c,
Orcutt 6458 (US).

Excluded Species

R. alata Cham. & Schl. Linnaea 3: 3. 1828=Anetanthus Hiern. Benth. &
Hooker, Gen. PI. 2: 941. 1873-1876.

R. alternifolia Pers. Pers. Syn. 2: 164. 1807. Not Russelia, as the description
states, "with alternate leaves and solitary flowers."

R. americana E. A. Raeuschel, Nom. Bot. Lipsiae. ed. III. 180. 1797=sar-
mentosa Jacq. Kew Index 2: 767. 1895.

R. capensis L. f. Carolus von Linne f. Suppl. PI. 175. 1781 = Vahlia capensis
Thunb. fide De Candolle Prodromus 4: 53. 1830.

R. depressa Thunb. Flor. Capens. 479-480. 1823 is Ruellia depressa Thunb.,
misquoted by Lindley, Bot. Reg. 22 (n.s. 9): 1882. 1836.

R. flammea Raf. C. S. Rafinesque, New Fl. N. Amer. 2: 71. 1837=Macran-
thera flammea (Bartram) Pennell. F. W. Pennell, Studies in the Agalinanae,
a subtribe of the Rhinanthaceae II. Bull. Torrey Bot. Club 40: 405. 1913.

R. peruviana Spruce, PI. Exs. n. 4515=Verbenaceae. Benth. & Hooker,
Gen. PI. 2: 941. 1873-1876.

R. racemosa (Ruiz & Pav.) Wetts. Engl. & Prantl, Natiir. Pflanzenf. IV. 3b:
63. 1891 = Gomara racemosa Ruiz. & Pav. H. Ruiz & J. Pavon, Syst. Veg. 1:
162. 1798.

Exsiccatae

Species numbers in parentheses. Other numbers are collectors' numbers.

Abbott, R. Q. 63 (45), 121 (41).

Aguilar, M. 68 (25), 194 (24), 464 (4).

Aguirre, A. & Reko, B. 312 (10).

Allen, P. H. 92 (22), 1081 (22), 1156 (22), 1582 (23), 2770 (22), 5624 (22).

Arsene, G. s. n. (10), (11), (12), (41).

Auber, A. 13 (22).

Bailey, L. H. 604 (11).

Baker, C. F. 101 (26), 197 (26), 580 (26), 880 (26), 2168 (26), 2215 (26).

Balls, E. R. 4272 (32).

Barclay, G. W. s. n. (14).

Barnes, C. R., Chamberlain, C. J., & Land, W. J. G. 47 (10).

Barnes, C. R. & Land, W. J. G. 302 (41), 660 (10).

286 FIELDIANA: BOTANY, VOLUME 29

Bartlett, H. H. 10542 (51), 10776 (51), 11516 (22), 11531 (22), 11716 (22), 11999

(4), 12263 (24), 12665 (24), 12852 (4).
Bartlett, H. H. & Lasser, T. 16349 (28).
Basil, H. B. s. n. (22).
Bigelow, J. s. n. (26).
Biolley, P. 71 (24), 17331 (22).
Bonner, J. 31 (5), 37 (5).
Botteri, M. 28 (10), 123 (32), 293 (32).
Bourgeau, M. 1983 (10), 2163 (32).
Brandegee, F. S. s. n. (39), (41), (42); 428 (42).
Brenes, A. M. 168 (26), 232 (22), 532 (22), 803 (22), 12696 (24), 14408 (22),

17004 (22).
Broadway, W. E. 8006 (10).

Calderon, S. 184 (22), 827 (22).

Carleton, M. A. 462 (24).

Carlson, M. C. s. n. (5), (10); 24 (22), 170 (22), 789 (22), 1305 (10), 1494 (52),
2762 (5), 2779 (10), 2781 (5), 2782 (10), 2787 (10), 3012 (41), 3032 (43),
3049 (17), 3079 (17), 3081 (17), 3108 (16), 3111 (45), 3117 (45), 3119 (12),
3130 (10), 3132 (44).

Carter, A. 2621 (48), 2687 (48).

Celestino, Bro. 53 (22).

Chanek, M. 231 (4).

Chaves, D. 63 (26).

Choussey, F. 13 (22).

Chun, W. Y. 6982 (10).

Collins, G. N. & Kempton, J. H. 21 (14).

Conzatti, C. s. n. (43); 835 (10), 4478 (47), 4525 (47), 5275 (43).

Conzatti, C. & Gonzalez, V. 1068 (43).

Cook, O. F. & Doyle, C. B. 21 (22).

Cook, O. F. & Griggs, R. F. 305 (24).

Cooper, G. P. 39 (22).

Cornman, L. R. 2025 (22).

Curtiss, A. H. 753 (22).

Davidson, M. E. 472 (23).

Deam, C. s. n. (5), (11), (12), (27), (46); 41 (5), 183 (35), 6069 (50), 6335 (35).

Diquet, L. s. n. (11), (12), (13), (14).

Dodge, C. W., Alfaro, A. & Thomas, W. S. 6245 (22).

Dodge, C. W. & Allen, P. H. 17324 (22).

Dodge, C. W., Steyermark, J. A. & Allen, P. H. 16967 (22).

Dodge, C. W. & Thomas, W. S. 6223 (24), 6223a (24), 6434 (22).

Drouet, F. & Richards, D. 3943 (31).

Duges, A. s. n. (41); 28 (41), 376 (41).

Duss, Pere 1348 (10).

Edwards, M. T. 816 (30).
Elmore, F. H. Ell (22).
Emrick, G. M. 177 (15), 181 (6).
Ervendberg, L. C. 223 (10), 263 (10).

Ferris, R. S. 5416 (14), 5614 (21), 6082 (37).
Fisher, G. L. 35323 (46), 45184 (30).
Foster, M. B. 900 (51).

Galeottii, H. 1109s (32).

Gamier, A. s. n. (22); 58 (26), 267 (26), 4297 (26).

Gaumer, G. F. 403 (22), 1746 (22).

Gaumer, G. F. & sons 23660 (22).

Gentle, P. H. 385 (4), 647 (4), 952 (22), 1140 (4), 2175 (4), 2248 (22), 2269 (4),

4060 (24).
Gentry, H. S. 851m (31), 1128 (30), 1193 (30), 1287 (3), 2374 (30), 5076 (39),

5121 (14), 5189 (21), 5235 (21), 5563 (39), 5657 (14), 6484 (41).

CARLSON: THE GENUS RUSSELIA 287

Classman, S. F. 1559 (22).

Goldman, E. A. 62 (10), 467 (4).

Goll, G. P. 99 (24), 143 (24).

Grant, V. 761 (26).

Greenman, J. M. 73 (10), 82 (10), 185 (10).

Greenman, J. M. & M. T. 5024 (22), 5899 (50).

Gregg, J. 997 (14), 1185 (21).

Haenke, T. 1066 (34).

Hammerly, B. J. 278 (41).

Hartweg, Th. 501 (32).

Hayes, S. 36 (22).

Heriberto, Bro. 130 (22).

Hermann, J. 5067 (22).

Herndon, W. s. n. (26).

Hewitt, W. P. 14 (30).

Heyde, E. T. 525 (22).

Heyde, E. T. & Lux, E. 2943 (25).

Hinton, G. B. 1167 (46), 2111 (41), 2112 (41), 3191 (45), 7297 (45), 9188 (15),
10465 (12), 10921 (49), 11105 (41), 11393 (15), 11585 (15), 11773 (33),
12892 (41), 13197 (41), 13845 (20), 15620 (41), 15841 (21), 16057 (15),
16127 (6), 16129 (15).

Hitchcock, A. S. s. n. (22).

Hitchcock, C. L. & Stanford, L. R. 6886 (51).

Howell, J. T. 10397 (21).

Humboldt & Bonpland s. n. (16).

Jack, J. G. 4074 (10).

Johnson, E. P. 115 (27).

Johnson, H. 144 (22).

Johnston, I. M. 4062 (47).

Johnston, J. R. 550 (22), 1561 (22), 1743 (10).

Jones, M. E. 394 (21), 23169 (30), 23170 (38), 23217 (14), 24429 (41), 24430 (41),

27409 (41).
Juzepczuk, S. 1399 (42).

Karwinski, W. F. von 1093 (51), 1094 (43).

Keating, M. s. n. (41).

Kellerman, W. A. 5164 (22), 7609 (22).

Kenoyer, L. A. s. n. (51), A240 (51), A247 (39), A370 (51), 822 (10), 1578 (43),

2261 (51).

Kenoyer, L. A. & Crum, H. 3647 (51).
Killip, E. P. 3192 (22), 13816 (22).
Knoblock, J. 5623 (30).
Kunth, C. S. s. n. (32), (46).

Lamb, F. H. 479 (14), 578 (38).

Langlasse, E. 723 (15).

Langman, I. K. 2783 (41), 3127 (41), 3298 (41), 3360 (17), 3433 (10), 3693 (41),

3740 (41), 3825 (10), 3916 (10).
Lankester, C. H. k6 (24).

Leavenworth, W. C. & Hoogstraal, H. 1534 (45).
Lehmann, F. C. 1722 (24).
Leon, J. 459 (22), 871 (24), 10656 (22).
Le Roy, P. V. 153 (32).
Le Sueur, H. 1381 (30).

Liebmann, F. s. n. (32), 9426 (22), 9431 (10), 9434 (2).
Lundell, C. L. 526 (22), 526B (22), 576 (4), 1112 (4), 1331 (22), 1338 (4), 1644 (4),

2457 (25), 3507 (25), 6131 (22), 6706 (25), 6728 (22), 6868 (24), 6869 (24),

6925 (22), 6928 (22), 7850 (4), 12285 (43).
Lundell, C. L. & A. A. 7231 (40), 7288 (51), 7354 (22), 7849 (22), 7850 (4), 12166

(40), 12170 (51), 12257 (40), 12285 (42), 12317 (8), 12396 (5), 12398 (5),

12401 (10), 12402 (10), 12406 (10), 12422 (10), 12487 (51), 12487a (51),

288 FIELDIANA: BOTANY, VOLUME 29

12489 (51), 12490 (1), 12491 (40), 12568 (45), 12580 (17), 12581 (17), 12582
(17), 12595 (17), 12597 (16), 12646 (41), 12649 (5), 12650 (10), 12651 (10),
12652 (40).
Lyonnet, P. E. 369 (41), 608 (12).

Macbride, J. F. 2771 (22).

MacDaniels, L. H. 818 (24).

Maltby, F. S. 2 (14), 114 (21).

Mart, M. 1961 (11).

Martfnez-Calderon, G. 391 (10), 472 (10).

Matuda, E. 136 (11), 1144 (32), 1643 (18), 2088 (18), 2088a (18), 2563 (18),

3524 (22), 5188 (18), 17251 (10) 17273 (10), 17336 (10).
Maxon, W. R. 7654 (26).
Maxon, W. R., Harvey, A. D. & Valentine, A. T. 7366 (26), 7434 (26), 7451 (26),

7665 (26).

McVaugh, R. 10180 (14).
Mell, C. D. s. n. (22), (27); 38 (22).
Mexia, Y. 98 (21), 225 (21), 225^ (21), 1097a (21), 1122 (21), 1238 (37), 1815

(37), 8827 (45), 9164 (27).
Meyer, W. C. 117 (22).

Meyer, W. C. & Rogers, D. 2490 (51), 2664 (5).
Miller, J. W. s. n. (22).
Mohr, C. 307 (32).

Molina, A. 964 (24), 3078 (24), 3350 (24).
Moore, H. E. Jr. 1791 (51), 2531 (10).
Morley, T. 753 (26).
Muller, F. 1369 (10), 4098 (10).

Nelson, E. W. 319 (10), 385 (10), 538 (43), 857 (27), 1202 (43), 1293 (43), 2143 (41),
4289 (21), 6947 (45).

O'Neill, H. 8739 (22).

Orcutt, C. R. 3139 (10), 3299 (5), 4477 (41), 4674 (15), 6332 (40), 6458 (52),

6556 (10).

Orozco, J. M. 236 (22), 333 (24).
Ortega, J. G. 88 (41), 4637 (21), 5340 (41), 5658 (41), 6062 (21), 6491 (21), 6605

(14), 6760 (21).

Palmer, E. s. n. (14); 126 (12), 149 (12), 249 (30), 252 (17), 259 (51), 509 (33),

540 (41), 954 (21), 1448 (7).
Palmer, W. & Riley, J. H. 854 (22).
Paul, Bro. 149 (22).
Peck, M. E. s. n. (22).
Pelley, R. S. 15 (22).
Piper, C. V. 6021 (22).

Pittier, H. 498 (22), 2125 (22), 5384 (22), 13650 (22).
Pringle, C. G. s. n. (11), (12), (41); 2149 (41), 2568 (12), 5086 (5), 6081 (10),

8445 (41), 8657 (12), 8712 (41), 8934 (12), 9540 (41), 9621 (43), 10148 (44),

10367 (44), 11069 (41), 13152 (41), 13400 (41).
Purpus, C. A. s. n. (43), (51); 205 (42), 436 (10), 514 (41), 1276 (32), 2378 (5),

3270 (43), 3270a (43), 3952 (43), 5327 (5), 5749 (10), 6039 (2), 6242 (10),

6316 (10), 6882 (46), 7288 (46), 8063 (5), 8567 (10), 10833 (22).

Record, S. J. s. n. (22).

Rene, A. 12 (26), 69 (26).

Renson, C. 172 (22).

Robbins, J. W. s. n. (22).

Rodriguez, J. V. 150 (22), 862 (22), 924 (22), 1698 (24), 2704 (23), 3150 (24),

3151 (24).

Rose, J. N. 2508 (12), 3506 (41).
Rose, J. N. & Painter, J. H. 10026 (43).
Rose, J. N., Standley, P. C. & Russell, P. G. s. n. (31); 12712 (30), 13701 (21),

14313 (14), 14354 (38).

CARLSON: THE GENUS RUSSELIA 289

Rovirosa, J. N. 151 (27), 418 (27), 893 (27).

Rowell, C. M. Jr. 3171 (10).

Rowlee, W. W. & Stork, H. E. 134 (22), 686 (22), 846 (22).

Rusby, H. H. s. n. (43); 9 (12), 12 (12).

Russell, P. G. & Souviron, M. J. 18 (12).

Schiede, C. & Deppe, F. 1186 (5).

Schipp, W. A. S653 (4), 671 (25).

Seaton, H. E. s. n. (10).

Seler, E. 43B (41), 705 (10), 1747 (43), 4965 (10).

Shank, P. J. & Molina, A. 4583 (26).

Shannon, W. C. 271 (10).

Sharp, A. J. 441460 (41).

Shimek, B. & Smith, C. L. 20 (26).

Skutch, A. F. 951 (7), 1448 (7), 1770 (24), 2085 (7), 4134 (22), 4670 (22).

Smith, A. P. 2371 (22).

Smith, C. L. s. n. (26); 1100 (22).

Smith, H. H. 1361 (22).

Smith, J. D. 1642 (24), 2943 (22).

Smith, J. G. 303 (10), 418 (27).

Smith, L. C. 154 (43), 400 (43).

Solis, F. 82 (24), 328 (22), 366 (24).

Standley, P. C. 487 (24), 1349 (24), 1666 (22), 8184 (26), 8443 (26), 9437 (26),
9449 (26), 9486 (26), 10109 (26), 10595 (26), 19288 (22), 20037 (22), 20099
(22), 20412 (25), 20722 (24), 20883 (23), 21630 (24), 22722 (22), 23312 (22),
23565 (22), 23891 (22), 24508 (22), 26325 (22), 26478 (22), 27000 (22),
27904 (22), 29141 (22), 29313 (22), 32188 (22), 35904 (22), 36033 (22),
40070 (22), 55919 (23), 56230 (23), 58262 (22), 59584 (50), 60235 (22),
60277 (22), 60401 (22), 60668 (22), 62200 (10), 63289 (22), 63592 (10), 63944
(22), 64578 (22), 64947 (22), 67099 (7), 67165 (7), 67275 (7), 72264 (22),
72282 (22), 73994 (50), 74660 (22), 74754 (22), 75484 (22), 75705 (22),
75953 (22), 76941 (19), 77067 (19), 77093 (19), 77591 (22), 77615 (22),
77716 (22), 78254 (25), 80371 (50), 80390 (50), 80427 (50), 84737 (7), 84762
(7), 84856 (7), 86690 (7), 87111 (7), 87129 (7), 87796 (10), 88178 (10), 88923
(22), 89070 (10), 89384 (22).

Standley, P. C. & Chacon, J. 5241 (22), 6489 (23).

Standley, P. C. & Lindelie, H. 0. 7416a (22).

Standley, P. C. & Padilla, E. 2478 (22), 2495 (22), 2840 (22), 3353 (24), 3454 (22).

Stanford, L. R., Lauber, & Taylor 2036 (51).

Stanford, L. R., Retherford, K. L. & Northcraft, R. D. 826 (51).

Steere, W. C. 2287 (22), 2484 (22).

Steyermark, J. A. 29308 (23), 29464 (50), 33214 (7), 37737 (7), 37982 (9), 38497
(22), 39398 (29), 42917 (50), 42980 (36), 44056 (25), 44626 (22), 45281 (22),
49555 (24), 52174 (22).

Stork, H. E. 2788 (26), 3328 (22).

Thieme, C. 5384 (24).

Tonduz, C. 7326 (22), 8775 (22).

Velasco, L. V. 9007 (22).

Viereck, H. W. 482 (51).

Von Hagen, C. & W. 1210 (24).

Von Rozynski, H. W. 41 (51), 41a (51), 485 (51), 729 (5), 759 (10).

Von Tuerckheim, H. 1198 (50), 2249 (13).

Watson, S. 474a (22).

West, J. 3505 (41).

White, G. 77 (22), 297 (22).

Wiggins, I. L. 7511 (30).

Williams, L. 8521 (10), 9056 (27), 9251 (27), 9254 (27), 9321 (27), 9848 (27),

16039 (22).
Williams, L. & Molina, A. 10051 (24), 10118 (24), 12252 (22), 13290 (22), 14330

(24), 15843 (24).

290 FIELDIANA: BOTANY, VOLUME 29

Williams, R. S. 185 (22).

Wilson, C. L. 251 (24).

Woodson, R., Allen, P. H. & Seibert, R. 1477 (22).

Wooton, E. D. s. n. (51).

Wright, C. s. n. (22).

Wright, W. G. 1203 (21).

Yuncker, T. G., Dawson, R. F. & Youse, H. R. 5572 (23).
Yuncker, T. G., Koepper, J. M. & Wagner, K. A. 8719 (29).

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CARLSON: THE GENUS RUSSELIA 291

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292

FIELDIANA: BOTANY, VOLUME 29

Index to Species, Varieties and Forms

New species, varieties and forms are printed in bold-faced type, previously
published accepted names in roman type, synonyms in italic type.

acuminata 248

alata (excluded) 285

alternifolia (excluded) 285

americana (excluded) 285

campechiana 246

capensis (excluded) 285

chiapensis 257

coccinea 251

coccinea f. stipitata 253

colombiana 259

conzattii 280

cuneata 255

deamii 253

depressa (excluded) 285

elongata 245

ephedroides 269

equisetiformis 224

flammea (excluded) 285

flavoviridis 259

floribunda 256

floribunda var. pubescens 257

furfuracea 274

glandulifera 253

grandidentata 281

hintoni 281

jaliscensis 253

juncea 244

laciniata 250

lanceifolia 250

leptopoda 258

longif olia 268

longisepala 254

maculosa 275

multiflora 251

obtusata 277

ovatifolia 256

oxyphylla 267

paniculata 251

parvif olia 282

pennelliana 283

pennelliana yar. pilosa 283

peruviana (Excluded) 285

polyedra 283

pringlei 278

pubescens 273

purpusii 245

racemosa (excluded) 285

retrorsa 275

retrorsa f. nudicostata 277

rotundifolia 271

rotundifolia f. velutina 271

rugosa 271

sarmentosa 259

sarmentosa var. nicaraguensis. 265

sarmentosa var. oaxacensis. . . . 266

sarmentosa f. eglandulata 263

sarmentosa f. pubescens 264

sarmentosa f. velutina 265

serratifolia 251

sonorensis 268

sonorensis var. pubescens 269

staleyae 284

Scrophularia coccinea 251

standleyi 257

steyermarkii 272

subcoriacea 247

syringaef olia 247

tabacensis 259

tenuis 258

tepicensis 273

teres 248

ternifolia 279

tetraptera 254

trachypleura 275

verticillata 269

villosa . . . 279

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