FLORA OF GUATEMALA
PART IX
FLORA OF GUATEMALA
PAUL C. STANDLEY
Late Curator of the Herbarium
Field Museum of Natural History
LOUIS 0. WILLIAMS
Chairman, Department of Botany
Field Museum of Natural History
AND
DOROTHY N. GIBSON
Supervisor of the Herbaria
Field Museum of Natural History
FIELDIANA: BOTANY
VOLUME 24, PART IX, NUMBERS 1-4
Published by
FIELD MUSEUM OF NATURAL HISTORY
1970-1973
The research on Part IX of the "Flora of Guatemala" was made possible
through generous grants from the National Science Foundation to Field Museum
of Natural History, Louis O. Williams, principal investigator.
Library of Congress Catalog Card Number: 48-3076
PRINTED IN THE UNITED STATES OF AMERICA
BY FIELD MUSEUM PRESS
9
-
FLORA OF GUATEMALA
PAUL C. STANDLEY
AND
LOUIS 0. WILLIAMS
9
The Lihrarv of the
at Uroana-Clianripaigi,
FIELDIANA: BOTANY
VOLUME 24, PART IX, NUMBERS 1 AND 2
Published by
FIELD MUSEUM OF NATURAL HISTORY
JUNE 26, 1970
FLORA OF GUATEMALA
PART IX
FLORA OF GUATEMALA
PAUL C. STANDLEY
The Late Curator of the Herbarium
AND
LOUIS 0. WILLIAMS
Chief Curator, Botany
FIELDIANA: BOTANY
VOLUME 24, PART IX, NUMBERS 1 AND 2
Published by
FIELD MUSEUM OF NATURAL HISTORY
JUNE 26, 1970
PUBLICATION 1100
Library of Congress Catalog Card Number: 48-3076
PRINTED IN THE UNITED STATES OF AMERICA
BY FIELD MUSEUM PRESS
Flora of Guatemala - Part IX, Numbers 1 and 2
TUBIFLORAE
PAUL C. STANDLEY AND Louis O. WILLIAMS
The Tubiflorae, in the sense of Engler and Melchior, Syllabus der
Pflanzenfamilien (12th edition), in Guatemala contains 16 families.
Some of these families are important in the flora, with numerous
genera and species. The group, as a whole, is most conspicuous in
the flora toward the end of the rainy season.
The families of this order are often very closely related. Because
of this relationship, due to parellel development or not, it is often
difficult, if not impossible, to write a family description of one family
which will exclude all those members of a related family. It is, in
fact, sometimes difficult to relate a genus to a family and to be sure
that it does not belong in another.
These families of the Tubiflorae, with exceptions, have corollas
that are (or are said to be) tubular, hence Tubiflorae; the flowers are
perfect; the leaves are simple; the stamens are usually not adnate and
are said to be free; the ovary is superior, and of united carpels; there
are no stipules. Exceptions occur in every one of these characters,
hence a key to the families in which exceptions occur: I, those that
have compound leaves; II, those in which the ovary is inferior or
appears to be so; III, those in which the stamens (or anthers) are
connate; IV, those in which unisexual flowers occur; and V, the
family in which stipules (actually pseudostipules) occur.
The key to the families of the Tubiflorae was prepared by Mary
James as part of a project carried on under a Shinner Foundation
scholarship at Field Museum.
PROVISIONAL KEY TO TUBIFLORAE
Leaves scale-like, lacking chlorophyll or seemingly so; parasites (or saprophytes).
Vines, slender and leafless, usually yellow. .
Not vines, usually not slender and not yellow.
Ovules numerous in each locule • Orobanchaceae.
Ovule one in each locule Lennoaceae.
2 FIELDIANA: BOTANY, VOLUME 24
Leaves not scale-like, chlorophyll present; sometimes parasites (or saprophytes).
Leaves opposite.
Flowers zygomorphic.
Stamens the same number as the corolla lobes.
Stigma one.
Ovule one in each locule Solanaceae.
Ovules more than one in each locule.
Ovules two or more in each locule.
Locule one Boraginaceae.
Locules two or more Labiatae.
Ovules numerous in each locule.
Placentation axile Solanaceae and Scrophulariaceae.
Placentation parietal Gesneriaceae.
Stigmas two or more.
Ovule one in each locule.
Locule one Pedaliaceae.
Locules two or more.
Corolla lobes imbricate Verbenaceae.
Corolla valvate or contorted Polemoniaceae.
Ovules more than one in each locule.
Ovules two or more in each locule.
Locule one.
Corolla lobes imbricate Pedaliaceae.
Corolla valvate or contorted Acanthaceae.
Locules two or more Polemoniaceae.
Ovules numerous in each locule.
Corolla lobes imbricate.
Locule one Pedaliaceae and Scrophulariaceae (rare).
Locules two Scrophulariaceae and Polemoniaceae.
Corolla valvate, convolute or contorted.
Locule one Acanthaceae.
Locules two or more Polemoniaceae.
Stamens not the same number as the corolla lobes.
Stigma one.
Ovule one in each locule Solanaceae.
Ovules more than one in each locule.
Ovules two or more in each locule Labiatae.
Ovules numerous in each locule.
Placentation axile.
Locule one Pedaliaceae and Scrophulariaceae (rare).
Locules two or more Scrophulariaceae.
Placentation parietal Gesneriaceae.
Stigmas two or more.
Ovule one in each locule.
Locule one Pedaliaceae.
Locules two or more Verbenaceae.
Ovules more than one in each locule.
Ovules two or more in each locule.
Corolla lobes imbricate . . . . Pedaliaceae.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 3
Corolla lobes valvate Acanthaceae.
Ovules numerous in each locule.
Placentation axile.
Corolla lobes imbricate.
Locule one Pedaliaceae and Scrophulariaceae (rare).
Locules two or more
Scrophulariaceae and Bignoniaceae (rare).
Corolla valvate, convolute, or contorted Acanthaceae.
Placentation parietal Martyniaceae and Bignoniaceae (rare).
Flowers actinomorphic.
Stigma one.
Ovule one in each locule Solanaceae.
Ovules more than one in each locule.
Ovules two or more in each locule.
Locule one Boraginaceae.
Locules two or more Labiatae.
Ovules numerous in each locule.
Placentation axile Solanaceae.
Placentation parietal.
Sepals valvate Gesneriaceae.
Sepals imbricate Hydrophyllaceae and Gesneriaceae (rare).
Stigmas two or more.
Corolla lobes imbricate Verbenaceae.
Corolla lobes valvate Polemoniaceae.
Leaves alternate.
Flowers zygomorphic.
Stamens same number as corolla lobes.
Stigma one.
Ovule one in each locule Solanaceae.
Ovules more than one in each locule.
Ovules two or more in each locule Boraginaceae.
Ovules numerous in each locule.
Placentation axile Solanaceae and Scrophulariaceae.
Placentation parietal Gesneriaceae.
Stigmas two or more.
Corolla lobes valvate Polemoniaceae.
Corolla lobes imbricate Scrophulariaceae.
Stamens not the same number as the corolla lobes.
Ovule one in each locule Solanaceae.
Ovules numerous in each locule
Solanaceae, Scrophulariaceae and Lentibulariaceae.
Flowers actinomorphic.
Stigma one.
Ovule one in each locule Solanaceae and Convolvulaceae.
Ovules more than one in each locule.
Ovules two or more in each locule. .Boraginaceae and Convolvulaceae.
Ovules numerous in each locule.
Placentation axile Solanaceae.
Placentation parietal Hydrophyllaceae and Gesneriaceae.
4 FIELDIANA: BOTANY, VOLUME 24
Stigmas two or more.
Corolla lobes imbricate Verbenaceae and Convolvulaceae.
Corolla lobes not imbricate.
Corolla lobes plicate-contorted Convolvulaceae.
Corolla lobes valvate Polemoniaceae.
EXCEPTIONS
I. FAMILIES OF TUBIFLORAE IN WHICH COMPOUND LEAVES MAY OCCUR.
Stamens the same number as the corolla lobes.
Stigma one Labiatae.
Stigmas two or more.
Corolla lobes imbricate Verbenaceae.
Corolla lobes valvate Polemoniaceae.
Stamens not the same number as the corolla lobes.
Stigma one Labiatae.
Stigmas two or more.
Ovule one in each locule Verbenaceae.
Ovules numerous in each locule Bignoniaceae.
II. FAMILIES OF TUBIFLORAE IN WHICH INFERIOR OVARIES MAY OCCUR.
Stigma one; parietal placentation Gesneriaceae.
Stigmas two or more; axile placentation Pedaliaceae.
III. FAMILIES OF TUBIFLORAE IN WHICH THE STAMENS MAY BE CONNATE (mostly
by anthers).
Stigma one.
Ovule one in each locule Solanaceae.
Ovules numerous in each locule.
Placentation axile Solanaceae and Scrophulariaceae.
Placentation parietal Gesneriaceae.
Stigmas two or more.
Corolla lobes imbricate Scrophulariaceae and Bignoniaceae.
Corolla lobes valvate, convolute, contorted Acanthaceae.
IV. FAMILIES OF TUBIFLORAE IN WHICH UNISEXUAL FLOWERS MAY OCCUR.
Stigma one.
Corolla lobes convolute, plicate-contorted Convolvulaceae.
Corolla lobes imbricate Boraginaceae and Convolvulaceae.
Stigmas two or more Verbenaceae and Convolvulaceae.
V. FAMILY OF TUBIFLORAE IN WHICH STIPULES OR PSEUDOSTIPULES MAY OCCUR.
One family Solanaceae.
CONVOLVULACEAE. Morning Glory family
PAUL C. STANDLEY and Louis O. WILLIAMS
Herbs, vines (usually twining), shrubs or rarely trees, the sap usually milky;
leaves simple but often lobate, pinnate or pectinate, or reduced to scales in Cuscuta,
STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 5
^stipulate; inflorescence axillary, the flowers solitary or in dichasia, heads or pan-
cles; flowers perfect, regular or slightly zygomorphic, often large and showy but
'ugaceous; sepals 5, free, imbricated, equal or not, usually persistent and often
iccrescent in fruit, corolla sympetalous, tubular, funnelform, campanulate or sal-
perform, the limb with 5 lobes or teeth or almost entire, often induplicate in bud
ind with distinctly limited longitudinal midpetaline areas or stripes; anthers 5,
iistinct, inserted on the base of the corolla tube and alternate with the lobes, linear
jr oblong, 2-celled, introrse; disc annular or cupular, sometimes 5-lobate, or none;
wary superior, mostly 2-3-celled and each cell biovulate, or 4-6-celled and the
jells uniovulate, rarely 1-celled with 4 ovules; the ovary of 2-4 almost free carpels;
;tyle filiform, simple or bifid or with 2 distinct styles; stigma capitate or bilobate
3r the stigmas 2 and globose, ellipsoid or linear; fruit 4-1-celled, usually capsular
ind dehiscent by valves, rarely transversely or irregularly dehiscent or indehiscent;
seeds often fewer than the ovules, often pubescent, with sometimes scanty but very
lard endosperm.
The Convolvulaceae, worldwide but most numerous in the tropics'
contains some 40 or 50 genera. The largest of these genera is Ipomoea
tvhich contains some 300-400 species, which are often common and
showy in Central America in both dry and wet regions. Ipomoea
batatas, the sweet potato, is one of the important food plants in the
tropics of the world. It is commonly grown in Central America.
Several members of this family are grown as ornaments and numer-
ous weeds occur in the family. One other genus, Stictocardia, is
known in Central America.
The family is much in need of critical revision. It is a natural one
and the genera are not easy to delimit. Most of the work on the Con-
volvulaceae in recent years has been carried out by the brilliant
Argentine botanist Carlos Alberto O'Donell (1912-1954). Post-
humous works published in Lilloa (29: 19-376. 1959) indicate
O'Donell's thinking along generic lines. It was a great tragedy that
this man of such great promise died so young.
Plants slender twining parasites, without any green coloring, the stems yellow or
orange; leaves reduced to scales Cuscuta.
Plants not parasitic, the leaves green and normally developed.
Ovary and fruit deeply bilobate, or the carpels distinct; small creeping plants
with minute flowers; leaves reniform, sericeous beneath Dichondra.
Ovary and fruit entire, not lobate.
Fruits indehiscent, somewhat baccate or dry, or dehiscent but the seeds sur-
rounded by orange-colored pulp; woody vines.
Leaves cordate or subcordate at the base; seeds usually 1 ; corolla white.
Turbina.
Leaves obtuse or acute at the base.
Fruit indehiscent; flowers purple, 3.5-5 cm. long; leaves glabrous beneath
or nearly so Maripa.
Fruit dehiscent; flowers greenish, 1 cm. long; leaves densely sericeous
beneath. . . .Itzaea.
6 FIELDIANA: BOTANY, VOLUME 24
Fruit dehiscent, dry; seeds never surrounded by pulp.
Style 1 and bifid, or the styles 2 and distinct.
Plants scandent, woody; styles bifid; stigmas capitate Bonamia.
Plants not scandent, herbaceous, small; styles 2; stigmas elongate.
Evolvulus.
Style 1, entire.
Sepals very unequal, the outer ones much broader and concealing the
inner ones, ovate, green; herbaceous vines; leaves obtuse or acute at
the base; flowers white Aniseia.
Sepals subequal, or but slightly unequal, the outer ones often shorter or
narrower than the inner.
Stigmas elliptic or oblong; small or large vines, the flowers usually
small lacquemoniia.
Stigmas globose or biglobose.
Capsule transversely dehiscent Operculina.
Capsule opening by vertical valves.
Stamens and style exserted.
Corolla salverform, white or blue Calonyction.
Corolla funnelform or tubular, rarely salverform, red, orange,
yellow, bronze, or greenish Quamoclit.
Stamens and style not exserted.
Pollen smooth; herbaceous vines; flowers yellow or white.
Merremia.
Pollen echinate; plants various in habit, often woody; flowers
various in color, most often blue or purple, rarely white or
yellow Ipomoea.
ANISEIA Choisy
Plants herbaceous, prostrate or scandent; leaves linear to ovate or elliptic,
often mucronate, entire; flowers small, axillary, solitary or in few-flowered dichasial
inflorescences; sepals 5, herbaceous, unequal, the 3 outer ones larger, often decur-
rent on the pedicel; corolla funnelform, the limb 5-dentate or subentire, with 5
longitudinal pubescent stripes outside; stamens and style included; ovary glabrous,
2-celled, the cells bi-ovulate; disk small or none; style 1, slender, the stigma bi-
globose; capsule globose, 2-celled, 4-valvate, 4-seeded.
About five species, one in the Old World tropics, the others
American. One other species is known from southern Central
America.
Aniseia cernua Moric. PL Nouv. Amer. 56. 1838; Standl. &
Steyerm. Field Mus. Bot. 22: 270. 1940. Jacquemontia chiapensis
Brandeg. Univ. Calif. Publ. Bot. 6: 60. 1914.
Wet thickets or wet meadows bordering lakes, 800 m. or less;
Jutiapa; Escuintla. Southern Mexico; British Honduras; El Sal-
vador; Honduras; South America.
A slender herbaceous vine, the stems prostrate and rooting or usually twining,
appressed-pilose; leaves narrowly oblong or linear-lanceolate, 3-10 cm. long, 1.5
STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 7
cm. broad or narrower, acute or rounded and mucronate at the apex, acute at the
base, somewhat fleshy when fresh, appressed-pilose with whitish hairs or glabrate,
short-petiolate; peduncles axillary, mostly 1-flowered, shorter than the leaves;
bracts linear, 2-4 mm. long; sepals appressed-pilose, the 3 outer ones ovate, acute
or acuminate, broadly rounded or subcordate at the base, 9-13 mm. long, the 2
inner ones ovate, acuminate; corolla white, 1.5 cm. long or larger; capsule 2 cm.
long; seeds black, glabrous.
Called "bejuco de pescado" in El Salvador, the tough stems being
used there for stringing fish. A. martinicensis (Jacq.) Choisy and its
var. nitens (Choisy) O'Donell occur in Central America.
BON AM I A Petit-Thouars
Reference: Tin Myint & Daniel B. Ward, A taxonomic revision of
the genus Bonamia, Phytologia 17: 121-239, figs. 1968.
Herbaceous or woody vines, or the plants sometimes prostrate and not twin-
ing, glabrous or pubescent; leaves entire; flowers cymose, the cymes axillary and
short-pedunculate, sometimes forming a terminal panicle, the bracts small, sepals
obtuse or rarely acute, subequal or the outer ones larger and orbicular-cordate;
corolla campanulate, the limb plicate, 5-angulate, stamens shorter than the corolla,
the filaments filiform, often dilated at the base, the anthers oval or oblong; ovary
2 celled, 4-ovulate, the style filiform, bifid, or the styles 2 and free, the stigmas
capitate; capsule globose or ovoid, 4-valvate, membranaceous or coriaceous; seeds
4 or by abortion fewer, glabrous or pilose.
Species about 45, in tropical regions of both hemispheres. Only
three species have been found in Central America.
Sepals 1-1.5 cm. long B. sulphured.
Sepals 0.4-0.5 cm. long B. brevipedicellata.
Bonamia brevipedicellata Myint & Ward, Phytologia 17: 188,
fig. 1968.
Forest edges and clearings, near sea level, endemic, Machaca,
British Honduras (type Schipp 1210).
Woody vines to 15 m. or more long and 2-3 cm. in diameter. Leaves elliptic
to ovate-elliptic, acute or acuminate, rounded or obtuse at the base, 8-12 cm. long
and 4-7 cm. broad, glabrous above, densely puberulent below, the petioles 2-3 cm.
long; inflorescence axillary, a compact many-flowered cyme, up to 10 cm. long or
perhaps more with short pedicels to 3 mm. long; sepals subequal, ovate-orbicular,
4-5 mm. long and 3-4 mm. broad, densely puberulent; corolla campanulate with a
narrow cylindrical tube, 1-1.2 cm. long, hirsute on the interplicae; stamens in-
cluded; styles free to the base, subequal, 1 cm. long or less, stigmas peltate sub-
globose.
Bonamia sulphurea (Brandegee) Myint & Ward, Phytologia
17: 178. 1968. Breweria sulphurea Brandegee, Univ. Cal. Publ. Bot.
4: 384. 1913.
FIG. 1. Bonamia sulphured. A, habit, natural size; B, calyx (imbricated) with
styles, X 2; C, corolla dissected to show stamens, X 2; D, foliar pubescence, much
enlarged.
8
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 9
Rocky, brushy hillsides, 200-900 m.; Zacapa; Chiquimula.
Mexico (Veracruz) ; Honduras.
A woody vine, climbing over shrubs, the stems covered with short, brownish
or yellowish, ascending hairs, glabrous in age; leaves short-petiolate, rather thick,
entire, elliptic-oblong to oval, 4-7. 5 cm. long, rounded and mucronate at the apex,
rounded at the base, thinly appressed-pilose above, densely fulvous-pilose or to-
mentose beneath; flowers numerous, in small cymes, these short-pedunculate or
sessile, the pedicels long and slender; sepals 1-1.5 cm. long, lance-oblong or ovate-
oblong, acute, appressed-pilose; corolla white, 1.5 cm. long, hirsute outside; imma-
ture capsule hirsute.
CALONYCTION Choisy
Large vines, usually herbaceous throughout, the stems sometimes covered with
fleshy prickles; leaves large, thin, entire or lobate, cordate at the base; flowers large,
white or blue, opening in the evening and closing in the morning; sepals coriaceous
or herbaceous, the outer ones often with subulate appendages; corolla long-salver-
form, the tube usually cylindric or nearly so; stamens and style exserted; styles
united, the stigmas globose; capsule globose, 2-celled, 4- seeded.
Half a dozen species, widely dispersed in tropical regions. Prob-
ably only the following occur in Central America. Hardly distinct
from Ipomoea.
Sepals with conspicuous subulate green appendages at the apex; stems with thick
aculeate appendages C. aculeatum.
Sepals without subulate appendages.
Stems long hirsute with hollow hairs, flowers blue C. clavatum.
Stems glabrous or with scale-like pubescence.
Calyx lobes subequal, mostly less than 1 cm. long; stems glabrous; leaves not
lobate C. ventricosum.
Calyx lobes very unequal, more than 1 cm. long; stems with scale-like pubes-
cence; leaves lobate C. sp.
Galonyction aculeatum (L.) House, Bull. Torr. Bot. Club 31 :
590. 1904. Convolvulus aculeatus L. Sp. PI. 155. 1753. Ipomoea bona-
nox L. Sp. PL ed. 2. 228. 1762. I. aculeata Kuntze, Rev. Gen. PI. 2:
442.1891. Luna blanca (Pete"n) . Moon-flower.
Moist or wet thickets, often in second growth, sometimes running
over rocky banks, 800 m. or less; Pete"n; Alta Verapaz; Izabel;
Zacapa; Santa Rosa; Escuintla; Solola; Suchitepequez ; Retalhuleu;
San Marcos. Southern Florida; Mexico; British Honduras to El
Salvador and Panama; West Indies; South America; Old World
tropics.
A small or large herbaceous vine, usually scandent, glabrous, sometimes 20 m.
long or more and covering large trees, the sap milky; stems densely armed with
short fleshy prickles to smooth; leaves long-petiolate, rounded-ovate, thin, 5-15 cm.
10 FIELDIANA: BOTANY, VOLUME 24
long, entire or rarely trilobate, abruptly short-acuminate, deeply cordate at the
base; peduncles bearing 1-several flowers, usually elongate but shorter than the
leaves; sepals green and fleshy, about 1 cm. long, the outer ones (at least) termi-
nated by a long slender green horn-like appendage; corolla tube very slender, 10-
12 cm. long, the limb white, 8-10 cm. broad, each lobe with a broad green median
stripe; capsule ovoid, pointed, 2 cm. long; seeds black, glabrous or nearly so.
There is a problem involved in the selection of the proper name
for this plant if placed in the genus Ipomoea. It is possible that /.
tuba (Schlecht.) G. Don is the proper one.
Called "panal de nifio" in Honduras; "nata," "naxh" (Chiapas);
"nicua," "cracion" (Yucatan); "zutub" (Yucatan, Maya); "campa-
nilla blanca," "pitoreta," "bejuco de tabaco," "flor de luna," "galan
de noche," "garza" (El Salvador). In the evening, when in full bloom,
the plant is a showy and handsome one. It is often grown for orn-
ament in temperate regions where it is not hardy, but must be grown
from seeds each summer. When the proper stage of dusk or darkness
has arrived, the buds open rapidly, in only a few minutes. The milky
sap of this plant and of various other Convolvulaceae has long been
used in Central America for coagulating the latex of Castillo, rubber
trees. The tough stems often are employed in place of string,
especially for hanging tobacco leaves to cure (hence the name bejuco
de tabaco).
Calonyction clavatum G. Don, Gen. Syst. 4: 264. 1838. Ipo-
moea clavata v. Oostr. & Macbride, Field Mus. Bot. 11: 3. 1931.
Luna morada.
Moist or dry forest or thickets, 800 m. or less; Pete'n. Southern
Mexico; British Honduras; Nicaragua; Costa Rica; western South
America, southward to Ecuador.
A large herbaceous vine, the stems hirsute with very long and soft, hollow,
spreading, whitish hairs; leaves long-petiolate, the petioles long-hirsute; leaf blades
glabrous or nearly so, rounded ovate, large, acute or acuminate, deeply cordate at
the base, entire; peduncles mostly 1-flowered, densely hirsute, much shorter than
the petioles, the pedicels stout, glabrous, usually longer than the peduncles, fistu-
lose-thickened above; sepals 2.5-3 cm. long, muticous, glabrous, lance-oblong, ob-
tuse or subacute; corolla blue, about 12 cm. long, the limb 10 cm. broad, the tube
dilated upward, 2.5 cm. broad in the throat, glabrous; capsule broadly ovoid,
2.5 cm. long, glabrous; seeds densely covered with long hairs.
Called "morning-glory" and "gloria de la mafiana" in British
Honduras. The latter name is a Spanish translation of the English
one, and probably of recent introduction. The usual Spanish name
for plants of this family in most parts of Central America is "cam-
FIG. 2. Calonydion aculeatum. A, habit, X 1A', B. leaf from another collection,
natural size. The fruit is also X 1A- Stems are not always aculeate.
11
12 FIELDIANA: BOTANY, VOLUME 24
panilla." The vine is an exceptionally beautiful one, one of the most
gorgeous of all the morning-glories, its huge blossoms of an exquisite
shade of blue rarely seen in flowers.
Calonyction ventricosum Hallier f., Bot. Jahrb. 16: 556. 1893,
nomen; Bull. Herb. Boiss. 5: 1027. 1897. Ipomoea santillanii
O'Donell, Anal. Inst. Biol. Mex. 12: 93. 1941. Quilamul; campana.
Moist or wet thickets, often in roadside hedges, 200-1,800 m.;
Chiquimula; Jutiapa; Santa Rosa; Guatemala; Chimaltenango;
Suchitepe'quez; Quezaltenango. Southern Mexico; El Salvador;
Costa Rica.
A large, coarse herbaceous vine, glabrous throughout or nearly so; leaves long-
petiolate, thin, entire, rounded-ovate, usually caudate-acuminate, deeply cordate
at the base; peduncles often equaling the leaves, divided into 2 long scorpioid
branches, several-flowered, the pedicels mostly short, thickened above; bracts whit-
ish or pale green, sometimes tinged with pink, membranaceous, broadly spatulate,
large and concealing the flower buds; sepals ovate, glabrous, rounded at the apex,
muticous, 1 cm. long; corolla white, the lower part of the tube slender, 3 cm. long,
the upper portion dilated, 2.5 cm. long and 2 cm. broad, the limb 6 cm. broad
or more.
A showy and rather handsome plant. It is particularly plentiful
and conspicuous on the otherwise sterile and uninteresting slopes of
Cuesta de la Conora in the Department of Jutiapa.
Calonyction sp.
Probably edges of clearings or thickets, Alta Verapaz, Steyermark
M999; 45121.
Coarse herbaceous vine; the stems covered with scale-like pubescence to about
3 mm. long, glabrous with age; leaves trilobate, often to near the middle, cordate,
broadly ovate-suborbicular in outline, to 15 cm. long and nearly as broad, petioles
to about 10 cm. long, with sparse scale-like pubescence or none; the calyx lobes in
fruit very unequal and large, coriaceous, from about 1.5 cm. long to 4 cm. long;
immature capsule to 3 cm. long; seeds pubescent with scale-like hairs.
These specimens have been called C. clavatum and C. ventricosum,
which they are not.
CUSCUTA L. Dodder
References: T. G. Yuncker, Revision of the North American and
West Indian species of Cuscuta, Univ. 111. Biol. Monog. 6, nos. 2-3:
1-142. 1921; The Genus Cuscuta, Mem. Torr. Bot. Club 19: 113-331.
1932; Cuscuta, No. Am. Fl. ser. 2, 4: 1-51. 1965.
Small, twining, parasitic plants without chlorophyll, the stems filiform, yellow
or orange (rarely greenish), attached by haustoria to the host plant; leaves reduced
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 13
:o minute scales; inflorescence of cymose clusters of small, usually white flowers;
^alyx gamosepalous and five-parted or rarely the sepals nearly free; corolla usually
arceolate or campanulate, the lobes various and usually 5, with scale-like append-
iges attached at the base within and opposite the stamens to form a corona or
sometimes lacking; ovary 2-celled and each cell biovulate; styles distinct (rarely
inited), terminal, the stigmas capitate to linear; capsule indehiscent or circum-
scissle near the base.
The genus has perhaps as many as 150 species. The host pref-
erence of those of Guatemala is little known and it is assumed that
nost occur on a variety of hosts. Germination of the seeds is in the
Around, the seedling plants soon attach themselves to host plants
md the attachment with the soil withers. Certain species cause
considerable damage in cultivated crops, especially in flax, beets, and
chiles but we do not know of damage to crops by Cuscuta in Guate-
nala.
Hapsule indehiscent; section Clistogrammica.
Capsule globose with a conspicuously thickened stylopodium; flowers rather
thick and fleshy, papillate or glandular C. indecora.
Capsule depressed-globose, not definitely thickened at the apex.
Calyx lobes acute C. yucatana.
Calyx lobes obtuse.
Corolla lobes obtuse C. obtusiflora.
Corolla lobes acute C. campestris.
Capsule circumscissile; section Eugrammica.
Styles stout and more or less subulate, tapering to the globose or conic ovary.
Calyx deeply cleft, the lobes distinct or nearly so C. boldinghii.
Calyx not deeply lobate, definitely gamosepalous.
Calyx lobes much broader than long at maturity; stigmas large and con-
volute C. rugosiceps.
Calyx lobes about as broad as long; stigmas not conspicuously enlarged
or convolute C. mlapemis.
Styles slender, usually of about the same thickness throughout.
Calyx lobes, and usually also the corolla lobes, obtuse.
Flowers about as broad as long C. tinctoria.
Flowers longer than broad.
Corolla saccate, subglobose C. corymbosa.
Corolla not saccate, cylindric C. cozumeliensis.
Calyx lobes, and usually also the corolla lobes, acute.
Pedicels mostly longer than the flowers, in lax inflorescences; lobes of the
corolla about equaling the tube C. saccharata.
Pedicels not longer than the flowers, in dense compact inflorescences;
lobes of the corolla mostly shorter than the tube . . . C. costaricensis.
Cuscuta boldinghii Urban, Repert. Sp. Nov. 16: 38. 1919.
Growing on various herbs, 600 m. or less; Santa Rosa; Escuintla;
Retalhuleu; Solola. Southern Mexico; Honduras; Nicaragua; West
[ndies; Costa Rica. Venezuela. Florida.
14 FIELDIANA: BOTANY, VOLUME 24
Plants slender; flowers about 2 mm. long, sessile or nearly so, in dense clusters;
calyx lobes equaling or longer than the corolla tube, distinct or nearly so, imbricate,
oblong or oval, obtuse and mucronate, or obtuse and bearing a horn-like projection
from its dorsal surface near the apex; corolla lobes about equaling the campanulate
tube, erect or spreading, oblong or oval, obtuse, irregularly dentate at the apex
and with a subapical horn-like projection; stamens shorter than the corolla lobes,
the anthers small, oval-ovate, shorter than the stout subulate filaments; scales
reaching the filaments, oblong or broadly ovate, short-lacerate; styles stout, subu-
late, much longer than the globose ovary, the stigmas capitate; capsule globose,
circumscissile, bearing the withered corolla at its apex; seeds oval, 1 mm. long.
The species is noteworthy for the horn-like projections on the
perianth lobes. The plant is called "hierba mala" in Honduras and
reported to be poisonous to stock. The genus yields a substance
called cuscutin, but there is no clear proof that it has been found
poisonous in the United States, although it has been suspected.
Cuscuta campestris Yuncker, Mem. Torr. Bot. Club 18: 138,
fig. 1932; No. Am. Fl. ser. II, pt. 4: 5. 1965.
Parasite in clearings near sea level, British Honduras (Gentle
791 It). Said by Yuncker to be distributed throughout the range of
the genus, but no specimens in our herbarium determined by him.
The type locality is Texas. The description is that of Yuncker.
"Stems medium; flowers up to about 1.5-2 mm. long from the base of the
flower to the corolla sinuses, appearing much larger when in fruit, smooth or with
scattered, pellucid, gland-like cells, on pedicels mostly shorter than the flowers,
in glomerulate-cymose clusters; calyx about as long as the corolla tube, the lobes
overlapping at the base, but not markedly angled at the sinuses, ovate to oval-
ovate, commonly as long as wide, obtuse; corolla tube campanulate, soon enlarg-
ing about the maturing capsule, the lobes triangular to sublanceolate, about as
long as the tube, spreading to reflexed, often granulate, the tips acute, inflexed;
stamens shorter than the lobes; filaments slightly subulate, as long as or somewhat
longer than the oval anthers; infrastamineal scales reaching the filaments, ovate-
oblong, abundantly fringed with medium-length processes, bridged below the
middle; style slender to slightly subulate, as long as or longer than the globose
ovary; stigmas capitate; capsule depressed-globose, the withered corolla remaining
about the lower half; seeds ovate in outline, about 1.5 mm. long; hilum short,
oblong; embryo slender, with about 2 coils."
We have followed Prof. Yuncker in using the name C. campestris
for this plant, although we think it possible that the proper one may
be C. arvensis Beyrich.
Cuscuta corymbosa R. & P. var. grandiflora Engelmann,
Trans. Acad. St. Louis 1 : 483. 1859. Tripa de gallina; cabello de ledn;
pelo de ledn; bej'uco de mesquino; pelo de angel; cancam (Quecchi);
fideos; cabellos de angel; barba de ledn.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 15
FIG. 3. Cuscuta corymbosa var. grandiflora. A, habit of the parasite, natural
size; B, flower, X 5; C, corolla dissected to show stamens in natural position, X 5;
D, pistil, X 5.
Parasitic on shrubs or coarse herbs, 2,700 m. or lower; Alta
Verapaz; El Progreso; Jalapa; Escuintla; Guatemala; Sacatepequez ;
Chimaltenango; Solola; Suchitepe"quez; Retalhuleu; Quezaltenango.
Southern Mexico; El Salvador to Costa Rica; northwestern South
America.
Stems bright orange (as in other species), the corolla white; flowers 4-8 mm.
long, on pedicels of about equal length, in rather lax, corymbose-paniculate inflores-
16 FIELDIANA: BOTANY, VOLUME 24
cences; calyx about reaching the middle of the corolla tube, the lobes short, broad,
obtuse, slightly imbricate; corolla lobes one-fourth to one-half as long as the tube,
ovate-oblong, obtuse, erect or spreading; scales thin, narrow, reaching the middle
of the tube, dentate or with a few scattered processes; capsule globose, circum-
scissile, capped and surrounded by the withered corolla; seeds 1-1.5 mm. long.
The Maya name "canlecay" has been reported from Yucatan for
this or a related species. The vernacular names cited above are, of
course, applied indiscriminately to all species, and all are alike in
general appearance and color. In some parts of Guatemala, espe-
cially in the Pacific coastal plain, the plants often are seen in great
abundance, densely covering shrubs and tall herbs. The plants are
often used in Guatemala for dying textiles. Dieseldorff states that
about Coban the plants are employed in domestic medicine to treat
eruptions on the face. This variety has been reported from Guate-
mala as C. americana L.
Cuscuta costaricensis Yuncker, Mem. Torr. Club 18: 227. 1932.
Bejuco de mesquino. Central Mexico; Costa Rica.
Stems slender; flowers 3-4 mm. long, more or less papillate, subsessile, in dense
glomerulate clusters; calyx loose about the corolla tube, the lobes broadly ovate,
acute or acuminate, slightly imbricate; corolla lobes broadly ovate or suborbicu-
late, erect or spreading, shorter than the campanulate tube, acute or acuminate
or usually cuspidate; stamens shorter than the corolla lobes, the filaments equaling
or exceeding the oval anthers; scales oblong, reaching the stamens, fimbriate; styles
slender, much longer than the globose ovary; capsule depressed-globose, circum-
scissile; seeds 1.7 mm. long, ovoid.
Cuscuta cozumeliensis Yuncker, Bull. Torr. Bot. Club 49: 108,
/. 2. 1922.
Parasitic on shrubs, 600 m. or less; Suchitepequez; Retalhuleu.
Mexico, Yucatan, the type from Cozumel Island.
Stems rather coarse; flowers 2-3 mm. long, the pedicels equaling or shorter
than the flowers, in cymose clusters; calyx shorter than the corolla tube, the lobes
ovate-orbiculate, imbricate, somewhat carnose toward the base; corolla lobes erect
or spreading, imbricate, obtuse or rarely subacute, half as long as the cylindric tube;
stamens shorter than the corolla lobes, the filaments equaling or slightly longer
than the oval anthers; scales reaching the stamens or shorter, oblong, moderately
laciniate; styles slender, much longer than the depressed-globose ovary; capsule
circumscissile.
Cuscuta indecora Choisy, Mem. Soc. Phys. Hist. Nat. Geneve
9:278, LS.f.S. 1841.
On low herbs, near sea level; British Honduras (Malfredi Lagoon,
Schippll61). Mexico.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 17
Flowers 2-3 mm. long, fleshy, papillose-hispid, the pedicels longer than the
flowers; calyx lobes triangular-ovate, acute or obtuse, shorter than the corolla tube;
corolla campanulate, the lobes erect or spreading, triangular, acute, with inflexed
tips; scales equaling or longer than the corolla tube, ovate or subspatulate, deeply
fimbriate; stamens shorter than the corolla lobes, the anthers broad, oval, about
equaling the filaments; styles equaling or slightly longer than the globose pointed
ovary, divaricate in fruit; capsule globose, thickened at the apex, surrounded by
the withered corolla; seeds 1.7 mm. long, roundish or broader than long, some-
what scurfy.
Cuscuta jalapensis Schlecht. Linnaea 8: 515. 1833. Fideos;
cabellos de angel; barba de leon; coralillo; flor de cadena.
On shrubs and coarse herbs, 1,300-3,000 m.; Jalapa; Chimal-
tenango; Huehuetenango ; Totonicapan; Quezaltenango ; San Marcos.
Southern Mexico.
Stems of medium thickness, orange; flowers 2.5-5 mm. long, white or greenish
white, the pedicels equaling or shorter than the flowers, densely clustered; calyx
usually shorter than the campanulate corolla, the lobes imbricate, ovate, obtuse,
somewhat thickened and verrucose along the middle; corolla lobes ovate, obtuse,
shorter than the tube, erect or reflex ed; scales ovate-oblong, about equaling the
tube or exserted, fimbriate; stamens shorter than the corolla lobes, the anthers
ovate, about equaling the filaments; styles strongly subulate, mostly shorter than
the globose-conic ovary; capsule globose, circumscissile, surrounded by the with-
ered corolla, the styles conic, widely divergent; seeds 1.5 mm. long, ovoid.
This species seems to be confined in Guatemala to the higher
mountains, and often is found as high as the Cupressus forests. Near
Palojuno (Quezaltenango) the senior author's attention was at-
tracted one day by what he assumed to be great masses of shrubs
with orange flowers about a mile away. Walking across the fields to
see what this unknown and remarkably showy plant might be, he
was disgusted when he reached the spot to find that these solid
masses of color consisted of Cuscuta plants covering Sambucus bushes.
Cuscuta obtusiflora HBK. var. glandulosa Engelm. Trans.
Acad. St. Louis 1 : 492. 1859.
On low shrubs, 600 m. or less; Jutiapa (Steyermark 31988).
United States; Mexico; Costa Rica; West Indies.
Stems of medium thickness; flowers 2 mm. long, glandular, subsessile, in com-
pact glomerulate clusters; calyx enclosing the corolla tube, the lobes unequal,
rounded-ovate, obtuse, the margins usually irregularly serrulate, not imbricate;
corolla lobes triangular-ovate, acute or subobtuse, spreading or reflexed, shorter
than the campanulate tube; stamens shorter than the corolla lobes, the filaments
equaling or longer than the oval or cordate anthers; scales oblong, fimbriate at the
apex, scarcely reaching the stamens; styles rather stout and subulate, shorter than
18 FIELDIANA: BOTANY, VOLUME 24
the globose ovary or about equaling it; capsule depressed globose, indehiscent, the
styles subulate and divergent; seeds ovate, 1.5 mm. long.
Cuscuta rugosiceps Yuncker, Univ. 111. Biol. Mongr. 6, nos.
2-3: 117. /.J, 70,155. 1921.
On shrubs or coarse herbs, 1,600-3,300 m.; Sacatepe'quez ;
Quich^ ; San Marcos; Huehuetenango. Southern Mexico.
Stems coarse; flowers glabrous, 4-6 mm. long, sessile, in compact clusters;
calyx large, campanulate, almost as long as the corolla tube, the lobes short, usu-
ally broader than long, unequal, obtuse, imbricate, sometimes carinate; corolla
lobes ovate, obtuse, spreading, shorter than the campanulate tube; scales reaching
the filaments, fimbriate; stamens shorter than the corolla lobes, the anthers oval,
about equaling the filaments; ovary small, somewhat conic, tapering into the subu-
late styles, the stigmas large and more or less convolute; capsule circumscissile,
with a very thick apex, giving the capsule a conic appearance, surrounded by the
withered corolla; seeds rounded, compressed, 1.4 mm. long.
Cuscuta saccharata (Engelm.) Yuncker, Mem. Torrey Club 18:
239. 1932. C. gracillima var. saccharata Engelm. Trans. Acad. Sci.
St. Louis 1:489. 1859.
Parasitic on herbs, 200-550 m.; Zacapa; Chiquimula. Southern
Mexico; Honduras; Costa Rica.
Stems very slender; flowers 2-2.5 mm. long, densely papillate (as also the
stems, pedicels and bracts), the slender pedicels longer than the flowers, in dense
compact clusters; calyx about equaling the corolla tube, the lobes ovate-lanceolate,
acute or acuminate; corolla lobes erect or spreading, about equaling the campanu-
late tube; stamens shorter than the corolla lobes, the slender filaments longer than
the oval anthers; scales ovate, reaching the stamens, short-fimbriate; styles slender,
longer than the globose ovary; capsule globose, thin, irregularly circumscissile;
seeds oval, 1 mm. long.
Cuscuta tinctoria Mart, ex Engelm. Trans. Acad. Sci. St. Louis
1:480. 1859.
On shrubs or trees; Sacatep6quez ; Quiche^ Totonicapan (Lehmann
1682). Mexico.
Flowers smooth, 4-5 mm. long, sessile or short-pedicellate, single or in dense
glomerules of many flowers; calyx lobes unequal, orbicular, obtuse, imbricate,
about equaling the corolla, sometimes carinate; corolla campanulate, becoming
urceolate in fruit, thinner toward the base, the lobes ovate or almost oblong, ob-
tuse, imbricate, erect or usually spreading; scales reaching the filaments; stamens
shorter than the corolla lobes, the filaments about equaling the oval-oblong anthers;
styles usually longer than the globose ovary, exserted in fruit; capsule depressed-
globose, circumscissile; seeds 1.5 mm. long, angulate.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 19
Cuscuta tinctoria var. kellermaniana Yuncker, Univ. 111.
Biol. Monogr. 6, nos. 2-3: 32, /. 16. 1921.
Known only from the type, Volcan de Agua, Sacatepe'quez,
2.700 m., W. A. Kellerman 7567; on Ceanothus.
Like the species, but the scales oblong, sparsely fimbriate; styles
shorter than the depressed-globose ovary and capsule, not exserted.
Cuscuta yucatana Yuncker, Bull. Torr. Bot. Club 62: 511.
1935. Coralillo.
On Houstonia serpyllacea, 2,500 m.; Jalapa (Montana Mira-
mundo, Steyermark 3264.8) . Known otherwise only from Valladolid,
Yucatan.
Stems slender; flowers 2 mm. long, the pedicels equaling or longer than the
[lowers, in compound umbellate cymes; calyx longer than the corolla tube, the
lobes triangular-ovate, acute, not imbricate; corolla lobes oblong-lanceolate, very
sharply acute, erect in young flowers, the tips inflexed, soon becoming reflexed, the
lobes equaling or longer than the campanulate tube, more or less grandular-papil-
ate; filaments slender or slightly subulate, mostly 2-3 times as long as the oval-
elliptic anthers; scales oblong or subspatulate, reaching the stamens, fimbriate;
styles equaling or longer than the depressed-globose ovary; capsule depressed-
globose, not circumscissile; seeds 0.8-1 mm. long, oval.
The distribution of this species is remarkable, for it would be hard
to imagine two regions more unlike than the localities from which the
slant is known at present. Even though both specimens known, the
type and Steyermark 32648, have been determined by Prof. Yuncker
*ve are inclined to suspect that the Guatemalan collection represents
mother species.
DICHONDRA Forster
Reference: B. C. Tharp & Marshall C. Johnston, Recharacteriza-
:ion of Dichondra and a revision of North American species, Brit-
tonia 13: 346-360. 1961.
Slender perennial herbs, glabrous or sericeous; leaves small, cordate-orbicular
)r reniform, entire, long-petiolate; flowers minute, solitary and pedicellate in the
eaf axils; sepals subequal, distinct, usually spatulate; corolla broadly campanu-
ate, deeply 5-fid, the lobes induplicate; stamens shorter than the corolla, the
^laments filiform, the anthers small; ovary bilobate, the lobes distinct, 2-celled,
jiovulate; styles 2, between the lobes, filiform, the stigmas capitate; capsules 2,
nembranaceous, erect, usually 1-seeded, indehiscent or irregularly bivalvate; seeds
mbglobose, smooth; cotyledons oblong-linear, 2-plicate.
Perhaps a dozen species, widely distributed in tropical and warm-
:emperate regions. Only the following is known from Central
\merica.
20 FIELDIANA: BOTANY, VOLUME 24
Dichondra sericea Sw. Prodr. Veg. Ind. Occ. 54. 1788. D. repens
var. sericea Choisy in DC. Prodr. 9: 451. 1845.
Usually in moist or dry open, often rocky slopes in open forest,
sometimes in moist shaded habitats, 1,200-2,500 m. Alta Verapaz;
Chiquimula; Jalapa; Jutiapa; Escuintla; Guatemala; Sacatepe"quez ;
Chimaltenango; Solola; Huehuetenango; Quezaltenango ; San Mar-
cos. Arizona; Mexico; Central America; the West Indies and South
America.
Plants very slender, repent, rooting at the nodes, the stems usually short but
sometimes 30 cm. long or more, appressed-pilosulous or glabrate; leaves long-petio-
late, reniform or cordate-orbicular, mostly 8-20 mm. broad, broadly rounded at
the apex and often emarginate, deeply cordate at the base, green on both surfaces,
sparsely pilosulous on both surfaces with subappressed hairs to sericeous below;
peduncles shorter than the petioles, filiform; sepals oblong or spatulate-oblong,
obtuse, green, pubescent, about equaling the corolla, shorter than the capsule,
about 2 mm. long; corolla yellowish; capsules densely pilose.
We have followed the work of Tharp and Johnston, cited above,
with some reservations. Their work is based on relatively little of
the material now available and limited in extent — consequently open
to some question.
EVOLVULUS L.
Reference: Simon Jan van Ooststroom, A monograph of the genus
Evolvulus, in Med. Bot. Mus. & Herb. R. Univ. Utrecht 14: 1-267.
1934.
Plants usually slender, annual or perennial, herbaceous or suffrutescent, not
twining, sometimes creeping; leaves small, often narrow, entire; flowers in axillary,
peduculate, 1-several-flowered dichasia, sometimes solitary, pedunculate or sessile
in the leaf axils; sepals 5, free, equal or subequal; corolla small, rotate, funnelform,
or salverform, blue or white, the limb plicate, usually subentire, the lobes with a
pilose longitudinal stripe outside; stamens 5, the filaments filiform, inserted at the
mouth of the corolla tube; anthers ovate to oblong or linear; ovary 2-celled, each
cell 2-ovulate, sometimes 1-celled and 4-ovulate; styles 2, slightly united at the
base or wholly free, each style 2-cleft; stigmas long, terete, filiform or subclavate;
capsule globose or ovoid, 4-valvate, 4-1-seeded; seeds small, smooth or minutely
verrucose; cotyledons almost flat, the radicle incurved.
About 100 species, widely distributed in both hemispheres,
mostly in the warmer regions.
Inflorescence not pedunculate, the flowers solitary or fasciculate from leaf axils.
Plants repent, the stems rooting at the nodes; leaves broadly oval or orbicular.
E. nummularius.
Plants erect or ascending; leaves oblong-ovate to linear.
Leaves oblong-ovate; stems pilose with long, lax, spreading hairs. . .E. ovatus.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 21
Leaves linear to oblong; stems strigose or sericeous E. scricem.
Inflorescence pedunculate, axillary or sometimes terminal.
Pubescence of the stem spreading, leaves variable.
Corolla 3-7 mm. broad E. alsinoides.1
Corolla 10 mm. broad, the leaves 12-20 mm. broad E. tennis.1
Pubescence of stem appressed E. filipes.1
Evolvulus alsinoides L. Sp. PL ed. 2. 392. 1762. Convolvulus
alsinoides L. Sp. PL 157. 1753. C. linifolius L. Amoen. Acad. 4: 306.
1759. Quiebra-cajete; cenicito.
Usually in open, moist or dry places, sometimes in thickets or on
rocky banks, often in pine forest, frequently a weed in cultivated
ground, 200-1,900 m.; Baja Verapaz; Alta Verapaz; Izabal; Jalapa;
Jutiapa; Santa Rosa; Escuintla; Guatemala; Sacatepe"quez ; Chimal-
tenango; Solola; Suchitep£quez; Huehuetenango. Southwestern
United States; Mexico; British Honduras to El Salvador and Pana-
ma; widely distributed in temperate and tropical America and in the
Old World.
Plants slender, annual or perhaps sometimes perennial, erect to prostrate, often
much branched, pilose with appressed or spreading hairs; leaves mostly oblong to
elliptic-oblong or lanceolate and 1-2.5 cm. long, usually 1 cm. broad or less, on
very short petioles, obtuse or acute, rounded or acute at the base, generally rather
densely appressed-pilose, especially beneath; inflorescence axillary, shorter than
the leaves or often much longer, 1-few-flowered, peduncles almost filiform; pedicels
equaling or longer than the calyx; sepals lanceolate, acute or acuminate, 1.5-3 mm.
long, pilose; corolla pale blue or sometimes white, 3-5.5 (-7) mm. broad; ovary
glabrous; capsule globose, with 4 or fewer seeds, these black, smooth.
Called "oreja de raton" in El Salvador; "xiatiu" (Yucatan,
Maya).
S. J. Ooststroom indicates that there are four varieties of this
species in Guatemala: var. griesbachianus Meissn., var. debilis (HBK.)
v. Ooststr., var. adscendens (House) v. Ooststr., and var. acapulcensis
(Willd.) v. Ooststr. The differences seem slight if not non-existent.
Van Ooststroom recognized 15 varieties of this species, world-wide.
Evolvulus filipes Mart. Flora 24, Beibl. 2: 100. 1844.
Moist shaded places, or sometimes a weed in cultivated fields,
200-2,000 m.; Zacapa; Jalapa; Jutiapa; Santa Rosa; Guatemala;
Huehuetenango. Mexico; Honduras to Panama; Jamaica; South
America.
1 The junior author is not at all sure that the Central American material tradi-
tionally named as E. alsinoides and E. filipes represents other than a single weedy
and somewhat variable species. Evolvulus tennis likewise seems doubtfully distinct
from E. alsinoides, as represented by Central American material.
22 FIELDIANA: BOTANY, VOLUME 24
Plants annual, erect or ascending, usually 30 cm. high or less, often densely
branched, the stems sparsely appressed-pilose; leaves sessile or on very short peti-
oles, linear or narrowly lanceolate, 1-2.5 cm. long, mostly 2-5 mm. broad, acute or
subobtuse, acute at the base, very sparsely pilose or glabrous above, sparsely ap-
pressed-pilose beneath; peduncles longer or shorter than the leaves, mostly 1-2-
flowered, the pedicels equaling or shorter than the calyx, reflexed in fruit; sepals
lanceolate, glabrous or sparsely pubescent, ciliate, 2-2.5 mm. long; corolla pale
blue or white, at most twice as long as the sepals, 3-4.5 mm. broad; ovary glabrous;
capsule little exceeding the sepals, globose or ovoid, with 4 or fewer seeds, these
brownish black, smooth.
The Maya name of Yucatan is recorded as "yax-cumil."
Traditionally E. filipes has been considered to be distinct from
E. alsinoides, having smaller flowers, narrower more glabrous leaves,
pubescence of the stems appressed. The junior author finds that
these characters are not consistent and believes that so far as the
Central American material is concerned only one somewhat variable
and weedy species is involved. Until monographic work can be done
it would be as well to name all pedunculate Evolvulus in Central
America as E. alsinoides.
Evolvulus nummularius L. Sp. PL ed. 2. 391. 1762. Convolvu-
lus nummularius L. Sp. PI. 157. 1753. Volvulopsis nummularium
Roberty, Candollea 14: 28. 1952. Hierba de pesar (Pete"n, fide
Lundell) ; cuartillo.
Moist shaded places, often among cobblestones in streets, fre-
quent on roadside banks or a weed in cultivated fields, 1,200 m. or
less; Pete"n; Izabal; Zacapa; Chiquimula; Jutiapa; Santa Rosa;
Escuintla; Guatemala; Suchitepe"quez; Retalhuleu. Mexico; British
Honduras to El Salvador and Panama; West Indies; South America;
Old World tropics.
Plants perennial, herbaceous, repent, rooting at the nodes, the stems short or
often 50 cm. long, pilose with short spreading curved hairs; leaves distichous,
short-petiolate, broadly oval to orbicular, 4-15 mm. long, rounded or emarginate
at the apex, rounded or subcordate at the base, glabrous or often sparsely appressed-
pilose beneath; flowers 1-2 in each leaf axil, the pedicels 2-6 mm. long, recurved in
fruit; sepals 2.5-4 mm. long, ovate-oblong, obtuse or subacute, sparsely pilose or
glabrous, ciliate; corolla white, 5-7 mm. broad; ovary glabrous; capsule globose,
equaling or slightly longer than the sepals, with 4 or fewer seeds.
Evolvulus ovatus Fernald, Proc. Amer. Acad. 33: 89. 1898.
Dry brushy plains or hillsides, 200 m. ; Zacapa. Western Mexico ;
Colombia and Venezuela to Brazil.
FIG. 4. Evolvulus sericeus. A, habit of plant, natural size; B, flower with
corolla in natural position, X 3; C, calyx, X 4.
23
24 FIELDIANA: BOTANY, VOLUME 24
Plants annual, erect, 30 cm. high or less, sparsely branched, the stems villous
with long spreading lax hairs; leaves short-petiolate or the upper ones sessile, ob-
long-ovate, 1-3 cm. long, obtuse or subacute, rounded or subcordate at the base,
sparsely appressed-pilose on both sides with very long, lax hairs; inflorescence
axillary, flowers 1-2, the pedicels shorter than the calyx, reflexed in fruit; sepals
lanceolate, acute, 4-5 mm. long, pilose; corolla blue, 5 mm. broad; ovary glabrous;
capsule depressed-globose, shorter than the sepals, 4-seeded; seeds smooth, dark
brown.
Evolvulus sericeus Swartz, Prod. Veg. Ind. Occ. 55. 1788.
Savannas or open grassy places, sometimes on rocky slopes, on
serpentine outcrops, or in pine forest, 2,000 m. or less; Pete"n; Alta
Verapaz; Zacapa; Chiquimula; Santa Rosa; Huehuetenango. South-
western United States; Mexico; British Honduras to El Salvador and
Panama; West Indies; South America.
Plants perennial, erect or ascending, mostly 25 cm. high or less, often densely
branched from the base, sometimes suffrutescent below, the stems slender but stiff,
grayish-sericeous; leaves erect or erect-spreading, sessile or nearly so, linear or
linear-lanceolate or elliptic, sericeous, at least beneath, sometimes glabrous on the
upper surface, 1-2.5 cm. long, 1-8 mm. broad; flowers solitary or few in the leaf
axils, sessile or short-pedicellate, reflexed in age; sepals narrowly oblong-lanceolate,
acuminate, 3-5 mm. long, densely appressed-pilose; corolla white or pale blue,
7-12 mm. broad; ovary glabrous; capsule globose, 1-4-seeded; seeds brown, smooth.
The common form of the species in Guatemala has leaves pubes-
cent on both surfaces. Two collections are referable to the following
variety:
Evolvulus sericeus var. discolor (Benth.) Gray, Syn. Fl. 2,
pt. 1: Suppl. 436. 1886. E. discolor Benth. PL Hartw. 6. 1839.
Grassy open slopes, 2,200 m.; Huehuetenango. Southwestern
United States; Mexico.
Differing from the common form (in Guatemala) of the species
in having broader leaves which are glabrous rather than pubescent
on the upper surface.
Evolvulus tenuis Mart, ex Choisy, Mem. Soc. Phys. Hist. Nat.
Geneve 78. 1837.
Dry or moist, brushy slopes, 1,400 m. or less; Chiquimula; Quezal-
tenango(?). Yucatan; Costa Rica; South America.
Plants erect or ascending, annual or somewhat suffrutescent at the base, the
stems sometimes more than a meter long, rather sparsely branched, the stems
densely pilose with long pale ascending hairs; leaves mostly ovate or oblong-ovate
and 1.5-3.5 cm. long, 1-2 cm. broad, acute or acuminate, sparsely or often densely
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 25
appressed-pilose; peduncles longer or shorter than the leaves, 1 few-flowered the
pedicels usually longer than the calyx; sepal lanceolate, acute or acuminate, 2. 5
3 mm. long; corolla blue or white, 10 mm. broad, ovary globose, glabrous.
This is presumably the plant reported by Hemsley from Llano de
San Juan de Dios, Quezaltenango, as E. villosus Ruiz & Pavon,
Bernoulli 181. The species is very closely related to E. alsinoides L.
IPOMOEA L.
References: Homer Deliver House, The North American Species
of the genus Ipomoea, Ann. N. Y. Acad. Sci. 18: 181-263. 1908.
Eizi Matuda, El Ge"nero Ipomoea en Mexico, An. Inst Biol 24-
85-145. 1964, 25: 45-76. 1965, 26: 83-106. 1965.
Twining or creeping, rarely erect herbs or shrubs, rarely trees. Leaves alter-
nate, entire, angled, lobate, or palmately or pinnately divided, usually petiolate;
the flowers usually large and showy, usually fugacious; the inflorescences usually
axillary, rarely aggregated in terminal panicles; sepals coriaceous or foliaceous,
equal or unequal, persistent; corolla funnelform or campanulate, rarely salverform,'
the limb entire or 5-angulate, rarely 5-lobate, usually induplicate; stamens in-
cluded, rarely exserted; pollen said to be spinulose; ovary 2-4-celled; style included,
the stigmas 1-2, globose-capitate; capsule 2-4-valvate, 2-4-seeded; seeds often
densely lanate or pubescent.
The genera Quamoclit, Turbina, Exogonium, and Calonyction are
sometimes included in Ipomoea but are kept separated in this flora—
however, in each case names in Ipomoea are provided in the synonomy .
There are perhaps not fewer than 400 species of Ipomoea which
are widely distributed, the majority of them in tropical regions.
There are currently credited to Mexico, Central America, and
Panama, in our index to this region, about 250 species (and a few
varieties). There are to be expected from Guatemala additional
species as collections continue, for this genus is difficult to collect and
has been neglected.
Two accounts of Ipomoea are useful in the study of the Guate-
malan species, both mentioned above. Mr. House's treatment was
prepared more than 60 years ago when Central America was rela-
tively little known. Professor Matuda's account of the Mexican
species is helpful for those of Guatemala.
Species of Ipomoea seem to be widely ranging and there is little
doubt that when the South American species have been compared
more critically with those of North America some changes will be
required in the nomenclature used here. Many Ipomoeas occur at
relatively low elevations and experience indicates that it is from
these regions that most wide-ranging plants are found.
26 FIELDIANA: BOTANY, VOLUME 24
The Ipomoeas produce some of the showiest and most beautiful
flowers of Guatemala, and occur in great abundance almost anywhere
at low and at middle elevations, more abundantly in the foothills and
on the plains. In some places along the foothills or lowlands of
Escuintla and Santa Rosa one rides for miles in the morning with
solid sheets of morning-glory flowers on each side of the road. The
flowers of all or most species open in the evening and close usually
before noon. In the higher parts of central and western Guatemala
there are few Ipomoeas, or only an occasional plant may be found.
The flowers are much visited by bees. The people of Chiquimulilla
(Santa Rosa) claim that the superior flavor of honey produced there
is due to the fact that it comes largely from Ipomoea flowers. The
most common vernacular name current in Guatemala for this genus
is "campanilla," but other terms, such as "quilamul," sometimes are
used, and even "alegria de la manana," which is probably a transla-
tion of the usual English name.
KEY TO IPOMOEA
Plants not twining, erect or climbing shrubs or trees, or sometimes almost wholly
herbaceous.
Leaf blades obtuse or rounded at the base; trees; corolla white; seeds with white
hairs on the dorsal angles only.
Sepals and corolla glabrous 7. pauciflora.
Sepals and corolla pubescent.
Corolla and sepals tomentose outside; sepals 2-3 cm. long ..../. murucoides.
Corolla and sepals puberulent; sepals 1 cm. long or shorter . . .7. arborescens.
Leaf blades cordate or subcordate at the base; corolla white or purple-pink;
seeds with black hairs on all surfaces 7. fistulosa.
Plants twining or repent, usually herbaceous throughout or nearly so.
Sepals herbaceous, at least above, sometimes subcoriaceous but then covered
with long, fleshy, subulate tubercles, often elongated and densely pilose;
ovary usually 3-celled.
Sepals covered with long spreading setae or fleshy tentacular outgrowths.
Leaves deeply lobate, the lobes 3-7.
Sepals oblong, obtuse, 10-14 mm. long 7. setosa.
Sepals linear-lanceolate, about 25 mm. long 7. silvicola.
Leaves entire.
Sepals all long-attenuate 7. silvicola.
Sepals, at least the inner ones, very obtuse 7. crinicalyx.
Sepals merely pilose or hirsute, or glabrate, never with fleshy outgrowths.
Corolla copiously pubescent outside; leaves densely sericeous beneath.
7. mairetii.
Corolla glabrous; leaves rarely if ever sericeous.
Sepals glabrous or with only short appressed hairs 7. indica.
Sepals hirsute, at least below, with long spreading hairs.
Corolla small, only 2-3 cm. long; inflorescences dense and head-like,
much shorter than the leaves 7. meyeri.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 27
Corolla large, 4-8 cm. long; inflorescences usually open, often long-
pedunculate.
Sepals abruptly contracted into a long linear tip, 2-3 cm. long; leaves
trilobate 7. nil.
Sepals acute or gradually acuminate, usually shorter.
Inflorescences short-pedunculate; stems usually glabrous; sepals
acuminate 7. indica var. variabilis.
Inflorescences long-pedunculate; stems usually hirsute; sepals
merely acute or sometimes obtuse.
Stems creeping, rooting at the nodes, densely long-hirsute; sepals
lance-linear 7. ophioides.
Stems scandent, not rooting at the nodes, short-hirsute; sepals
broader.
Inner sepals mostly 12-15 mm. long, obtuse; leaves entire;
corolla 6-8 cm. long 7. tyrianthina.
Inner sepals mostly 9-12 mm. long, acute; leaves trilobate or
entire; corolla 4-6 cm. long 7. purpurea.
Sepals coriaceous, membranaceous, or obscurely herbaceous, usually not elon-
gate, most often glabrous but sometimes pubescent; ovary usually 2- or 4-
celled.
Leaves deeply palmate-lobate or pinnate-lobate almost to the base or to the
costa, the lobes narrow; plants erect or usually twining.
Leaves sessile, divided to the base into filiform segments; plants usually
not twining 7. capillacea.
Leaves petiolate, the segments linear or often much broader; plants usually
twining.
Lobes of the leaves less than 4 cm. long; small vines.
Sepals cuspidate; peduncle not coiled 7. costellata.
Sepals rounded at the apex; peduncles usually spirally coiled at the
base 7. pulchella.
Lobes of the leaves 5-8 cm. long; large vines.
Leaves parted to the base, the terminal lobe 3-parted .... 7. hete.rodoxa.
Leaves not parted to the base, the lobes all entire 7. digitata.
Leaves entire, or lobate to the middle or usually much less deeply, rarely more
deeply lobate (7. stolonifera) but the plants then creeping.
Plants prostrate or creeping, not twining, herbaceous.
Leaf blades cordate or sagittate at the base.
Sepals pilose or ciliate, about 2 mm. long 7. minutiflora.
Sepals glabrous, 5-10 mm. long.
Leaves sagittate 7. reptans.
Leaves rounded-cordate 7. asarifolia.
Leaf blades rounded at the base.
Flowers white; leaves mostly oblong, sometimes deeply lobate, mostly
1-2 cm. wide 7. stolonifera.
Flowers pink or purple; leaves suborbicular, mostly 6 cm. wide or more,
never lobate 7. pes-caprae.
Plants twining or trailing, at least the tips twining.
Sepals in an thesis 5 mm. or less long; corolla blue to purple or yellow.
Corolla blue or pink to purple.
Calyx rugose on dorsal veins; corolla blue or purple, about 20-25 mm.
long 7. aristolochiaefolia.
Calyx lobes not rugose; corolla pink, about 15 mm. long. .7. perplexa.
Corolla yellow, 8-25 mm. long.
28 FIELDIANA: BOTANY, VOLUME 24
Corolla about 8 mm. long; leaves mostly as broad as long, rounded
at the apex or abruptly acute; plants often prostrate.
/. minuti flora.
Corolla 20-25 mm. long; leaves longer than broad, acute or acumi-
nate; plants twining /. microsepala.
Sepals in anthesis 5 mm. long or usually much longer; corolla rarely
yellow.
Corolla yellow or orange.
Calyx lobes elliptic or oblong, subequal, 3-4 mm. broad . . . /. flavida.
Calyx lobes broadly ovate to suborbicular, unequal, more than 4 mm.
broad /. aurantiaca.
Corolla never yellow.
Leaves densely and conspicuously tomentose or sericeous beneath.
Corolla glabrous or essentially so.
Sepals aristate at the apex /. leucotricha.
Sepals rounded at the apex, not aristate.
Sepals sericeous /. tuxtlensis.
Sepals glabrous.
Leaves sagittate; anthers near the base of the corolla.
I. squamosa.
Leaves not sagittate; anthers near middle of corolla.
Leaves cuneate at the base; flowers red or carmen.
/. steerei.
Leaves cordate at base.
Leaves suborbicular-ovate; flowers lilac /. teruae.
Leaves trilobate; flowers white with red, or red.
/. tuxtlensis.
Corolla copiously, often densely pilose.
Sepals 5-15 mm. long; leaves glabrous or only slightly pubescent
above.
Sepals narrowly lanceolate, acuminate; stems muriculate,
pilose with brownish hairs; corolla purple. . . ./. villifera.
Sepals suborbicular; stems pale, whitish tomentose at first;
corolla violet-pink or white /. carnea.
Sepals almost or quite 2 cm. long; leaves densely whitish-
tomentose on both surfaces /. ptaecana.
Leaves glabrous beneath or inconspicuously pubescent, never densely
sericeous or tomentose.
Sepals conspicuously aristate at the apex.
Sepals minutely aculeolate along the costa; petioles minutely
aculeolate; leaves triangular-hastate, entire. . . ./. setifera.
Sepals and petioles not aculeolate; leaves not triangular-hastate.
Sepals hirsute or at least ciliate I. triloba.
Sepals glabrous.
Stems mostly creeping and rooting, only the tips of the
stems twining; plants usually not flowering. ./. batatas.
Stems not rooting at the nodes, twining throughout, the
plants flowering abundantly.
Corolla about 2-3 cm. long; leaves usually lobate.
/. trifida.
Corolla 3.5-7 cm. long; leaves usually not lobate.
Sepals 1 cm. long or shorter.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 29
Corolla 3.5-5 cm. long; stamens inserted below mid-
dle of the corolla; outer calyx lobes cochleate.
/. tiliacea.
Corolla 6-7 cm. long; stamens inserted near the
throat; outer calyx lobes not cochleate.
/. seducta.
Sepals 1.3 cm. long or longer.
Sepals oblong to oval, obtuse; peduncles about 1 cm.
long /. santae-rosac.
Sepals narrowly lanceolate, subaristate; peduncle
essentially none 1. sessilis.
Sepals not aristate at the apex, usually rounded or obtuse.
Stems setose or setose-hispid.
Calyx lobes glabrous.
Calyx lobes ovate, obtuse, unequal, 10-20 mm. long.
/. sepacuitensis.
Calyx lobes lanceolate to elliptic, acute, subequal, 25-30
mm. long /. contrerasii.
Calyx lobes setose /. setosa.
Stems not setose nor setose-hispid.
Stems woody throughout; plants glabrous; corolla white.
/. pauci flora.
Stems wholly or chiefly herbaceous.
Flowers subtended by a large leaf-like reniform bract 2-5
cm. long /. suffulta.
Flowers never subtended by a large leaf like bract.
Sepals in anthesis about 5 mm. long, narrow, lanceolate
or ovate-lanceolate, imbricate only below.
Corolla glabrous outside, 5-6 cm. long; sepals mostly
oblong-lanceolate, white marginate ..../. tricolor.
Corolla puberulent outside, about 3 cm. long; sepals
lanceolate- ovate /. parasitica.
Sepals in anthesis usually longer, mostly oval to orbicu-
lar, sometimes elliptic-oblong, usually strongly im-
bricated for most their length.
Upper leaves sessile or short-petiolate; corolla salver-
form /. purga.
Upper leaves all or mostly long-petiolate; corolla fun-
nelform.
Stems, at least below, bearing numerous fleshy tuber-
cles; sepals 6 mm. long /. parasitica.
Stems not tuberculate.
Leaves sagittate, the terminal portion oblong or
triangular to almost linear.
Sepals very unequal, the outer ones less than
half as long as the inner; peduncles several-
flowered . . . /. anisomeres var. sagittiformis.
Sepals subequal, the outer ones but little shorter
than the inner; peduncles mostly 1-flow-
ered /. sagittata.
Leaves narrowly to broadly ovate, cordate.
Corolla 5-8 cm. long, white, pink, purple, or
brown-purple.
Sepals 12-18 mm. long.
30 FIELDIANA: BOTANY, VOLUME 24
Peduncle less than 2 cm. long; inflorescence
1-2-flowered /. armentalis.
Peduncle 10-20 cm. long; inflorescence usu-
ally many-flowered. . . ./. demerariana.
Sepals 6-10 mm. long.
Stems conspicuously papillate on older
growth; stamens only a third as long as
corolla /. anisomeres.
Stems smooth; stamens extending to the
middle of the corolla or beyond.
Seeds long-pectinate-pilose; corolla
brown-purple /. armcnlalis.
Seeds glabrous or short-pubescent; corolla
rose-purple or white.
Corolla purple; leaves glabrous or
nearly so, or sparsely pubescent.
Peduncles 0-5 mm. long. ./. lindenii.
Peduncles 70 mm. long or usually
much longer.
Corolla campanulate or funnel-
form; anthers not reaching
the middle of corolla.
/. squamosa.
Corolla salverform; anthers reach-
ing the throat or exserted.
/. signata.
Corolla white; leaves densely hirtellous
beneath /. saxorum.
Corolla about 4-4.5 cm. long, sometimes longer,
usually white, sometimes tinged with pink
or yellow.
Stems woody with age; under surface of leaf
with microscopic black dots; sinus at
base of leaf shallow /. microsticia.
Stems herbaceous; under surface of leaves
without black dots.
Sinus at base of leaf shallow; flowers yellow.
/. flavida.
Sinus at base of leaf deep ; flowers not yel-
low 7. wallii.
Ipomoea anisomeres Rob. & Bartl. Proc. Am. Acad. 43: 57.
1907.
Moist or wet thickets at 500 m. or less; Zacapa (type, Deam 318);
Pet^n (Contreras ^37). Possibly British Honduras. Mexico (Vera-
cruz).
A slender herbaceous vine, glabrous, the stems to about 3 mm. in diameter,
smooth at first becoming prominently papillate with age. Leaves oblong-ovate to
broadly ovate, acute or acuminate, cordate and the basal lobes broad and rounded,
petioles slender, 3-6.5 cm. long; inflorescence several-many-flowered corymbs or
cymes, the peduncles 4-14 cm. long, the pedicels slender, and up to 2.5 cm. long,
slightly thickened above; calyx coriaceous, up to about 1 cm. long, the lobes very
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 31
unequal, the outer ones orbicular or transversely oval, 1-3 mm. long, the inner
lobes oblong to ovate, obtuse, 8-10 mm. long at an thesis; corolla white with a
purple throat, narrowly infundibuliform, about 7 cm. long, the limb 4-5 cm. broad;
stamens unequal in length, one-third as long as the corolla or less; style as long as
the stamens; capsule unknown.
Two numbers, Deam 318 and 319 are cited as types. We select
318 as the lectotype in Gray Herbarium.
Ipomoea anisomeres var. sagittiformis L. Wms. Fieldiana,
Bot. 32:185. 1970.
Trailing on the ground or in thickets, 600 m. or less, Izabal (type
Steyermark 381*85}. Endemic.
Like the typical variety described above except: leaves smaller,
sagittiform, pubescent, about 8 cm. long with the terminal lobe
linear-oblong, about 1-2 cm. broad; the exterior calyx lobes 3-5 mm.
long; the corolla purple. The capsule is suborbicular, about 1 cm.
long, the seeds subvelutinous with a longer sericeous coma along the
lateral margins; these last characters not known for the typical
variety.
Ipomoea arborescens (Humb. & Bonpl.) G. Don, Gen. Syst. 4:
267. 1838. Convolvulus arborescens Humb. & Bonpl. ex Willd. Enum.
PL 1:204. 1809.
Dry, rocky thickets, 900 m.; Chiquimula; Zacapa; Jutiapa (near
Jutiapa, Standley 60491). Western and southern Mexico. El Sal-
vador.
Small tree, the branches densely and minutely puberulent, the bark whitish;
leaves slender-petiolate, ovate or oblong-ovate, mostly 5-14 cm. long, acute or
acuminate, shallowly cordate at the base, finely pubescent above or glabrate,
densely and minutely pubescent beneath, the veins conspicuous and closely reticu-
late; peduncles usually short and several-flowered; sepals oval, obtuse or rounded
at the apex, 8-12 mm. long, tomentulose; corolla white, 5 cm. long, glabrous or
glabrate outside; capsule ovoid, 2 cm. long; seeds black, the angles densely long-
pilose.
Three collections of this species have been made in Guatemala,
one by the senior author, and on the label it is stated that the tree is
abundant about Jutiapa. The specimen is immature and may well
be this species, which is known to occur on both sides of Guatemala,
or it could be the common I. murucoides. The peduncles in /.
arborescens are usually several flowered while those of /. murucoides
usually bear a single flower.
32 FIELD IANA: BOTANY, VOLUME 24
Ipomoea aristolochiaefolia (HBK.) G. Don, Gen. Hist. PI. 4:
277. 1838; O'Donell, Lilloa 29: 114, fig. 4. 1959. Convolvulus aris-
tolochiaefolia HBK. Nov. Gen. & Sp. PI. 3: 102. 1819. I. tuerckheimii
Vatke ex Donn.-Sm. Bot. Gaz. 40: 8. 1905 (type from Coban,
Tuerckheim 386} . Campanilla; quiebra cajete.
Moist thickets, 600-2,000 (?) m.; Alta Verapaz; Jalapa; Santa
Rosa; Zacapa. Mexico; El Salvador; Costa Rica; Venezuela; Colom-
bia and Ecuador to Argentina.
A slender vine, the stem puberulent with short, erect hairs, becoming glabrous,
internodes 5-10 cm. long. Leaves with short petioles 1-5 cm. long, the blades
5-8 cm. long and 2-4 cm. broad, cordate, acuminate, glabrous or sparsely puberu-
lent above; peduncle very slender, 3-6 cm. long, puberulent or not, often passing
between the basal lobes of leaf, usually several-f owered; sepals oval, acute or ob-
tuse, rugose on the dorsal veins, nearly equal or the outer ones shortest, mostly
3-5 mm. long; corolla campanulate, glabrous, 20-25 mm. long; ovary conic, bi-
locular, the style longer than the stamens; capsule ovoid, 1 cm. long, seeds pubes-
cent, black or brown.
Ipomoea armentalis L. Wms. Fieldiana, Bot. 32: 185. 1970.
Thickets, cornfields and forest edges, 500-1,200 m.; Chiquimula;
Jutiapa (type, Standley 76005); Escuintla; Huehuetenango. Mexico
(Chiapas). Madre de maiz; campanula.
Delicate twining herbaceous or suffrutescent vines with puberulent or glabres-
cent stems to about 1 mm. in diameter. Leaves oblong-ovate to ovate, cordate,
acuminate, pilose-puberulent on both surfaces, the blade 3-8 cm. long and 1.5-
4.5 cm. broad, petiole slender, shorter than the blade; inflorescence axillary, 1-
(rarely 2-) flowered, longer than the subtending leaf, peduncle 0.1-1.2 cm. long,
puberulent, the slender pedicels 1-4 cm. long, puberulent; calyx about 1.5 cm.
long at anthesis and in fruit, coriaceous, glabrous, the lobes unequal to subequal,
oblong or oblong-oblanceolate, obtuse, 8-12 mm. long, with the outer lobes often
shorter than the inner ones; corolla dark purple to blue, narrowly campanulate,
about 5-6 cm. long, tip of bud puberulent; capsule lanceolate-ovoid, the thick style
base persisting, 1.5-2 cm. long; seeds subtrigonous, black, puberulent over the
whole surface, except the lateral margins with long coma of comb-like hairs, about
9 mm. long and 4 mm. broad.
The only species in Guatemala with the margins of the seeds
pectinate-ciliate; the one-flowered inflorescences are uncommon in
the genus.
Ipomoea asarifolia (Desr.) Roem. & Schult. Syst. 4: 251. 1819.
Convolvulus asarifolius Desr. in Lam. Encycl. 3: 562. 1789. Hierba
del carbunclo.
Moist or wet, open places, 200 m. or less; Pete"n (La Libertad).
Mexico (Chiapas); Honduras; Panama; West Indies; South America;
tropical Asia and Africa.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 33
Plants herbaceous, trailing, not twining, glabrous, the stems very coarse and
thick, rooting at the nodes; leaves succulent, rounded-cordate or subreniform,
often 10 cm. broad, rounded at the apex, entire; peduncles short or elongate,
several- flowered, the pedicels short or elongate, rather slender; sepals unequal,
5-10 mm. long, elliptic or oval, subcoriaceous, rounded at the apex, mucronate;
corolla purple, 6-8 cm. long, glabrous; seeds blackish, glabrous.
This species is noteworthy for its irregular and interrupted dis-
tribution. It is frequent in Panama but apparently rare in other
parts of continental North America, or at least has not been collected.
Ipomoea aurantiaca L. Wms. Fieldiana, Bot. 32: 187. 1970.
Moist slopes and thickets, alt. 1,000-2,000 m., known only from
Chiapas (Breedlove 13362, type) and from Huehuetenango (Steyer-
mark 51015).
Delicate twining glabrous herbs with stems to about 2 mm. in diameter.
Leaves oblong-lanceolate, acuminate, truncate at the base, membranaceous or
submembranaceous, the blade when mature 4.5-12 cm. long and 1.5-6 cm. broad,
the petioles very slender (0.5 mm. in diameter), 1-1.5 (4) cm. long; inflorescence
axillary, about as long as or longer than the subtending leaves, few-flowered,
basically cymose, the peduncle about as thick as the stem, 2.5-7 cm. long, the
pedicels slender, somewhat furfurescent, mostly less than 2 cm. long; calyx about
8-10 mm. long, the lobes unequal, broadly ovate to suborbicular, obtuse, coria-
ceous, the outer ones somewhat furfurescent dorsally, 4-6 mm. long, the inner
lobes 8-10 mm. long, the margins chartaceous; corolla orange or yellow, tubular-
campanulate, glabrous, 5-6 cm. long and about 1 cm. in diameter near the throat,
lobes broadly ovate, about 1 cm. long; capsule unknown.
There are only two species of Ipomoea in Guatemala with large
yellow flowers — this and /. flaivda.
Ipomoea batatas (L.) Poir. Encycl. 6: 14. 1804. Convolvulus
batatas L. Sp. PI. 154. 1753. Camote; is or iz (Quiche", Maya, Quecchi) ;
Us (Ixil) ; on (Jacaltec, Chuje) ; om (Poconchi) . Sweet potato.
Cultivated commonly at low and middle elevations, and often
naturalized in waste ground or moist thickets or fields; perhaps native
of America, now cultivated in most warmer regions of the earth;
noted as escaped in Sacatepe"quez and Quezaltenango, and doubtless
to be found in various other departments.
Perennial from a large fleshy root, glabrous or nearly so; stems usually trailing,
a meter long or more; leaves very variable in outline, ovate to rounded, cordate at
the base, entire or dentate or lobate, often deeply lobate, with acute lobes; pedun-
cles equaling or shorter than the petioles, few-flowered; sepals oblong, acute or
rounded and cuspidulate, 7-10 mm. long, somewhat unequal; corolla pale purple
or almost wholly white, about 5 cm. long, glabrous; capsule 2-celled, glabrous;
seeds glabrous.
The usual name for the sweet potato in Mexico and Central
America is "camote," a word of Nahuatl origin. In Cuba this vege-
34 FIELDIANA: BOTANY, VOLUME 24
table is called "boniato," a word said to be of Antillean origin. In
Puerto Rico and in other parts of tropical America it is called
"batata," perhaps a Carib word. It may be noted that the term ba-
tata belongs properly to the sweet potato, but in Central America
and elsewhere it was transferred to the Irish potato when that was
introduced from South America, and in most parts of the earth the
Irish potato now is known by the name that belongs rightfully to the
sweet potato. The name of this plant has given its name to a pueblo
of the Department of Chiquimula, Camotan. Sweet potatoes are
grown abundantly in some parts of Guatemala and large quantities
are eaten, but they are not a highly important food, and in some re-
gions they seldom are seen. The only departments with a large
production are Alta Verapaz, Sacatepe"quez, Izabal, and Pete"n. Al-
though often of good quality, most of the local sweet potatoes are
small, compared with those marketed in the United States. They
often are sold boiled in the markets, and by the upland Indians are
eaten more or less as a delicacy. In Guatemala sweet potatoes are
especially esteemed for preparing a dessert known as dulce de camote,
which sometimes is rather good, but generally is too highly flavored
with cinnamon. There are a number of distinct color varieties of
sweet potatoes in Guatemala. A camote bianco when grated or mashed
is almost snow-white when cooked, and is much esteemed for des-
serts, likewise the camote morado, which is of a handsome deep purple
color when put on the table. The sweet potato doubtless has been
grown in Central America for many centuries. It is closely related to
I. tiliacea, and is believed by some persons to have originated from
that species. In cultivation the plants often bear no flowers, and
fertile capsules are a great rarity.
Ipomoea capillacea (HBK.) G. Don, Gen. Syst. 4: 267. 1838.
Convolvulus capillaceus HBK. Nov. Gen. & Sp. 3: 97. Feb. 1819.
Ipomoea muricata Cav. Icon. 5: 52, t. 478, fig. 2. 1799, non Jacq. 1798.
I. armata Roem. & Schult. Syst. 4: 214. Spring 1819. /. murica-
tisepala Matuda, An. Inst. Biol. Me"x. 34: 124. 1964, nom. illeg.
Open grassy places in pine forest, sometimes in wet meadows or
on limestone, 800-2,000 m.; Zacapa; Chiquimula; Guatemala;
Quiche" ; Huehuetenango. Southwestern United States; Mexico;
El Salvador; Honduras; Costa Rica; Panama. Northwestern South
America.
Perennial from a small tuberous root, the stems erect, simple or sparsely
branched, usually 40 cm. long or less, glabrous throughout or nearly so; leaves
sessile, pedately parted into filiform segments 1-2 cm. long; peduncles equaling
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 35
or shorter than the leaves; inflorescence 1-flowered; sepals 4-5 mm. long, oval or
broadly ovate, obtuse, sparsely or densely muricate dorsally; corolla bright pink
or purple, about 2.5 cm. long; capsule globose, glabrous, 4-5 mm. long; seeds
minutely puberulent.
Easy of recognition by the erect habit, small flowers, and filiform
divisions of the leaves. Called "pinito" in El Salvador.
Ipomoea carnea Jacq. Enum. PI. Carib. 13. 1760.
Rocky brushy hillsides, 200-500 m.; Zacapa. Mexico (Yucatan) ;
Nicaragua; Costa Rica; West Indies; northern South America.
A large woody vine, the branches usually stout and thick, pale, whitish-
tomentose at first; leaves broadly ovate to suborbicular, 5-15 cm. long, rounded
and apiculate at the apex or subacute, shallowly cordate at the base, pilosulous
and densely puncticulate above or glabrate, pale-tomentose beneath, petiole usu-
ally shorter than the blade; peduncles about equaling the petioles or shorter,
several-flowered; sepals suborbicular, 5 mm. long, tomentose; corolla violet-pink
or white, 4-7 cm. long, pilose outside; capsule ovoid, 1.5 cm. long, glabrous; seeds
densely covered with very long, soft, spreading hairs.
The Maya name of Yucatan is recorded as "chocobcat."
Ipomoea contrerasii L. Wms. Fieldiana, Bot. 32: 189 1970.
Thickets at 300 meters or less, Pete"n (type, Contreras 36^0) .
Twining herbaceous vines, the slender stems, the petioles and the peduncles
with long spreading hirsute pubescence, the hairs 4-5 mm. long. Leaves broadly
ovate-cordate, short acuminate, with 6-8 pairs of lateral nerves, the blades 8-12
cm. long and 7.5-10 cm. broad, glabrous, submembranaceous, cordate base very
open, the petioles slender, hirsute, 9-11 cm. long; inflorescence axillary, 1-flowered,
the peduncles 4-5 cm. long, hirsute, pedicels 3-4.5 cm. long, swollen upward (in-
flated?), glabrous, 4-5 mm. in diameter at the apex; sepals chartaceous, the mar-
gins thin and membranaceous, about equal in length, narrowly oblong-lanceolate,
acute, 2.5-3 cm. long and 0.8-1 cm. broad; corolla blue, campanulate, the limb
rotate, 10-12 cm. long in anthesis, glabrous, stamens inserted about 2.5 cm. above
the base of the corolla, the filaments bearded at the base, the anthers reaching to
about the middle of the corolla or less; capsule unknown.
This is one of the largest flowered of the Guatemalan morning
glories, easily distinguished by a combination of large flowers,
spreading hairs on the stems, petioles and peduncles, and the large
calyx.
Ipomoea costellata Torr. Bot. Mex. Bound. Surv. 149. 1859.
Brushy plains or limestone hillsides, 250-1,500 m.; Zacapa; Chi-
quimula; Huehuetenango. Southwestern United States; Mexico.
Plants annual, erect or ascending, the larger plants twining, very sparsely
pilosulous or almost glabrous; leaves small, short petiolate, cleft almost to the base
36 FIELDIANA: BOTANY, VOLUME 24
into several linear, entire or lobate segments 1.5 cm. long or shorter, strigillose;
peduncles filiform, longer than the leaves, 1-flowered; sepals subequal, 4 mm. long,
linear-oblong or lance-oblong, acute, glabrous, subcoriaceous; corolla 12 mm.
long, glabrous, the limb purple, the tube white; capsule subglobose, 6 mm. long;
seeds glabrous.
This is a diminutive plant, in general appearance unlike most
Central American Ipomoeas, but several similar species occur in
Mexico.
Ipomoea crinicalyx S. Moore, Trans. Linn. Soc. II. 4: 402,
t. 27, f. 1. 1895; O'Donell, Lilloa 29: 139, t. 9. 1959. /. seleri Millsp.
Bot. Jahrb. 36, Beibl. 80: 23. 1905 (type from Yucatan).
British Honduras (El Cayo). Southern Mexico; Brazil ; Argentina.
A small or large, herbaceous vine, glabrous or sparsely short-pilose; leaves
long-petiolate, rounded-cordate, abruptly acuminate, shallowly or deeply cordate
at the base, entire, more or less punctate and minutely papillose beneath along the
linear-oblong or lance-oblong, acute, glabrous, subcoriaceous; corolla 12 mm.
veins; peduncles shorter or longer than the petioles, inflorescence a 1 -few-flowered
cyme, the pedicels slender, usually elongate, minutely puberulent; sepals herba-
ceous, about 1.5 cm. long, acute or obtuse, puberulent and puncticulate, densely
covered with long spreading green fleshy spine- like appendages; corolla purple or
blue, glabrous, 6-9 cm. long; capsule ovoid, glabrous, 12-15 mm. long; seeds pilose
on the angles.
Ipomoea demerariana Choisy in DC. Prodr. 9: 361. 1845.
Wet thickets or forest, often in wooded swamps, 700 m. or usually
less; Izabal; Alta Verapaz. British Honduras; along the Atlantic
coast to Panama; Lesser Antilles; northern South America.
A large vine, sometimes climbing to a height of 15 meters, herbaceous or some-
what woody below, the stems as much as 2.5 cm. thick, glabrous or nearly so;
leaves long-petiolate, rounded-cordate, 8-20 cm. broad, abruptly acute or cuspi-
date-acuminate, often purplish beneath, shallowly or deeply cordate at the base,
glabrate, sometimes short-pilose beneath along the veins; peduncles 10-20 cm.
long or more, usually many-flowered; sepals broadly ovate, 15-18 mm. long, obtuse
or subacute, purple or lavender, finely pubescent or almost glabrous, subequal;
corolla 5-8 cm. long, purple, glabrous; capsule subglobose, glabrous, 1 cm. long;
seeds covered with long soft hairs.
This is an exceptionally showy and handsome morning-glory, and
attains perhaps a greater size than any other Guatemalan species.
It has been confused, rather strangely, with Stictocardia campanulata
(L.) House, and has been reported from British Honduras and
Panama under that name. Stictocardia, so far as available records
indicate, is not found north of Costa Rica and Panama.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 37
Ipomoea digitata L. Syst. ed. 10. 924. 1759.
Wet thickets, at or little above sea level; Izabal. British Hon-
duras; Honduras; Panama; West Indies; South America; Old World
tropics.
A stout vine, usually herbaceous, sometimes woody below, the stems sparsely
pubescent or glabrous; leaves slender-petiolate, somewhat succulent, palmately
lobate to or (ours) below the middle, cordate or truncate at the base, the lobes
usually 5-7, ovate or lanceolate, acute or acuminate, entire, 5-8 cm. long, glabrous
or sparsely pilose, pale beneath; peduncles usually longer than the petioles, pilosu-
lous or glabrous; inflorescence cymose, few-several-flowered; pedicels much longer
than the calyx; sepals equal, orbicular, subcoriaceous, 8-10 mm. long, glabrous or
sparsely pubescent; corolla rose-pink, 4-6 cm. long, glabrous; ovary glabrous, 2-
celled; seeds 4, long-pilose.
Known in Guatemala, Honduras, and British Honduras at sea
level and doubtless will be found along the Atlantic shore in Nicar-
agua and Costa Rica. It is rare in collections.
Ipomoea fistulosa Mart, ex Choisy, DC. Prodr. 9: 349. 1845;
O'Donell, Lilloa 29: 157, fig. 15. 1959. Batatas ? crassicaulis Benth.
Voy. Sulph. 5: 134. 1845. Ipomoea crassicaulis Rob. Proc. Am.
Acad. Sci. 51 : 530. 1916. Campanilla; mananita; campana; campanula
lila.
Usually in wet, open ground, especially in marshes about lakes or
along streams, often becoming naturalized in thickets, 1,000 m. or
less; found wild in Pete"n; Izabal; Chiquimula; Jutiapa; San Marcos;
and probably elsewhere; cultivated commonly for ornament in parks
and gardens of many parts of Guatemala. Southwestern Texas;
Mexico; British Honduras; El Salvador to Costa Rica; Panama;
South America; Old World.
An erect herb or shrub, sometimes 2.5 m. high or even more, the stems stout,
minutely puberulent or glabrous, fistulose; leaves long-petiolate, lance-cordate or
narrowly ovate-cordate, 10-25 cm. long, very narrowly long-acuminate, shallowly
cordate at the base, entire, minutely puberulent or glabrous; peduncles little longer
than the petioles, several-flowered, the pedicels short or elongate, thick; sepals
subcoriaceous, suborbicular, 5-6 mm. long, minutely puberulent or glabrate; corolla
deep pink or rose-purple, finely tomentulose, 5-8 cm. long; capsule subglobose,
2 cm. long or smaller, glabrous; seeds covered with very long, soft, spreading hairs.
Sometimes called "campanilla de arbol" and "campanola" in El
Salvador. The plant is not a particularly handsome or showy one,
but it is much planted in Guatemalan gardens, perhaps because it
thrives in spite of neglect. Its natural habitat in Central America
seems to be marshes about the borders of lakes, but it grows well,
even during the dry season, in well drained soil.
38 FIELDIANA: BOTANY, VOLUME 24
Dr. O'Donell, 1. c., believes that this species is native only in
South America but has been introduced as an ornamental plant in
North America and to other continents. It has the appearance of
a native plant in Guatemala.
Ipomoea flavida L. Wms. Fieldiana, Bot. 32: 190. 1970.
Probably moist thickets and forest edges, alt. about 1,000 m.;
known only from the type (Tuerckheim 3930)', Alta Verapaz. Ende-
mic.
Slender herbaceous vines with minutely puberulent stems to about 2 mm. in
diameter. Leaves ovate, the base shallowly cordate, apex acuminate, glabrous or
puberulent along or on the veins, principal lateral veins 5 alternate pairs, blade
3.5-11 cm. long and 2-7.5 cm. broad, petioles slender, puberulent, 2-6 cm. long;
inflorescence axillary, corymbiform, mostly 5-7-flowered, shorter or subequal to
the subtending leaves, peduncles 0.2-2 cm. long, the pedicels slender, at anthesis
about 2-2.5 cm. long; calyx about 12 mm. long, the lobes elliptic-oblong, obtuse,
coriaceous, subequal and 3-4 mm. broad, the outer dorsally puberulent or pilosu-
lose, the inner glabrous or nearly so; corolla narrowly infundibuliform, bright sul-
phur-yellow, glabrous except tip of lobes, about 4.5 cm. long and 1.5 cm. broad,
contracted to a short narrow tube at the calyx, anther reaching to about the middle
of corolla, about 5 mm. long, the style reaching middle of the corolla; capsule and
seeds unknown.
The only Ipomoea in Guatemala with large yellow flowers.
Another occurs in Mexico (Chiapas).
Ipomoea heterodoxa Standl. & Steyerm. Field Mus. Bot. 23:
82. 1944.
British Honduras, in moist thickets or open forest, little above
sea level; type from Maskall, Gentle 971. Mexico (Yucatan; Cam-
peche).
A slender twining herb, glabrous throughout; leaves long-petiolate, digitately
5-7-foliolate, the lower leaflets short-petiolulate or sessile, oblanceolate or linear-
oblanceolate, 4-8.5 cm. long, 8-18 mm. broad, slightly narrowed to the obtuse
apex, attenuate to the base, entire; terminal leaflets 6-12 cm. long, deeply 3-parted,
often almost to the base, cuneately long-attenuate at the base, the lobes entire;
inflorescence a few-several-flowered cyme exceeded by leaf petioles, the peduncle
to 1.5 cm. long, the slender pedicels less than 1 cm. long; sepals unequal, subcoria-
ceous, oval or suborbicular, the inner ones 5-6 mm. long, rounded and muticous
at the apex, the outer ones somewhat shorter; corolla white tinged with pink, or
pale pink throughout, glabrous, 3.5-4 cm. long, the limb scarcely more than 1.5 cm.
broad; capsule glabrous, ellipsoid-ovoid, 1 cm. long; seeds covered with long soft
spreading hairs.
A curious species allied to I. cairica (L.) Sweet.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 39
Ipomoea indica (Burm.) Merrill, Int. Rumph. Herb. Amb. 445.
1917. Convolvulus indicus Burm. Index Universalis Herb. Amb. 7:
[6]. 1755. Ipomoea congesta R. Br. Fl. Nov. Holl. ed. 1, 485. 1810;
O'Donell, Lilloa 29: 134. 1959. Convolvulus acuminatus Vahl, Symb.
Bot. 3: 26. 1794. Ipomoea cathartica Poir. in Lam. Encycl. Me'th.
Suppl. 4: 633. 1816. /. acuminata Roem. & Schult. Syst. Veg. 4: 228.
1819, non /. acuminata Ruiz & Pavon, 1799. I. vahliana House, Ann.
N. Y. Acad. Sci. 18: 204. 1908. Quilamul; sayun (Coban, Quecchi).
Moist or wet thickets, often a weed in second growth or in aban-
doned fields or hedge rows, 1,400 m. or less; Pete"n; Alta Verapaz;
Zacapa; Sacatepe'quez ; Suchitepe"quez; Retalhuleu; Huehuetanango ;
San Marcos; doubtless in other departments. British Honduras.
Pantropical, the type from southeast Asia, and widely cultivated.
In America from the southern United States to Argentina.
Plants herbaceous, twining, much branched and often elongated, rarely pros-
trate, the stems appressed-pilosulose to glabrate; leaves rounded-ovate, mostly
5-9 cm. long, usually acuminate, cordate at the base, entire or rarely trilobate,
appressed-pilosulous or glabrate; peduncles 2-20 cm. long, mostly shorter than the
petioles but sometimes longer; inflorescences 1-few-flowered cymes; sepals pale
green, herbaceous at least toward the apex, lance-ovate or ovate, 1-2 cm. long,
3-7 mm. broad, acuminate or long-acuminate, appressed-pilosulous or usually gla-
brate; corolla blue or purple, rarely white, glabrous outside, 5-7 cm. long or even
larger, the limb 6-8 cm. broad; capsule globose, 1 cm. broad or less, glabrous; seeds
glabrous, 3 mm. long.
Called "gloria de la manana" in British Honduras, a translation
of the English name that has come into use along the banana coast
of Central America, "quiebra-platos" (Veracruz). The species is a
variable one, especially in pubescence. House divided the material
that we refer here into three species, but they cannot be separated
definitely by the characters that he suggested.
Ipomoea indica var. variabilis (Schlecht. & Cham.) L. Wms.
Fieldiana, Bot. 32: 191. 1970. Convolvulus variabilis Schlecht. &
Cham. Linnaea 5: 116. 1830. Ipomoea variabilis Choisy in DC.
Prodr. 9: 383. 1845.
Moist or wet thickets, 700 m. or less; Izabal; Escuintla; San
Marcos. Southern Mexico.
A slender herbaceous vine, the stems and leaves glabrous or nearly so; leaves
thin, long-petiolate, cordate-ovate or triangular-ovate, 5-8 cm. long, acuminate
or long-acuminate, rather shallowly cordate at the base, sometimes subhastate,
entire; peduncles shorter than the petioles, 1-2 cm. long, 1-5-flowered, the bracts
green, linear-lanceolate; sepals lanceolate or ovate-lanceolate, herbaceous, 12-15
40 FIELDIANA: BOTANY, VOLUME 24
mm. long, long-setose or subsetose at the base; corolla bright purple, 5-6 cm. long,
glabrous, with a very broad limb; capsule 8-10 mm. in diameter; seeds glabrous.
Distinguished from the species by the long setose or subsetose
hairs at the base of the calyx.
Ipomoea leucotricha Donn.-gm. Bot. Gaz. 23: 10. 1897.
Alta Verapaz; type collected near Nenton, Heuhuetenango, 900-
1,200 m., Nelson 3512. Nicaragua; Costa Rica.
Plants twining, densely silvery-sericeous throughout except less so on the co-
rollas and the upper leaf surface; leaves long-petiolate, orbicular-cordate, 7-12 cm.
long, 6-10 cm. broad, entire or rarely lobate, sparsely pilose and green on the upper
surface; peduncles longer than the leaves, with numerous flowers; bracts linear-
lanceolate, 6 mm. long, the pedicels 1-2 cm. long; sepals 6 mm. long, subequal,
oblong-ovate, with a recurved mucro at the apex; corolla purple, 5-7 cm. long and
2.5 cm. broad at the throat, sericeous outside except in the plicae; ovary glabrous.
Ipomoea lindenii Mart. & Gal. Bull. Acad. Brux. 12, 2: 264.
1845; Matuda, An. Inst. Biol. 36: 84. 1965.
Thickets. Guatemala, Escuintla, Standley 24406, according to
Matuda. Mexico.
Twining plants with slender often suffruticose stems. Leaves cordate-ovate
to subtriangular, acuminate, glabrous to sparsely pubescent along the veins, 5-
10 cm. long and 2-6 cm. broad, petioles slender, 3-5 cm. long; inflorescence axillary,
1-2-flowered, the peduncles 1-5 mm. long, the pedicels to about 15 mm. long, gla-
brous to sparsely pilose; sepals unequal, oblong or lance-oblong, obtuse, 6-8 mm.
long, glabrous or nearly so; corolla infundibuliform to campanulate, glabrous ex-
cept toward the tip, 5-6 cm. long.
Prof. Matuda, possibly in error, reports the species from Guate-
mala based on Standley 24406 in 1922.
Ipomoea mairetii Choisy in DC. Prodr. 9: 374. 1845. Quiebra-
cajete.
Moist or rather dry thickets or forest, often in pine-oak forest,
650-1,900 m.; Zacapa; Chiquimula; Jalapa; Jutiapa; Santa Rosa;
Guatemala; Solola; Sacatepe"quez; Chimaltenango. Southern Mexico.
A woody vine, often large and climbing over trees, the stems densely sericeous
or somewhat tomentose; leaves long-petiolate, rounded-ovate or subreniform, 7-
15 cm. long or larger, acute or acuminate, shallowly or deeply cordate at the base,
often trilobate, usually green above and rather densely appressed-pilose, very
densely silvery-sericeous beneath; peduncles several-flowered, equaling or shorter
than the leaves, the inflorescence a few-flowered cyme, open or dense, the bracts
large, ovate, acute or acuminate, persistent, the pedicels 1-3 cm. long; sepals ovate
or broadly ovate, 2-2.5 cm. long, acute, densely sericeous; corolla bright rose-
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 41
purple, 7-8 cm. long, the tube densely white-pilose outside; capsule glabrous, glo-
bose, 1 cm. in diameter; seeds blackish, densely and finely pubescent.
One of the handsomest of the local Ipomoea species, abundant at
many places in the central region at middle elevations, often growing
in thin oak forest. It has been reported from Guatemala as 7.
tyrianthina Lindl.
Ipomoea meyeri (Spreng.) G. Don, Gen. Syst. 4: 275. 1838.
Convolvulus meyeri Spreng. Syst. 1: 597. 1825. /. chiapensis Brand-
egee in Univ. Calif. Publ. Bot. 6: 60. 1914 (type from Tonala,
Chiapas). /. brachypoda Benth. Bot. Voy. Sulph. 135. 1844.
Moist thickets, 150-500 m.; Alta Verapaz; El Progreso; Zacapa;
Chiquimula; Santa Rosa. Southern Mexico; British Honduras; Hon-
duras; Costa Rica; Panama; West Indies; northern South America.
A small, slender herbaceous vine, the stems glabrous or sparsely pilose; leaves
slender-petiolate or the upper ones short-petiolate, broadly or narrowly ovate, 4-
10 cm. long, acute or acuminate, cordate at the base, entire or hastately trilobate,
glabrate, often white-puncticulate beneath; inflorescence a few-several-flowered
cyme as long as or usually shorter than the subtending leaf, peduncles to 6 cm.
long (usually much shorter in Guatemalan material), the pedicels short, bracts
linear; sepals 1-2 cm. long, linear-lanceolate, acuminate, pilose or hirsute with long
pale spreading hairs with tuberculate bases, green, herbaceous; corolla blue or
purple with a white throat, 2.5-3 cm. long, glabrous; capsule globose, 8 mm. in
diameter; seeds densely and finely pubescent.
Called "camotillo" in British Honduras; the Maya names
"tsusuc" and "xhail" are reported from Yucatan. In Honduras
called "campanilla."
Ipomoea microsepala Benth. Bot. Voy. Sulph. 136. 1844. Prin-
cipe amarillo.
Wet to dry, sometimes rocky thickets, 1,600 m. or less; Solola;
Sacatepe'quez ; Suchitepe"quez; Retalhuleu; San Marcos. Western
and southern Mexico.
A small, slender herbaceous vine, climbing over shrubs, the stems sparsely hir-
sute with long spreading hairs or glabrous; lower leaves long-petiolate, the upper
on very short petioles, broadly ovate, mostly 3-6 cm. long, acuminate to obtuse,
deeply cordate at the base, entire, glabrous or nearly so; inflorescence a few-flow-
ered axillary cyme on usually long almost filiform peduncles exceeding the leaves,
the pedicels subfiliform, mostly less than 1 cm. long; sepals subequal, oblong, ob-
tuse, sphacelate, about 2 mm. long and 1 mm. broad; corolla bright yellow, 2-2.5
cm. long; capsule subglobose; seeds densely short-pilose.
A common plant of the western Pacific plains, easily recognized
by the small yellow flowers and elongate twining stems and very
small, sphacelate sepals.
42 FIELDIANA: BOTANY, VOLUME 24
Ipomoea microsticta Hallier f. Bull. Herb. Boiss. ser. 1, 7: 411.
1899; L. Wms. Fieldiana, Bot. 32: 191. 1970. I. glabriuscula House,
Bot. Gaz. 43:409. 1907.
Moist thickets or forest edges, 325-1,200 m.; Escuintla (type
Seler & Seler 2^27); Suchitepe"quez; Sacatepe"quez ; Santa Rosa;
Chiquimula; Jutiapa. Mexico; British Honduras; Costa Rica.
Herbaceous or suffrutescent vines, the branches becoming quite woody with
age, glabrous, terete. Leaves broadly ovate to suborbicular, shallowly cordate,
acuminate, obscurely puberulent below or glabrous, provided with abundant micro-
scopic "glands" below, the blades 3-10 cm. long and 2.5-11 cm. broad, the petioles
2.5-6 cm. long, slender, usually with two inconspicuous glands at the apex; inflores-
cence a 1 -several-flowered axillary cyme, the peduncles 2-8 cm. long, the pedicels
about 1 cm. long, angulate, subclavate; calyx subglobose, glabrous, the sepals
broadly elliptic to suborbicular, subcoriaceous, cochleate to somewhat concave,
6-8 mm. long; corolla campanulate, glabrous, pale pink to white, 4-5 cm. long;
stamens inserted near the base of the corolla and about a third as long; style ex-
ceeding the stamens; capsules broadly ovate, somewhat attenuate to the persistent
base of the style, 10-15 mm. long.
Said by Standley to be one of the most attractive morning-glories
of Guatemala. The campanulate corolla, subglobose calyx, and
shallowly cordate leaves are distinctive marks of this species.
The small "glands," described by Hallier, from the undersurface
of the leaves are on all the specimens we have seen. However, we
are not sure that these are not caused by a fungus.
Ipomoea minutiflora (Mart. & Gal.) House, Muhlenbergia 5:
71. 1909. Convolulus minutiflorus Mart. & Gal. Bull. Acad. Brux. 12,
pt. 2: 262. 1845. /. filipes Benth. ex Meissn. in Mart. Fl. Bras. 7:
274. 1869.
Moist shaded banks, brushy rocky slopes, wet meadows, some-
times a weed in cultivated ground, 1,200 m. or less; Zacapa; Chiqui-
mula; Jutiapa; Santa Rosa; Escuintla. Southern Mexico; British
Honduras to Nicaragua and Costa Rica; northern South America.
Plants annual, usually prostrate, very slender, the stems pilose or glabrate,
often densely leafy; leaf blades broadly ovate to subreniform, 1-3 (-6. 5) cm. long,
acute to rounded at the apex, shallowly or deeply cordate at the base, entire or
somewhat angulate, ciliate, sparsely pilosulous above, glabrous beneath, long-petio-
late to almost sessile; peduncles filiform, 1-3-flowered, usually longer than the
leaves, the pedicels recurved'in fruit; sepals 2 mm. long, acute, pilose; corolla yel-
low, 1-1.5 cm. long; capsule subglobose, glabrous, 4-5 mm. long; seeds blackish,
minutely puberulent or glabrate.
An inconspicuous little plant, usually creeping, noteworthy for
the very small, bright yellow corollas. It is to be expected in other
departments.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 43
Ipomoea murucoides Roem. & Schult. Syst. 4: 248. 1819. Siete
camisas; tonche; tutumuzco; tutumuste; siete pellejos; tutumuscuavo;
palo bianco; siete capas; tutusmuscuago; tutumuzcual.
Brushy, open, dry, often rocky slopes or plains or in open dry
forest, frequently in hedges, or in oak forest, 600-2,000 m.; Chiqui-
mula; Jalapa; Jutiapa; Santa Rosa; Guatemala; Sacatepe*quez ;
Quiche" ; Huehuetenango. Southern Mexico.
A tree, commonly 5-9 m. high, the crown low and spreading, the bark almost
smooth, white, the branches thick, the young branches densely white-tomentose;
leaves slender- petiolate, oblong-lanceolate, mostly 7-12 cm. long, long-acuminate,
obtuse or rounded at the base, entire, somewhat tomentose when young but soon
glabrate; inflorescence a white tomentose few-several-flowered terminal raceme,
or single flowers borne in the axils of leaves, the peduncles as short as the petioles
or almost none, the pedicels 2-4 cm. long; sepals 2-3 cm. long, subequal, coriaceous,
rounded at the apex, tomentose outside, sericeous within; corolla pure white, 7-
8 cm. long, tomentose outside; capsule 2.5 cm. long, ovoid, glabrous; margins of
the seeds densely set with long soft spreading white hairs.
It is said that the ashes make the best lye for soap-making. Cattle
are fond of the ashes, which they eat in place of salt. This species
belongs to the group of tree Ipomoeas that grow so profusely in the
dryer parts of western Mexico. It is common and conspicuous in
the central region, especially about Antigua, where it is often planted
for living fence posts or for hedges. It is more abundant in the west,
and is conspicuous on the arid hills about Sacapulas in Quiche*. The
trees are conspicuous because of their white trunks and branches,
which look as if they had been used as roosting places by birds. The
abundant large white flowers are very handsome, and the trees are
strangely unlike most members of the morning-glory family. The
flowers apparently remain open all day long and, once open, probably
do not close until they wither.
Ipomoea nil (L.) Roth, Cat. 1: 36. 1797; O'Donell, Lilloa 29:
197, L 1959. Convolvulus nil L. Sp. PL ed. 2. 219. 1762. Campanula;
campana.
Wet to dry, often rocky thickets, often in waste or cultivated
ground, 1,750 m. or less; Zacapa; Chiquimula; Jutiapa; Santa Rosa;
Sacatepe"quez; Guatemala; Quiche"; Huehuetenango. Florida; Mexi-
co; British Honduras to Panama; West Indies; South America; Old
World tropics.
A small or rather large, herbaceous vine, usually always annual, hirsute almost
throughout with sparse or often dense, usually pustulate-based, unicellular, brown-
ish hairs; leaves 5-15 cm. long, shallowly or deeply cordate at the base, ovate to
44
FIELDIANA: BOTANY, VOLUME 24
FIG. 5. Ipomoea nil. Habit of the plant, X
suborbicular, entire or trilobate (rarely 5-lobate), the lobes entire, abruptly acute
or acuminate; inflorescence 2-5-flowered, cymose, dense and head-like, peduncles
longer or shorter than the petioles; sepals subequal, 1.5-2.5 cm. long, linear and
herbaceous, the bases somewhat broadened, densely long-hirsute; corolla infundi-
buliform, blue or purple, 3-5 cm. long, glabrous, the limb 4-5 cm. broad; ovary
3-celled; style 17-30 mm. long, stigma biglobular; capsule globose, 8-12 mm. long;
seeds finely and densely pubescent.
One of the most common weedy species of Ipomoea through much
of the lowlands of Central America. The Maya name "itzotz-cabil"
is reported from Yucatan.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 45
Ipomoea ophiodes Standl. & Steyerm. Field Mus. Bot. 23: 82.
1944.
Moist thickets or forest, 200-400 m.; Santa Rosa (type from
region of La Morenita, northeast of Chiquimulilla, Standley 78884).
El Salvador.
A small herbaceous vine, the stems sometimes prostrate and rooting at the
nodes, densely hirsute with very long, spreading hairs, the internodes elongate;
leaves long-petiolate, cordate-ovate, 4-10 cm. long, 3-6.5 cm. broad, acute or acu-
minate, hirsutulous above, slightly paler beneath and puberulent, the petioles
puberulent and very sparsely hirsute; inflorescence 1-flowered, peduncles scarcely
1 cm. long, the slender pedicels 2.5-3.5 cm. long, minutely puberulent and very
sparsely hirsute; sepals subequal, subherbaceous, 12 mm. long, linear-lanceolate,
acuminate, sparsely long-hirsute; corolla deep purple, 8-9 cm. long, glabrous, the
limb 8 cm. broad.
Noteworthy for the very dense and long hairs of the stem, the
pubescence of the leaves consisting wholly of short and inconspicuous
hairs.
Ipomoea parasitica (HBK.) G. Don, Gen. Syst. 4: 275. 1838.
Convolvulus parasiticus HBK. Nov. Gen. & Sp. PI. 3: 103. 1819.
Campanilla; quebra-cajete.
Brushy slopes and thickets, 800-1,500 m.; Jalapa; Jutiapa; Santa
Rosa. Mexico through Central America. Northern South America.
A small or large, herbaceous vine, the stems glabrous or nearly so, often bear-
ing below wart-like fleshy tubercles; leaves long-petiolate, rounded-cordate, 6-10
cm. long, acuminate or rounded and cuspidate at the apex, strigillose above, gla-
brous beneath or nearly so, the petioles rarely tuberculate; inflorescence 5-12-
flowered, peduncles shorter or slightly longer than the petioles, stout, the pedicels
much thickened upward; sepals 4-5 mm. long, the outer ones lanceolate-ovate,
subacute and minutely cuspidate, minutely appressed-pilosulous, the inner ones
rounded; corolla sericeous outside, 3 cm. long, bright purple; stamens reaching to
about middle of the corolla; style as long as the stamens but breaking off shortly
above the capsule after corolla falls; capsule ovoid-globose, 1 cm. long; seeds mi-
nutely puberulent.
This species is very closely allied to /. tricolor Cav. but can be
distinguished by the smaller corollas which are sericeous outside,
and by the broader calyx lobes.
Ipomoea pauciflora Mart. & Gal. Bull. Acad. Brux. 122: 266.
1845; Walp. Rep. 6: 532. 1847; L. Wms. Fieldiana, Bot. 32: 192. 1970.
Ipomoea populina House, Ann. N. Y. Acad. Sci. 18: 227. 1908.
Dry thickets and hills, 400-900 m.; Chimaltenango; Chiquimula;
Progreso. Western and southern Mexico; Honduras; Nicaragua.
46 FIELDIANA: BOTANY, VOLUME 24
A large vine, woody throughout, the stems sometimes 2 cm. thick, glabrous or
nearly so; leaves mostly ovate, entire, 5-8 cm. long, acuminate or long- acuminate,
rounded or very shallowly cordate at the base, glabrous or nearly so; inflorescence
cymosely few-many-flowered, often on defoliate branches, peduncles 8-12 cm. long,
the pedicels 3 cm. long or shorter; sepals subequal, rounded-ovate, 5-8 mm. long,
glabrous; corolla white, often with a dark red or purple throat, about 6 cm.
long, glabrous, the limb 6-8 cm. broad; capsule thick-walled, ellipsoid, 2 cm. long,
1 cm. thick, 4-valvate, 2-celled; seeds covered with long soft spreading white hairs.
A most attractive woody vine that is in flower in dry regions from
about the end of the rainy season. It is exceedingly common in parts
of central Honduras.
Ipomoea perplexa L. Wms. Fieldiana, Bot. 32: 193. 1970.
In thickets, clearings, or forest edges, alt. up to 600 m.; Pete"n.
British Honduras (type, Bartlett 12868) and Honduras.
Slender twining herbaceous vines, stems about 1 mm. or less in diameter, gla-
brous. Leaves glabrous, small, narrowly ovate to broadly ovate in outline, cordate,
acute or acuminate, blade entire, dentate to trilobulate, 1.5-6 cm. long and 0.6-
4.5 cm. broad, petioles short, to about as long as the blade; inflorescence an axillary
few-several-flowered corymb, peduncles about as thick as the stem, 1.5-6 cm. long,
pedicels relatively stout, less than 1 cm. long; calyx coriaceous, glabrous, the lobes
subequal, about 4 mm. long, the outer ones usually somewhat shorter, oblong to
oblong-obovate or obovate, obtuse or truncate, the margins somewhat chartaceous,
usually with a minute inframarginal apicule at the center; corolla small, pink or
purple, narrowly funnelform, glabrous outside, 14-18 mm. long, stamens inserted
near the base of the corolla, unequal, reaching to about the middle, anthers 1-1.2
mm. long, filaments bearded at the base, stigma globose (or biglobose), large, about
0.8 mm. broad; capsule glabrous, subglobose, about 4-5 mm. long; seeds brown or
black, glabrous, smooth.
The smallest flowered of Central American Ipomoeas with pink or
purplish flowers. It is distinguished by the small flowers and the
small obtuse or truncate calyx lobes. The leaves are usually lobate
but lobate and entire leaves occur on the same stem.
Ipomoea pes-caprae (L.) Sweet, Hort. Lond. 35. 1818. Con-
volvulus pes-caprae L. Sp. PI. 159. 1753. See Figure 12.
On sea beaches; Izabal; Escuintla; Retalhuleu; San Marcos;
doubtless in all coastal departments. Common on tropical seashores
of both hemispheres.
Plants herbaceous, glabrous, somewhat succulent, the stems prostrate, thick,
sometimes 20 m. long or more, branched; leaves long-petiolate, suborbicular, 6-
10 cm. long, broadly rounded and emarginate at the apex, rounded or subcordate
at the base, entire; inflorescence 1 -several-flowered, peduncles stout, about equal-
ing the petioles, usually bifid, the pedicels long, slender; sepals oval or suborbicular,
1 cm. long, obtuse or rounded at the apex; corolla pink or purple, glabrous, 4-5 cm.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 47
long, the limb 5-8 cm. broad; capsule ovoid-globose, 1.5 cm. long; seeds densely
pubescent.
One of the most characteristic and widely distributed of tropical
strand plants, and found on most of the sandy seashores of Central
America. The stems are coarse, rope-like, and strong, so that they
can stand the wind and waves that prevail in such places. Along the
Pacific coast of Guatemala the leaves and stems wither during the
dry season.
American material of this species is said by van Ooststroom, and
O'Donell seems to agree (Lilloa 30: 46. 1960), to be a subspecies dis-
tinct from the typical subspecies which appears to be limited to east
Africa, tropical Asia, and the Malay Archipeligo. An acceptable
subspecific name for the American plant is subsp. brasiliensis (L.)
Oosts. If a varietal rank is desired the proper name would seem to
be var. emarginata Hallier f .
Ipomoea praecana House, Ann. N. Y. Acad. Sci. 18: 227. 1908.
Dry thickets or forest, 300-700 m. ; Zacapa; El Progreso. Southern
Mexico; Honduras; Nicaragua.
A stout vine, woody throughout, the stems thick, densely whitish-tomentose;
leaves on rather short petioles, suborbicular to broadly ovate, 7-20 cm. long,
rounded to acute at the apex, shallowly cordate at the base, densely whitish-to-
mentose on both surfaces, thick; inflorescence 3-5-flowered, peduncles equaling or
shorter than the petioles, the pedicels 5-10 mm. long; sepals 18-25 mm. long, ellip-
tic or oval, obtuse, densely white-tomentose outside, almost glabrous within; co-
rolla white, 6-13 cm. long, sparsely or densely white-pilose outside, the limb 6-10
cm. broad.
Ipomoea pulchella Roth, Nov. PL Sp. 115. 1821; O'Donell,
Lilloa 29: 221. 1959.
Moist thickets or open grassy places near lake shores, 500-600 m. ;
Jutiapa. Circumtropical, in America from southern United States
to Argentina.
Plants probably annual, very slender, twining or trailing, herbaceous, glabrous;
leaves long-petiolate, pedately 5-foliolate, the leaflets lanceolate or linear-lanceo-
late, sessile, 1-4 cm. long, acute, entire; peduncles filiform, equaling or often much
longer than the petioles, spirally coiled, 1-flowered; sepals greenish, ovate or oval,
5-7 mm. long, obtuse or rounded at the apex; capsule globose, 8-10 mm. long,
glabrous; seeds finely pubescent.
This species may be recognized at once by the delicate, conspic-
uously coiled peduncles, a character not found in any other Ipomoea
of Central America.
48 FIELDIANA: BOTANY, VOLUME 24
Ipomoea purga (Wender.) Hayne, Arzn. Gew. 12: t. 33, 3^.
1833; Convolvulus purga Wender. Pharm. Centralbl. 1: 457. 1830.
Quilamul; quiebra-cajete; nacta (Quecchi).
Wet to dry thickets, often in waste ground or in hedges, 2,000 m.
or lower, most frequent at low elevations; Alta Verapaz; Izabal;
Santa Rosa; Escuintla; Guatemala; Suchitep£quez; Retaljuleu; San
Marcos. Southern Mexico; El Salvador and Honduras to Costa Rica.
Usually a large, herbaceous vine, glabrous throughout or nearly so; lower leaves
long-petiolate, the upper ones almost sessile sometimes and clasping the stems; leaf
blades ovate or broadly ovate, thin, mostly 5-11 cm. long, acute or acuminate,
deeply cordate at the base, entire; peduncles usually longer than the subtending
petioles, 1-2-flowered ; sepals very unequal, the longest 8-9 mm. long, oval, very
obtuse and usually cuspidulate, glabrous; corolla red-purple or rose-purple, gla-
brous, salverform, 7-8 cm. long, the tube slender, the limb rotate or nearly so.
Known in El Salvador as "campanilla," "campanula," and
"mechoacan." This plant has long been known in pharmacy as the
source of the drug jalap or jalapa, obtained from the roots, and
formerly used extensively in medicine as a purgative. Large amounts
of the crude drug have been exported from Mexico, and some
amounts may still be exported to the United States. The purgative
properties are believed to exist in a resin that may be separated from
the large tuberous roots, which are merely dried for export. The
plant was used in medicine by the ancient Mexicans, and perhaps by
the people living farther southward. It is one of the handsomest of
the Ipomoeas because of its brilliantly colored, red-purple flowers.
A century and a half ago there was much confusion regarding the
source of the drug jalap, and for a while it was believed that it was
derived from Mirabilis jalapa, hence the erroneous drug name given
to that plant.
Ipomoea purpurea (L.) Roth, Bot. Abh. 27. 1787. Convolvulus
purpurea L. Sp. PI. ed. 2, 219. 1762. Pharbitis diversifolius Lindl.
Bot. Reg. 23: t. 1988. 1837. Ipomoea purpurea var. diversifolia
O'Donell, Lilloa 26: 385, fig. 1953. Campanilla; quiebra-cajete;
quilamul
Moist thickets or hedges, often in second growth or waste ground,
frequently a weed in cornfields, sometimes in oak forest, 1,300-2,100
m.; Alta Verapaz; Baja Verapaz; Jalapa; Escuintla; Guatemala;
Sacatepe"quez; Chimaltenango; Huehuetenango. United States;
Mexico; El Salvador to Costa Rica; West Indies; South America;
Old World tropics, where introduced.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 49
A slender herbaceous annual vine, the stems pilose or hirsute with spreading
or retrorse hairs; leaves broadly ovate or rounded, shallowly or deeply cordate at
the base, entire or often trilobate, long-petiolate, hirsute or hirsutulous on both
surfaces, mostly 2-10 cm. long and about as broad; cymes 1-5-flowered, peduncles
usually longer than the petioles, the pedicels short or elongate, sepals herbaceous,
lanceolate or oblong-lanceolate, 8-16 mm. long, acute, pilose or hirsute (especially
at the base); corolla glabrous, blue, purple, pink, or white, sometimes with vertical
stripes of different colors, 3-5 cm. long; ovary usually 3-celled; capsule depressed-
globose, 10 mm. long; seeds black, glabrous.
In Central America this morning-glory does not seem to become
so noxious a weed as it often does in the United States where it some-
times forms such dense tangles in cornfields that it is all but impos-
sible to force one's way through them, and in late summer it often is
necessary to cut and remove the vines before a potato crop can be
harvested. Like all or most Ipomoeas, this is a handsome plant, and
often is cultivated for ornament. In Guatemala they often are found
in cultivation and sometimes as escapees from cultivation forms with
"double" corollas, variously colored. Forms with cordate entire or
trilobate leaves are common in Guatemala and sometimes both leaf
forms are found on the same specimen.
Ipomoea reptans (L.) Poir. in Lam. Encycl. Suppl. 3: 460. 1813.
Convolvulus reptans L. Sp. PL 158. 1753. I. aquatica Forsk. Fl.
Aegypt. Arab. 44. 1775.
In wet soil or floating in water, at or little above sea level ; British
Honduras; Costa Rica; Panama; West Indies; South America; Old
World tropics.
Plants herbaceous, the stems hollow, rooting at the nodes, trailing on wet soil
or floating on water, glabrous or nearly so; leaves mostly hastate, with narrowly
triangular or lanceolate, terminal lobes and small spreading basal lobes, acute or
obtuse, shallowly cordate at the base, 4-12 cm. long, long-petiolate; peduncles
short, 1-few-flowered, the pedicels slender and greatly elongate; sepals subequal,
ovate-oblong, obtuse or subacute, mucronulate, glabrous, 7 mm. long; corolla gla-
brous, purple, 4-5 cm. long, with a broad limb; capsule small, globose, with 4 or
fewer seeds, these densely soft-pilose.
A widespread species but uncommon in our region. To be ex-
pected in Izabal and in Mexico.
Ipomoea sagittata Lam. 111. 1: 466. 1791.
In swamps, 300 m.; Huehuetenango (Cienaga de Lagartero,
Steyermark 51539}. British Honduras (Pueblo Nuevo, New River).
Southeastern United States; Mexico; West Indies.
50 FIELDIANA: BOTANY, VOLUME 24
Usually a small, herbaceous vine, the stems very slender, glabrous throughout
or nearly so; leaves slender-petiolate, 4-7 cm. long, very deeply hastate at the base,
entire, the terminal lobe almost linear, the basal lobes linear or lanceolate, often
almost as long as the terminal one; inflorescence 1- flowered, peduncles stout,
shorter than the petioles, the pedicels elongate, thick and dilated above; sepals
oval, 8-9 mm. long, glabrous, subcoriaceous, obtuse or rounded at the apex and
mucronulate; corolla purple, glabrous, about 6-7 cm. long, the limb 6 cm. broad
or larger.
Known in Guatemala from a single collection. The species
reaches its southern limit here, in continental North America.
Ipomoea santae-rosae Standl. & Steyerm. Field Mus. Bot. 23:
81. 1944. Campanula; campana.
Moist or wet thickets or dry, brushy slopes, 300-1,400 m.; Baja
Verapaz; Guatemala; Zacapa; Santa Rosa (type from Chiquimulilla,
Standley 79211); Quezaltenango; Huehuetenango. El Salvador;
Honduras; Nicaragua. Mexico (Chiapas).
A slender woody vine, glabrous throughout or nearly so; leaves long petiolate,
broadly ovate-cordate, 4-10 cm. long, 3.5-6.5 cm. broad, long- acuminate, deeply
cordate at the base, hirtellous beneath at the base of the blade, otherwise glabrous;
inflorescence a 1-3 flowered cyme, the peduncles mostly about 1 cm. long, the ped-
icels slender and thickened upward; sepals subequal, subcoriaceous, 18-30 mm.
long, oblong to oval, obtuse and short-aristate at the apex, somewhat thickened
and venose in fruit; corolla campanulate, red-purple or rose-pink, 6.5-9 cm. long,
glabrous, the tube 1.5 cm. or broader at the mouth; capsule 4-celled, subglobose, gla-
brous, 8-12 mm. long; seeds 4, fuscous, minutely puberulent.
The large sepals make this one of the most distinctive species of
Ipomoea in Central America.
Ipomoea saxorum Standl. & Steyerm. Field Mus. Bot. 23: 81.
1941. Quiebra cajete.
In thickets or woods, 150-1,000 m., endemic; Chiquimula (type
Steyermark 3025 4-) ; Huehuetenango ; Santa Rosa.
A herbaceous vine, the stems rather stout, densely reflexed or spreading pilose;
leaves rounded-ovate, 5-8 cm. long, 5-7 cm. broad, obtuse or acute, truncate or
subcordate at the base, densely hirtellous or short-pilose on both surfaces, petioles
shorter than the blades, 1-3.5 cm. long; inflorescence 1 -few-flowered cymes, pe-
duncles stout, 4-10 cm. long, densely spreading pilose, the stout pedicels 1.5-2.5
cm. long, pilose; sepals unequal, coriaceous, rounded at the apex, the outer ones
oval, 5 mm. long, sparsely hirtellous near the apex, the inner ones glabrous,
broader, 8 mm. long; corolla white, glabrous, 6-7 cm. long, the tube 1 cm. broad
in the throat; capsule subglobose, glabrous, 1 cm. long, broadly rounded at the
apex and abruptly contracted into the style; the style thickened at the base and
with a thickened subrotate point of dehiscence about 2 mm. above the capsule;
seeds glabrous.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 51
Ipomoea seducta House, Ann. N. Y. Acad. Sci. 18: 241. 1908.
Damp thickets and forest edges, 250-1,900 m.; Alta Verapaz
(type, Tuerckheim 7926); Huehuetenango. Mexico (Chiapas).
Twining herbaceous vines with slender glabrous stems. Leaves suborbicular-
ovate, acuminate, cordate, 5-10 cm. long and 3-7 cm. broad, glabrous, the petioles
1-1.5 cm. long, borne with the peduncle on a "short-shoot" about 1 mm. long, the
petioles of sterile leaves often much longer; inflorescence two or three-flowered with
only one flower developing, peduncles mostly less than 2 cm. long, the pedicels less
than 1 cm. long; sepals unequal, all apiculate, the outer ones broadly lanceolate,
4-6 mm. long, the inner ones 8-10 mm. long; corolla subcampanulate or funnel-
form, 5-7.5 cm. long, expanded from the base and about 1 cm. broad at the throat,
stamens reaching to the throat but not exserted, limb 4-5 cm. broad, style elon-
gated and filiform.
This species is related to /. purga but is distinguished by the sub-
campanulate corolla and the leaves and associated peduncle being
borne on a "short-shoot," which is the fused bases of the petiole and
peduncle. The ecology of the two species seems to be quite different
in Guatemala.
The shortening of petioles at fertile nodes is of interest and occurs
in several other species of related morning-glories, including /. purga.
Ipomoea sepacuitensis Donn.-Sm. Bot. Gaz. 56: 59. 1913.
Ixcajen (Quecchi).
Probably to be found in thickets and in edges of clearings; Alta
Verapaz (type from Finca Sepacuite, Cook & Griggs 590). British
Honduras.
Twining herb with herbaceous or suffrutescent stems, these with coarse setose
pubescence or glabrous with age. Leaves trilobate, the lobes to near the middle
or less, glabrous, with 7-9 nerves from near the base, the base cordate, the petioles
glabrous or sparsely setose, to about 10 cm. long, the blade 10-17 cm. long and
about as broad; peduncles coarse, setose or not, to about 15 cm. long; inflorescence
cymose, few-flowered; flowers white with rose within; calyx fleshy, glabrous, the
lobes very unequal in size, the outer ones broadly ovate and obtuse, 10-20 mm.
long, the inner one similar and 15-35 mm. long; corolla campanulate, about 5 cm.
long or perhaps more; capsule coriaceous, glabrous, to 25 mm. long or perhaps
larger at maturity.
Known from only four collections, two of these distributed as
different species of Calonyction.
Ipomoea sessilis L. Wms. Fieldiana, Bot. 32: 195. 1970.
Probably to be found in thickets as are most Ipomoeas, known
only from the type, between Nenton and Miramar, Huehuetenango,
Steyermark 51566. Endemic.
52 FIELDIANA: BOTANY, VOLUME 24
Delicate twining herbs with glabrous stems to 1 mm. in diameter. Leaves
broadly ovate-cordate, shortly acuminate, sparsely setose pubescent on both sur-
faces and ciliate, the blades 2-4 cm. long and about as broad, the petioles slender,
sparsely setose, shorter than the blades, 1.5-3 cm. long; inflorescences of a single
flower borne in the axiles of leaves, peduncle none or very short, the pedicel fleshy,
obscurely aculeolate and ridged (at least in dry material), up to about 0.8 cm. long;
sepals chartaceous to somewhat carnose, verrucose ridged at the base, narrowly
lanceolate, acuminate and subaristate, 1.5-2 cm. long, the outer two slightly longer
than the inner three; corolla rose-pink with white at base and in middle of lobes,
narrowly campanulate, about 5-6 cm. long, glabrous outside; capsule unknown.
The species is easily distinguished from the other species known
in Guatemala by the combination of long sepals which are relatively
narrow and the sessile one-flowered inflorescences.
Ipomoea setifera Poir. in Lam. Encycl. Meth. 6: 17. 1804;
O'Donell, Lilloa 39: 237, fig. 1959. Convolvulus ruber Vahl, Eclog. 2:
12. 1798. 7. rubra Millsp. Field Mus. Bot. 2: 86. 1900, non. /. rubra
Murr. 1791. 7. calidicola Standl. & L. Wms. Ceiba 3: 127. 1952;
7. serrulifera Standl. & L. Wms. I.e. 128.
Thickets and open forests at or near sea level; Izabal (Tuerckheim
II. 1128). British Honduras (Schipp 495). Nicaragua; South
American; tropical Africa.
A rather coarse, herbaceous vine, the stems short-hirsute, the other parts of
the plant glabrous; stems and petioles usually minutely aculeolate with short fleshy
protuberances; leaves long-petiolate, about 7-14 cm. long, varying from broadly
ovate-cordate to triangular-hastate, acute or obtuse, the basal lobes obtuse or
acute; inflorescence 1-5-flowered, foliaceous bracts at the bases of the pedicels,
peduncles generally much longer than the petioles, sometimes 20 cm. long, the
pedicels 1.5-2 cm. long; sepals unequal, glabrous, the three outer ones broadly
ovate, 15 mm. long, obtuse or subacute, mucronate, the two inner sepals 12 mm.
long, ovate-lanceolate, acute or acuminate; corolla rose-pink, 7 cm. long, glabrous;
ovary glabrous, 2-celled.
Ipomoea setosa Ker, Bot. Reg. 4: t. 335. 1818. 7. melanotricha
Brandegee, Univ. Cal. Publ. Bot. 4: 381. 1913.
Wet thickets and forest edges, 400 m. or less; Izabal; Santa Rosa.
Mexico; British Honduras; Honduras; Nicaragua. South America.
A large, coarse herbaceous vine, the stout stems and petioles hirsute or setose
with stout, somewhat fleshy trichomes; leaves large and broad, 10-20 cm. long,
deeply 3-7-lobate, cordate at the base, the lobes ovate or lanceolate, long-acumi-
nate; peduncles as long as the petioles or sometimes much longer, 3- 12- flowered;
pedicels 2.5-3 cm. long, thick and fleshy, setose; sepals oblong, obtuse, subcoria-
ceous, setose, 10-14 mm. long, enlarged in fruit; corolla pink or purple, 5-6 cm.
long; capsule 4-celled, 1.5-2 cm. long, 4-celled, 4-seeded; seeds silky pilose.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 53
The type, from which the illustration cited was prepared, was
grown in England from Brazilian seeds. While O'Donell (Lilloa 29:
241. 1959) does not give Central America in the range of the species
it does seem to match the illustration very well.
Ipomoea signata House, Muhlenbergia 3: 46. 1907.
Thickets and forest clearings, 1,500-2,000 m., in the highlands;
Quezaltenango; Huehuetenango (type, Nelson 3595}; Sacatepe"quez ;
Solola; Chimaltenango; Zacapa; El Progreso. Mexico.
Perennial, branched twining vines, the stems mostly 2-3 mm. in diameter,
glabrous or sparsely pubescent, the internodes 6-10 cm. long. Leaves ovate, cor-
date, acuminate, secondary nerves 6-8 pairs and with tertiary nerves connecting
them, the blades 6-14 cm. long and 4-8 cm. broad at maturity, sparsely pubescent
on both sides or glabrate, petioles slender, shorter than the blades; inflorescence
1 -few-flowered cymes, the peduncles mostly 5-10 cm. long but some to 20 cm. long,
pedicels mostly 2-3 cm. long; calyx mostly about 1 cm. long, the outer lobes lance-
ovate, apiculate, deep green with spathulate margins, the inner lobes similar but
longer; corolla salverform, glabrous outside, the tube 4-6 cm. long and 0.5-0.7 cm.
in diameter, the limb spreading and 3-4 cm. broad, stamens reaching the throat
or exserted, the anthers linear-oblong and 4-5 mm. long, pollen grains spinulose,
globose, about 0.1 mm. in diameter; capsules and seeds unknown.
Ipomoea signata has been mostly included in /. morelii, quite a
different species and one that is quite rare in Guatemala. This
species is outstanding among the Guatemalan Ipomoeas in having a
salverform corolla with the stamens exserted or at least reaching the
throat. The species is not uncommon in the highlands.
Ipomoea silvicola House, Bot. Gaz. 43: 411. 1907. Quiebra-
cajete; campanilla; nechao (Huehuetenango).
Moist or wet thickets or thin mixed forest, 600-2,000 m.; Chiqui-
mula; Jalapa; Santa Rosa (type from Rio de las Canas, Heyde & Lux
4.022); Escuintla; Chimaltenango; Guatemala; Solola; Suchitepe"-
quez; Huehuetenango; San Marcos. Southern Mexico; Honduras.
A large or small herbaceous vine, the stems thinly hirsutulous; leaves thin,
long-petiolate, orbicular-cordate, abruptly acuminate, entire or often trilobate,
5-13 cm. long, appressed-pilose above, more densely so beneath; inflorescence usu-
ally 1-3-flowered, peduncles shorter or longer than the petioles, the pedicels usually
long and slender, short-hirsute; sepals subequal, lanceolate, herbaceous, acuminate,
2-3 cm. long, pubescent and densely covered with long spreading green spine-like
tubercles; corolla bright purple, glabrous, 6-7 cm. long, the limb very broad.
House reports the species from Panama on the basis of a collection
made in July, 1860, by Sutton Hayes. A specimen of presumably the
54 FIELDIAN BA:OTANY, VOLUME 24
same collection in the Herbarium of Field Museum is labeled as col-
lected in woods along the road from Escuintla to Antigua, Guate-
mala. So far as we know, the species has not been found in Panama.
Ipomoea squamosa Choisy in DC. Prodr. 9: 376. 1845. 7. mo-
relii Duchass. & Walp. Linnaea 23: 752. 1850. 7. callida House,
Muhlenbergia 3: 42, t. 3. 1907. 7. wilsonii House, I.e. 44, 1. 1. 7. ves-
talii Standl. Contr. Arn. Arb. 5: 130, t. 17. 1933.
From swampy thickets and forest edges at sea level to mountain
forests and thickets at 2,100 meters. Izabal; Escuintla; Quezalte-
nango; Solola; Sacatepe"quez; Chimaltenango ; Guatemala; Jutiapa;
perhaps other departments. Mexico through all of Central America
and Panama to South America.
Twining herbaceous or suffrutescent vines. Leaves triangular in outline, sagit-
tate, or ovate-cordate, long-acuminate, the lateral lobes spreading, somewhat acu-
minate; the sinus broad, 6-12 cm. long and 3-8 cm. broad at the base, glabrous to
sparsely or densely sericeous-pilose pubescent on both surfaces, petioles slender,
pilose or glabrous; inflorescence usually longer than the subtending leaves, a rather
compact few-10-flowered cyme, peduncles to 10 cm. long, pedicels bracteate, slen-
der, ultimate segment to 1 cm. long; sepals coriaceous, unequal, ovate to sub-
orbicular, apex rounded or truncate, scarious marginate, glabrous, the outer ones
about 5 mm. long, the inner ones to about 1 cm. long; corolla infudibuliform, gla-
brous, pink to purple, (4)-7-8 cm. long, stamens inserted near the base of the
corolla and only about 2 cm. long.
We have followed Dr. O'Donell in using 7. squamosa for this
common, perhaps aggregate, Ipomoea. The several synonyms given
above, all of which have been applied to Guatemalan material, are
ones which he has indicated to belong here.
Ipomoea steerei (Standl.) L. Wms. Fieldiana, Bot. 32: 195. 1970
Exogonium steerei Standl. Carnegie Inst. Wash. Publ. 461: 83. 1935.
Known in Guatemala only from thickets in the vicinity of Tikal,
Pete"n. Mexico.
Twining herbaceous or suffrutescent vines with rather coarse stems up to 3-
4 mm. in diameter, densely sericeous-puberulent, becoming glabrous, the inter-
nodes 6-15 cm. long. Leaves elliptic to elliptic-ovate, obtuse, somewhat cuneate
at the base, mostly with 4-6 lateral nerves, the lower surface densely sericeous-
pubescent with appressed hairs and the papillate excrescences with clumps of pu-
bescence, the upper surface appearing green but sparsely appressed sericeous-
pubescent, the petioles short, sericeous- pubescent, 0.8-1.5 cm. long; inflorescences
axillary, a (one-) few-many-flowered dichasium either shorter or longer than the
subtending leaf, up to about 20 cm. long but usually less, the peduncle 1-6 cm.
long, puberulent, the pedicels mostly about 2 cm. long, thickened at the apex;
calyx coriaceous, about 1 cm. long at anthesis and in fruit, glabrous, the lobes un-
equal, the outer ones suborbicular or ovate, apex rounded, 5-8 mm. long, the inner
totftaensfsa
FIG. 6. Ipomoea steerei. A, habit of plant, X %', B, corolla dissected to
show anthers, X 1; C, ovary showing disc below and constriction at base of style,
X 5; D, calyx and pistil, X 1; E, seed with dense coma, X 1^; F, mature capsule,
X 13^; G, under surface of leaf enlarged to show dense sericeous pubescence.
S5
56 FIELDIANA: BOTANY, VOLUME 24
ones suborbicular or ovate, obtuse or retuse, cochleate, about 10 mm. long; corolla
red or carmine, narrowly campanulate, glabrous outside, 5-6 cm. long, the lobes
shallow and rounded, stamens inserted near the base of the corolla and reaching
nearly to the apex, the filaments bearded at the base; pistil glabrous, on a glandular
disc, the style about 5.5 cm. long, contracted and deciduous just above the ovary;
capsule broadly ovoid, with thickened persistent base of the style, the whole about
1.5 cm. long; seeds black, about 5 mm. long with very abundant coma of crisped
hairs attached on the back and near the apex.
The only species in Guatemala with sericeous leaves with obtuse
or cuneate bases and red or carmine flowers.
Ipomoea stolonifera (Cyrill.) Gmel. Syst. Vog. 1: 345. 1796.
Convolvulus littoralis L. Syst. ed. 10. 924. 1759, not Ipomoea littoralis
Blume, 1826. C. stoloniferus Cyrill. PL Rar. Neap. 1: 14. 1788.
/. littoralis Boiss. Fl. Orient. 4: 112. 1879.
Sandy seashores; Izabal. Southeastern United States; Mexico;
British Honduras to Panama; West Indies; South America; Old
World tropics.
Plants prostrate and creeping, succulent, glabrous; leaves long-petiolate, mostly
oblong or oval, rounded at the base, rounded or emarginate at the apex, 3-7 cm.
long, the blades sometimes shallowly or deeply 3-5-lobate; peduncles short, usually
1-flowered; sepals equal or the outer ones shorter, oblong, obtuse, mucronulate,
subcoriaceous, glabrous, 10-15 mm. long; corolla glabrous, 4-5 cm. long, white
with a purple throat; ovary glabrous, 4-celled; capsule globose, 1-1.5 cm. long, with
4 or fewer seeds, these tomentose.
A characteristic member of the seashore vegetation of the At-
lantic coast of Central America, but seldom occurring in abundance.
Ipomoea suffulta (HBK.) G. Don, Gen. Syst. 4: 276. 1838.
Convolulus suffultus HBK. Gen. & Sp. PI. 3: 102, t. 211. 1819.
Trailing over rocks, 800-1,200 m.; Huehuetenango. Southern
Mexico.
Small herbaceous vines, the stems sparsely short hirsute pubescent; leaves or
leaf-like bracts pilose on both surfaces with slender bulbose-based hairs, borne on
peduncles or "short shoots" 1-4 cm. long (and without buds in the axil on the main
stem), rounded-cordate or reniform-cordate, 2-5 cm. long and as broad or broader
than long, sessile on the peduncles and with usually a single flower borne in the axil
of the leaf or leaf-like bract; inflorescence l-(2-) flowered and except the corolla
often hidden in the conduplicate bract; calyx 5-6 mm. long, the sepals unequal, the
outer ones elliptic-oval, the inner ones oblong, obtuse, apiculate, glabrous; corolla
funnelform, 6-8 cm. long, glabrous outside, pink to purple.
A most unusual species in which the leaves or leaf-like bracts are
borne on peduncles or "short shoots." These are sessile or nearly so
and with usually a single nearly sessile flower in the axil. A single
collection is known from Guatemala.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 57
Ipomoea teruae Molina & Williams, Fieldiana Bot 32- 196
1970.
Vine climbing over trees, alt. 2,000 m.; Solola (type Williams,
Molina & Williams 25331). Endemic.
Large twining herbaceous vines, the puberulent stems to 4 mm. or more in
diameter. Leaves suborbicular-ovate, cordate, the apex acute or short acuminate,
the blade 4-7 cm. long and as broad, densely sericeous-pubescent below, glabrous
or nearly so above, petioles 1.5-4 cm. long, pubescent; inflorescence about as long
as or longer than the subtending leaves, dichasioid, few-flowered, peduncle stout,
5-8 cm. long, pedicels 1.5-2 cm. long, stout, puberulent; calyx glabrous, about 1 cm!
long, the lobes broadly ovate to suborbicular, apex obtuse or rounded, the outer
ones only slightly shorter, cochleate, coriaceous, the margins chartaceous; corolla
lilac, 4-5 cm. long, campanulate, glabrous outside, the stamens reaching to about
the middle; capsule and seeds unknown.
This attractive species may be distinguished from all those known
in Guatemala by the sericeous undersurface of the leaves, the sub-
equal, broad and cochleate calyx lobes.
Ipomoea tiliacea (Willd.) Choisy in DC. Prodr. 9: 375. 1845.
Convolvulus tiliaceus Willd. Enum. PL 1: 203. 1809. C. fastigiatus
Roxb. Hort. Beng. 13. 1814. /. fastigiata Sweet, Hort. Brit. 288.
1826. Quilamul.
Moist or dry thickets, often on rocky hillsides or in hedges, fre-
quent in waste places or a weed in cultivated grounds, 1,800 m. or
less; Pete"n; Chiquimula; Santa Rosa; Escuintla; Guatemala; Suchi-
tepe"quez; Retalhuleu; San Marcos; Quezaltenango. Southern Flori-
da; Mexico; British Honduras to El Salvador and Panama; West
Indies; South America.
A large or small, herbaceous vine, glabrous or sparsely pubescent; leaves long-
petiolate, cordate-ovate or rounded-cordate, 5-8 cm. long, acute, entire or trilo-
bate; inflorescences few-several-flowered cymes, peduncles equaling the petioles
or often much longer, the pedicels mostly short; sepals subcoriaceous, oblong to
ovate, mucronate or aristulate, unequal, the larger ones about 8 mm. long; corolla
purple, pink, or rarely white, usually with a dark throat, 3-5 cm. long, glabrous;
capsule subglobose, 2-celled, 8-10 mm. in diameter; seeds glabrous.
Known in El Salvador as "campanola" and "man to de Jesus";
"hebil" (Yucatan, Maya). One of the common weedy species of the
Central American lowlands.
Ipomoea tricolor Cav. Icon. 3: 5, t. 208. 1794. Campanilla;
quiebra-cajete.
Moist thickets, sometimes in hedges or waste ground, occasionally
planted for ornament, 800-1,850 m.; Jalapa; Jutiapa; Santa Rosa;
58 FIELDIANA: BOTANY, VOLUME 24
Guatemala; Chimaltenango. Mexico to Costa Rica; West Indies;
South America.
A small or large, herbaceous vine, glabrous throughout; leaves long-petiolate,
rounded-cordate, thin, entire or obscurely lobate, abruptly acuminate or obtuse;
peduncles stout, hollow, 2-5 cm. long, usually several-flowered, the stout pedicels
5-40 mm. long; sepals oblong-lanceolate, obtuse, white-marginate, 6-7 mm. long;
corolla blue with a white throat, 5-6 cm. long; the limb 6-8 cm. broad; capsule
globose-ovoid, 1-1.5 cm. long, glabrous; seeds glabrous.
The Maya name of Yucatan is reported as "yaxcelil." This
species has been reported from Guatemala under the name 1. violacea
L. The blue flowers are very handsome. Possibly an escape from
cultivation in many places.
Ipomoea trifida (HBK.) G. Don, Gen. Syst. 4: 280. 1838;
O'Donell, Lilloa 30: 58. 1960. Convolvulus trifidus HBK. Nov. Gen.
& Sp. PI. 3: 107. 1819. Campanilla; campana; cajetilla; sisicuch
(Coban, Quecchi).
Moist or wet thickets or hedges, in waste place or in fields, 2,500
m. or less; Pete"n; Alta Verapaz; Izabal; Jutiapa; Zacapa; Jalapa;
Santa Rosa; Escuintla; Guatemala; Sacatepe"quez; Suchitepe"quez;
Retalhuleu; Quezaltenango; San Marcos; doubtless in all depart-
ments. Southern United States; Mexico and British Honduras to
South America.
Small to often large much branched vines, the stems rounded or angled, 1-3
mm. in diameter, pubscent or usually glabrous. Leaves cordate with the blade
ovate, the margins undulate, sometimes to trilobate or rarely 5-lobate, 2-10 cm.
long, the mid-lobe ovate to lanceolate, the sinus deep or shallow, mostly glabrous
(ours) to densely pubescent, the petioles slender, about as long as the blades; in-
florescence cymose, few-(l-) many-flowered, the peduncles 2-25 cm. long, pedicels
3-15 mm. long; sepals coriaceous or straw-like, the outer ones elliptic to ovate,
obtuse or usually long acuminate, glabrous or sparsely pubescent, 4-10 mm. long
and half as broad, the inner sepals elliptic to ovate, acuminate and often cuspidate,
glabrous or sparsely pubescent, 5-12 mm. long and half as broad; corolla infundi-
buliform, glabrous, 2-4 cm. long; anthers reaching to about the middle of the
corolla; capsule subglobose, glabrous or sparsely pubescent, 5-7 mm. long.
A common and widely distributed species needing study. Per-
haps more than one species is included here.
Ipomoea triloba L. Sp. PI. 161. 1753. /. confertiflora Standl.
Carnegie Inst. Wash. Publ. 461: 83. 1935 (type from Rio Grande,
British Honduras, Schipp 1236}. Campanilla; quilamul; resicuch
(Coban, Quecchi).
STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 59
Moist or wet thickets or borders of forest, frequently in hedges or
a weed in cultivated ground, 1,500 m. or less, most abundant at low
elevations; Pete"n; Alta Verapaz; Zacapa; Chiquimula; Jutiapa;
Santa Rosa; Escuintla; Guatemala; Retalhuleu; Solola; Quezalten-
ango; Huehuetenango. Southern United States; Mexico; British
Honduras; El Salvador to Panama; West Indies; South America to
Argentina; naturalized in the Old World tropics.
A small or large, slender, herbaceous vine, sparsely or densely pubescent or
glabrate; leaves long-petiolate, small, mostly 3-5 cm. long but often larger, cordate
at the base, usually deeply 3-5-lobate, the lobes acute or acuminate, entire; in-
florescence 1-several-flowered, peduncles mostly longer than the petioles, the pedi-
cles slender, 1-2 cm. long; sepals oblong or oval, 5-6 mm. long, acute or acuminate
and often mucronate, usually pilose dorsally and long ciliate; corolla pink or pale
purple, with a dark red or dark purple throat, glabrous, commonly (1.5-)2.5-3.5
cm. long, the limb 1-2 cm. broad or more; capsule 2-celled, pilose or glabrous,
about 7 mm. in diameter; seeds glabrous.
This is perhaps the most abundant Ipomoea species of the Pacific
lowlands, where, through much of the year it covers almost com-
pletely thousands of acres of thickets. It is not a pretty plant, and
is decidedly weedy in nature.
This is the oldest name for an exceedingly abundant and variable
group of plants that extend from the southern United States through
Mexico, Central America, the West Indies, and South America to
Argentina. Ipomoea triloba in the broad sense includes /. trichocarpa
Ell. from the southern United States, I. ramonii Choisy from Cuba,
/. con/ertiflora Standl. from British Honduras, and I. trichocarpa var.
australis O'Donell from southern South America.
Ipomoea tuxtlensis House, Ann. N. Y. Acad. Sci. 18: 256. 1908.
Quiebra-cajete.
Moist or wet thickets, often in rocky places, 450 m. or less;
Pete"n; Alta Verapaz; Izabal. Southern Mexico; British Honduras.
A small or large, herbaceous vine, usually very densely pubescent throughout;
leaves long-petiolate, ovate-cordate or rounded-cordate, 5-15 cm. long, deeply
cordate to truncate at the base, entire or usually trilobate, the lobes acute or ob-
tuse, densely appressed-pilose above, densely sericeous beneath with lustrous hairs;
inflorescence few-many-flowered, peduncles very short, usually less than 2 cm. long,
the stout pedicels mostly 4-5 mm. long; sepals subequal, 10-12 mm. long, oblong-
ovate, the outer ones densely sericeous, rounded at the apex, the inner ones gla-
brous or nearly so, retuse; corolla wine-red, 3.5-6 cm. long, narrowly campanulate,
glabrous, the limb 2-3 cm. broad; capsule globose, glabrous, 1 cm. in diameter,
densely lanate with very long hairs.
60 FIELDIANA: BOTANY, VOLUME 24
Ipomoea tyrianthina Lindl. Bot. Reg. 24: Misc. 87. 1838.
Pharbitis tyrianthina Hook. Bot. Mag. 49: t. 4.024. 1843. P. longi-
pedunculata Mart. & Gal. Bull. Acad. 12, pt. 2: 271. 1845. /. longi-
pedunculata Hemsl. Biol. Cent. Am. Bot. 2: 389. 1882. Quiebra cajete.
Moist thickets, 1,600-3,800 m. (?); Quiche^ Huehuetenango;
Chimaltenango. Mexico; Honduras.
A small or large, herbaceous vine, the stems sparsely reflexed-hirsute or gla-
brate; leaves long-petiolate, orbicular-ovate, 6-10 cm. long, abruptly acuminate,
cordate at the base, entire or rarely trilobate, thinly hirtellous or short-pilose;
inflorescence 1-5-flowered, peduncles long and slender, often much longer than the
leaves, the pedicels 1-4 cm. long; sepals subequal, oblong- lanceolate, the outer
ones acute or rarely obtuse, the inner acute, 12-15 mm. long, long pilose pubescent
becoming glabrate above, the hairs usually from thickened bases, or the inner
sepals quite glabrous; corolla rose-purple, 6-8 cm. long, glabrous.
Closely related to Ipomoea purpurea (L.) Roth and perhaps not
distinct from it.
Ipomoea villifera House, Muhlenbergia 5: 70. 1909.
Wet thickets and forest, 1,100-1,400 m., endemic; Huehuetenango
(type, Nelson 3579; Molina 21404).
Stout, twining vines, the stems muriculate, pilose with brownish hairs, be-
coming glabrous. Leaves suborbicular, cordate, entire or trilobate, sparsely
appressed-pubescent above, more densely so below, the blade 7-12 cm. long and
as broad, the petiole slender, pubescent, 5-7 cm. long; inflorescence few-several-
flowered, subcapitate, villous, the peduncle at maturity exceeding the subtending
leaf, villous, the bracts in the inflorescence ovate to ovate-lanceolate, acute,
villous, 1-2.5 cm. long, the pedicels very short; sepals narrowly lanceolate, acumi-
nate, villous, 16-22 mm. long; corolla narrowly campanulate, purple, sparsely
villous outside on the tube and plicae.
A rare species from the mountains of Huehuetenango. Known
only from the two collections cited above. It is closely related to
/. mairetti Choisy.
Ipomoea wallii (Morren) Hemsl. Biol. Cent. Am. Bot. 2: 396.
1882. Batatas wallii Morren, Ann. Soc. Agric. Bot. Gand 2: 285.
1846.
Moist or wet thickets, 300 m. or less; Pete"n (El Paso, Lundell
1532). British Honduras; Honduras.
A small or large, herbaceous vine, glabrous throughout or nearly so; leaves
long-petiolate, cordate-ovate or deltoid-ovate, 6-15 cm. long, acute, rather deeply
cordate at the base; inflorescence usually several-flowered, peduncles usually much
longer than the petioles, sometimes short; sepals more or less unequal, subcoria-
ceous, ovate or oval, 7-8 mm. long, glabrous, rounded at the apex; corolla white,
4.5-6.5 cm. long, glabrous; capsule glabrous, 8 mm. long.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 61
No authentic material of this species is available, and it is not at
all certain that the material we have referred here has been named
correctly.
UNRECOGNIZED SPECIES
IPOMOEA BREVIPES Peter in Engler & Prantl, Pflanzenfam IV
Abt. 3a: 30. 1891. Hyponym.
No description of the species has been published, and it is merely
listed in a skeleton key prepared by Peter to cover a few species of
the genus. The native country is cited as Guatemala. The name,
like some others published by Peter at the same time, has no formal
status.
CALONYCTION CAMPANULATUM Hallier f., Bull. Herb. Boiss. ser.l,
5: 1050, t. 18, f. 2. 1897. Ipomoea setosa campanulata House, Ann
N. Y. Acad. Sci. 18: 219. 1908.
The identity of this species is not certain. We presume that of
the several specimens cited by Hallier the collection made near
Granada, Nicaragua, Levy 1316, and from which the illustration was
made may well be the type although it does not correspond too
closely to the description. There are two possibilities,— the species
may be a synonym of /. setosa as treated in this flora or it may be
the same as /. sepacuitensis but if transferred to Ipomoea it would
become a homonym of /. campanulata L.
CONVOLVULUS MOLLISSIMUS Bertol. Misc. Bot. 20: 11, t. 3. 1859.
The type from Volcdn de Agua, Velasquez.
We do not know what the species may be and the publication
cited is not available to us. We guess that it may be Ipomoea
mairetti.
IPOMOEA ROSTRATA Peter in Engler & Prantl, Pflanzenfam IV
Abt. 3a: 30. 1891, hyponym.
This is said to be based on Guatemalan material. No description
of the plant has been published.
IPOMOEA SERICOPHYLLA Peter, I.e. 31, hyponym.
Based on material said to have come from Guatemala but no
description of the species has been published. Possibly the same as
/. tuxtlensis House.
62 FIELDIANA: BOTANY, VOLUME 24
IPOMOEA TORTUGENSIS Peter, I.e. 31, hyponym.
The name is said to be based on Guatemalan material but the
collection is not mentioned and no description has been published
to validate the name.
ITZAEA Standley & Steyermark
Large woody vines; leaves petiolate, entire, densely sericeous beneath; flowers
small, green, in few-many-flowered, compact, short-pedunculate, axillary cymes,
these shorter than the petioles, or in fruit sometimes elongate; sepals subequal,
suborbicular, rounded or very obtuse at the apex and obscurely apiculate, sericeous
outside, glabrous within; corolla campanulate, shallowly 5-lobate, sericeous out-
side, the lobes induplicate valvate; stamens inserted in the throat of the corolla,
short-exserted, the filaments elongate, slightly dilated at the base; anthers oblong,
the cells contiguous, the connective narrow; ovary 1-celled, 4-ovulate, pilose;
styles 2, short, the stigmas large, globose-capitate; capsule subglobose, somewhat
ligneous, the pericarp dry, 3-4-valvate at the apex, finally separating into numerous
longitudinal linear segments suggesting the staves of a barrel; seeds 3-4, sur-
rounded by bright red pulp, glabrous.
The genus consists of a single species. The generic name is de-
rived from Itza, name of the race that founded the first Maya
Empire.
Itzaea sericea (Standl.) Standl. & Steyerm. Field Mus. Bot. 23:
83. 1944. Lysiostyles sericea Standl. Field Mus. Bot. 8: 322. 1931.
British Honduras; Pete"n, to be expected in Izabal; Atlantic coast
of Honduras, the type from Lancetilla Valley, Dept. Atlantida.
A large woody vine as much as 15 m. long, the stems densely and minutely
sericeous; leaves on slender petioles 1.5-3.5 cm. long, elliptic-oblong or elliptic-
ovate, 7-13 cm. long, 3.5-6.5 cm. broad, obtuse to acuminate, subcordate to acute
at the base, glabrous above, very densely lustrous-sericeous beneath; inflorescences
solitary, very shortly pedunculate, the pedicels mostly 3-4 mm. long; sepals 4 mm.
long; corolla 1 cm. long; capsule subglobose, about 13 mm. in diameter, glabrous.
When fresh, the fruit with its orange-red pulp is strongly sugges-
tive of that of the northern bittersweet (Celastrus scandens).
JACQUEMONTIA Choisy
Twining herbs, often densely pubescent; leaves petiolate, often cordate at the
base, entire or sometimes dentate or lobate; inflorescence axillary, umbelliform or
head-like cymes, rarely in scorpioid cymes or solitary, flowers small or medium-
sized; bracts small and linear or lanceolate or large and leaf like; sepals 5, equal or
unequal; corolla funnelform or campanulate, blue, lilac, or white, 5-dentate or al-
most entire; stamens and style included; style 1, filiform, the 2 stigmas elliptic or
oblong and complanate, rarely globose or linear; ovary 2-celled, 4-ovulate; disk
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA
63
FIG. 7. Itzaea sericea. A, habit, X 1A\ B, capsule from the side showing in-
durated calyx lobes, X 2; C, capsule from above, X 4; D, pubescence on upper
surface of leaf, much enlarged.
small or none; capsule globose, 2-celled, with 4 valves, 4-seeded; seeds glabrous,
tuberculate, or pilose.
About 60 species, in tropical America, Africa, Asia, and Australia.
One or two others are known from southern Central America. The
genus is very closely allied to Convolvulus.
Inflorescences head-like, the bracts large, foliaceous.
Bracts linear or lanceolate; corolla blue J. tamnifolia.
Bracts broadly ovate to rounded; corolla white J. hirtiflora.
64 FIELDIANA: BOTANY, VOLUME 24
Inflorescences cymose, lax and open or rather dense, the bracts small and incon-
spicuous.
Leaves rounded or very obtuse at the apex, obtuse or rounded at the base, thick
and fleshy, glabrous or nearly so J. havanensis.
Leaves acute or acuminate, usually cordate or subcordate at the base, thin, often
densely pubescent.
Sepals rounded at the apex, glabrous or nearly so; peduncles much shorter
than the cymes; corolla white. J. nodiflora.
Sepals acute or acuminate, or at least apiculate, often densely pubescent;
peduncles usually longer than the cymes.
Inflorescence glandular-pilose J. azurea.
Inflorescence without glandular pubescence.
Sepals mostly 10-1 5 mm. long; inflorescence lax J.mexicana.
Sepals all or chiefly 8 mm. long or shorter; inflorescence dense and com-
pact.
Outer bracts ovate J. houseana.
Outer bracts subulate or linear-lanceolate.
Leaves green and glabrate beneath J. pentantha.
Leaves densely velutinous-pubescent beneath.
Calyx 3-4 mm. long; leaves acute or subobtuse J. pinelorum.
Calyx 5-6 mm. long; leaves usually abruptly cuspidate-acuminate.
J. apiculata.
Jacquemontia apiculata House, Muhlenbergia 5: 66. 1909.
Moist or dry thickets, 900 m. or less; El Progreso; Retalhuleu;
Quezaltenango. Mexico; British Honduras.
A herbaceous vine, small or sometimes several meters long, the stems densely
pale-torn entose; leaves long-petiolate, broadly ovate or rounded-ovate, 2-5 cm.
long and often almost as broad, broad at the apex and usually abruptly cuspidate-
acuminate, shallowly or deeply cordate at the base, densely and finely soft-pubes-
cent above, densely torn entose beneath with whitish or brownish, soft hairs; the
inflorescence lax or dense, peduncles 2-8 cm. long, the pedicels short or sometimes
1-2 cm. long, the flowers few or numerous; sepals unequal, the outer ones spatulate-
obovate, 5-6 mm. long, subacute, pubescent, the inner ones shorter, ovate, acute;
corolla blue or white, 2 cm. broad, glabrous.
Jacquemontia azurea (Desr.) Choisy, Mem. Soc. Phys. Hist.
Nat. Geneve 6: 476. 1833. Convolvulus azurea Desr. in Lam. Encycl.
3: 554. 1789. J. guatemalensis Standl. & Steyerm. Field Mus. Bot.
23: 84. 1944 (type from Chiquimula, Steyermark 30066).
Grassy or brushy plains or hillsides, often in rocky places or in
pine forest, 200-1,500 m.; Alta Verapaz; Zacapa; Chiquimula;
Jutiapa; Huehuetenango. Southern Mexico; British Honduras;
Honduras; Costa Rica; Panama; South America.
A slender annual vine, the stems herbaceous, pilose with rather long, soft, more
or less spreading hairs; leaves long-petiolate, ovate-oblong to broadly ovate, 1-5
cm. long, acute to long-acuminate, shallowly cordate or truncate at the base, thinly
or densely pilose with long spreading di-or tribrachiate hairs, especially beneath;
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 65
peduncles about 3-flowered, very slender, often much longer than the leaves, the
pedicels usually much longer than the calyx, glandular-pilose; sepals lanceolate,
3-5 mm. long, pilose with long spreading gland-tipped hairs; corolla usually blue,
6-10 mm. long, glabrous; capsule globose, glabrous, 4-5 mm. long.
When well developed, the plant is elongate and twining. If
stunted, it sometimes is procumbent or even erect.
Jacquemontia havanensis (Jacq.) Urban, Symb. Antill. 3: 342.
1902. Convolvulus havenensis Jacq. Obs. Bot. 2: 25, t. 4.5, /. 3. 1767.
Known in our region only from Turneffe Island, British Hondu-
ras, growing on coral sand, frequent beneath coconut palms along the
beach, F. E. Egler 42-41. Yucatan Peninsula of Mexico; Cuba.
Plants suffrutescent, trailing or suberect, less than a meter long, the stems
puberulent; leaves short-petiolate, succulent, ovate to oblong, 1-4 cm. long,
rounded or subemarginate at the apex, rounded or obtuse at the base, minutely
puberulent or almost glabrous; inflorescences sometimes 1-flowered but usually in
short-pedunculate few-flowered cymes, the peduncles little longer than the
petioles, the pedicels longer than the calyx; outer sepals obovate or suborbicular,
rounded at the apex, 1.5 mm. long, the inner ones slightly longer; corolla 1 cm.
long, white; capsule 3 mm. long, the seeds ovoid, narrowly marginate.
The Maya name of Yucatan is recorded as "extabentuxiu."
Jacquemontia hirtiflora (Mart. & Gal.) O'Donell, Anal. Inst.
Biol. Mex. 12: 81. 1941. Ipomoea hirtiflora Mart. & Gal. Bull. Acad.
Brux. 12, pt. 2: 267. 1845. J. perryana Duchass. & Walp. Linnaea
23: 751. 1850. /. lactescens Seem. Bot. Voy. Herald 171. 1854. J.
platycephala Donn.-Sm. Bot. Gaz. 57: 423. 1914 (type from Cubil-
guitz, Alta Verapaz, Tuerckheim 4133}.
Moist or wet thickets or open rocky forest, 1,500 m. or lower;
Alta Verapaz; Santa Rosa; Huehuetenango. Southern Mexico;
British Honduras; Costa Rica; Panama.
A large or small vine, herbaceous, the stems stout, hirsute or glabrous; leaves
rounded-ovate to deltoid-ovate, 6-13 cm. long, abruptly short-acuminate or some-
times narrowly long-acuminate, shallowly or deeply cordate at the base, glabrous
above or sparsely short-pilose, finely appressed-pilose beneath or glabrate; pedun-
cles often much longer than the leaves, stout, the inflorescences capitate or at least
head-like, 7 cm. broad or smaller, many-flowered, very dense, the outer bracts
about 2 cm. long, orbicular, rounded or very obtuse at the apex, the inner ones
almost as broad, densely hirsute; sepals elliptic, acuminate, 12 mm. long; corolla
pure white, 3-4 cm. long, villous outside.
When in flower, this is a rather handsome plant, much showier
than other Central American species of the genus. Called "mano de
gato" in Veracruz.
66 FIELDIANA: BOTANY, VOLUME 24
Jacquemontia houseana Standl. Field Mus. Bot. 11: 140. 1932.
Known only from the El Cayo District of British Honduras, the
type collected near El Cayo, H. H. Bartlett 12928.
A herbaceous vine, the stems densely stellate-pilosulous; leaves slender-
petiolate, broadly ovate or ovate-elliptic, 2.5-5 cm. long, abruptly acute or acumi-
nate, shallowly or deeply cordate at the base, softly and densely stellate-tomentu-
lose on both surfaces; cymes dense, many-flowered, head-like, about 2.5 cm. broad,
the peduncles as much as 10 cm. long, the flowers sessile, densely crowded; bracts
foliaceous, ovate, equaling the sepals; sepals ovate or broadly ovate, 6-8 mm. long,
acuminate or long-acuminate, densely tomentulose; corolla blue, glabrous, almost
2 cm. broad.
Jacquemontia mexicana (Loes.) Standl. & Steyerm. Field Mus.
Bot. 23: 84. 1944. Aniseia mexicana Loes. Verh. Bot. Ver. Brandenb.
59: 155. 1917.
Brushy hillsides, 400 m. ; Santa Rosa (La Morenita, northeast of
Chiquimulilla, Standley 78873). Chiapas.
A small slender vine, herbaceous or suffrutescent, the young stems tomentu-
lose, becoming glabrate; leaves thin, on long slender petioles, broadly ovate, 3-11
cm. long, acuminate, usually abruptly so, shallowly or deeply cordate at the base,
thinly stellate-pubescent on both surfaces or glabrate; peduncles slender, about
equaling the leaves, lax and few- flowered, the bracts lanceolate, 7-10 mm. long;
outer sepals broadly ovate, 10-15 mm. long, acute or acuminate, puberulent,
usually subcordate at the base; corolla white, 2-3 cm. long, glabrous.
Jacquemontia nodiflora (Desr.) G. Don, Hist. Dichl. PI. 4:
283. 1838. Convolvulus nodiflorus Desr. in Lam. Encycl. 3: 557. 1789.
Campanilla; chata simpatica; loroquito de monte.
Moist or dry thickets or hedges, often in rocky places, 400-1,500
m.; Pete'n; Zacapa; Chiquimula; Jalapa; Jutiapa; Santa Rosa;
Guatemala; Quiche". Mexico; British Honduras; Honduras; Nicara-
gua; Costa Rica; West Indies; South America.
A small or rather large, slender vine, herbaceous or suffrutescent below, the
stems tomentose or almost glabrous; leaves on slender, short or elongate petioles,
oblong-ovate to broadly ovate, 3-5.5 cm. long, acute or acuminate, rounded or
shallowly cordate at the base, sparsely tomentulose or glabrate above, usually
densely tomentulose beneath, sometimes glabrate in age; cymes many-flowered,
dense, short-pedunculate, usually much shorter than the leaves; bracts small and
inconspicuous; sepals suborbicular, rounded and muticous at the apex, 2 mm. long,
glabrous or nearly so; corolla white, 10-12 mm. long, glabrous; capsule glabrous,
globose or ovoid, 4 mm. long.
Recently Lewis and Oliver (Ann. Mo. Bot. Gard. 52: 222. 1965)
have stated that this species must be included in Convolvulus based
upon a study of pollen. We prefer to leave it in Jacquemontia until
•tj
p-l-
FIG. 8. Jacquemontia tamnifolia. A, habit, X 1A\ B, corolla expanded, X 2;
C, calyx and pistil, X 2.
67
68 FIELDIANA: BOTANY, VOLUME 24
a more thorough study is made of generic characters in this group of
Convolvulaceae.
Jacquemontia pentantha (Jacq.) D. Don, Hist. Dichl. PI. 4:
283. 1838. Convolvulus pentanthus Jacq. Coll. Bot. 4: 210. 1790.
J. violacea Choisy, Mem. Soc. Phys. Hist. Nat. Geneve, 61. 1838.
J. violacea var. guatemalensis Meissn. in Mart. Fl. Bras. 7: 296. 1869
(type Friedrichsthal 1).
Dry to wet thickets in second growth, 1,500 m. or less; Suchite-
pe"quez; Retalhuleu; Quezaltenango; Huehuetenango. Southern
Florida; Mexico; British Honduras; Honduras; Costa Rica; Panama;
West Indies; South America.
Usually a small herbaceous vine, the stems pubescent or almost glabrous;
leaves slender -petiolate, ovate or broadly ovate, 3-6 cm. long, acute to long-
acuminate, shallowly or deeply cordate at the base, glabrate, at least in age, some-
what paler beneath; peduncles slender, exceeding the leaves, the cymes dense and
almost head-like, few-many-flowered, the bracts small and inconspicuous, the
pedicels shorter than the calyx; sepals ovate or lanceolate, acute or acuminate,
4-5 mm. long; corolla blue or white, 2 cm. broad, glabrous; capsule globose, about
equaling the sepals.
Maya names of Yucatan are "akilxiu," "sacmiz," "yaax-hebil,"
and "yaxhal."
Jacquemontia pinetorum Standl. & Steyerm. Field Mus. Bot.
22: 372. 1940.
Izabal (type Steyermark 38533).
Plants somewhat woody, prostrate or ascending, branched, the young branches
densely velutinous-pilose; leaves long-petiolate, deltoid-ovate, 2-2.5 cm. long,
acute or subacuminate, subcordate at the base, grayish, densely stellate, velu-
tinous-pilose on both surfaces; peduncles longer than the leaves, the cymes dense,
few-many-flowered, 12 mm. broad, the pedicels short, the bracts small, subulate;
sepals 3-3.5 mm. long, densely pilose, rounded-ovate, muticous, rounded at the
apex; corolla lavender-purple, 8 mm. long, glabrous.
Jacquemontia tamnifolia (L.) Griseb. Fl. Brit. W. Ind. 474.
1861. Ipomoea tamnijolia L. Sp. PI. 162. 1753. Thyella tamnifolia
Raf. Fl. Tellur. 4: 84. 1836.
Moist or dry thickets, about 900 m.; Chiquimula; Jutiapa.
Southern United States; Mexico; British Honduras; Honduras;
Panama.
A small or large, herbaceous vine, sometimes 6 m. long, hirsute almost
throughout, densely so in the inflorescence; leaves slender-petiolate, ovate or
broadly ovate, 3-10 cm. long, acute or abruptly acuminate, cordate at the base,
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 69
FIG. 9. Maripa nicaraguensis. A, habit, X K; B, corolla expanded, X 1J^;
C, calyx and style, X 1J^; D, bud, X 1^; E, pubescence from the corolla, much
enlarged.
sometimes glabrate; peduncles often much longer than the leaves, the inflorescence
head-like, 3-4 cm. broad, many-flowered, the flowers sessile or nearly so; outermost
bracts ovate, 1-3 cm. long; sepals linear, 10-15 mm. long; corolla blue, 1 cm.
broad; capsule globose, glabrous, much exceeded by the sepals.
MARIPA Aublet
Large woody vines, glabrous except in the inflorescence; leaves alternate,
coriaceous, petiolate, entire, not cordate; flowers medium-sized or rather small, in
70 FIELDIANA: BOTANY, VOLUME 24
many-flowered cymes, these usually corymbose or paniculate at the ends of the
branches, the bracts scale-like or none; sepals ovate, obtuse, coriaceous, subequal,
appressed to the base of the fruit; corolla campanulate, the limb plicate, shallowly
5-fid; stamens included or exserted, the anthers oblong or linear; disk annular;
ovary 2-celled, 4-ovulate; fruit ovoid, oblong, or globose, fleshy or indurate, inde-
hiscent, usually by abortion 1-seeded; seed erect, glabrous.
About ten species, all in tropical America. One other Central
American one occurs in Panama.
Maripa nicaraguensis Hemsl. Biol. Cent. Am. Bot. 2: 382. 1882.
Moist or wet forest, 1,000 m. or less; Pete"n; Alta Verapaz; Izabal.
British Honduras, along the Atlantic coast to Costa Rica and South
America.
A large woody vine, climbing to a height of 18 m., glabrous or essentially so
except in the inflorescence; leaves coriaceous, ovate or oblong, 7-12 cm. long,
abruptly acute or cuspidate-acuminate, obtuse to subcordate at the base, short-
petiolate; flowers rose or rose-purple, 3.5-5 cm. long, in small dense terminal
panicles, the branches and pedicels at first ferruginous-puberulent; corolla shortly
5-lobate, strongly constricted below the middle, strigillose-puberulent outside
above the middle, sparsely and minutely lepidote below the middle; stamens
included, the filaments dilated and barbate at the base; ovary glabrous; fruit sub-
globose or broadly ellipsoid, about 3 cm. long, very hard and woody.
A monograph of the genus is being prepared. It is to be ex-
pected that this may be found to be the same species that occurs in
the coastal regions of South America, under an older name.
Called Indian Honey in British Honduras.
MERREMIA Dennstedt
Reference: Carlos A. O'Donell, Revisi6n de las especies americanas
de "Merremia." Lilloa 6: 467-554. 1941.
Plants usually herbaceous and twining, small or often very large; leaves entire,
lobate, or palmately compound and with 3-7 leaflets, glabrous or pubescent;
flowers solitary and axillary or in few-flowered cymes or dichasia, the bracts linear
or lanceolate; sepals usually subequal; corolla campanulate, large or small, white,
yellow, or purple; filaments equal or nearly so, usually glandular at the base, the
anthers spirally twisted in age; ovary usually glabrous, 2-3-carpellate, 4-6-ovulate;
style filiform, the stigma globose or biglobose, never exserted; capsule 2-4-celled,
longitudinally dehiscent by 4-6 valves, the pericarp thin and fragile; seeds 4-6,
glabrous or pubescent.
About 60 species, in the tropics of both hemispheres. Probably
all the Central American species are included in the following treat-
ment.
FIG. 10. Merremia aturiensis. A, habit, X Vz\ B, corolla expanded showing
stamens and pistil, X 2; C, calyx, X 2; D, anther, X 15.
71
72 FIELDIANA: BOTANY, VOLUME 24
Leaves reduced to scales; stems not twining M. aturensis.
Leaves normally developed; stems twining.
Leaves entire.
Flowers umbellate; sepals 6-8 mm. long M. umbellata.
Flowers solitary or on 2-flowered peduncles; sepals 20-30 mm. long.
M. discoidesperma.
Leaves compound or deeply lobate.
Leaves not compound, deeply lobate but the lobes united near the base.
Divisions of the leaves entire; corolla yellow M. tuberose.
Divisions of the leaves lobate; corolla white M. dissecta.
Leaves palmately compound, the segments distinct.
Leaflets entire; pubescence not glandular M. aegyptia.
Leaflets evidently dentate or crenate; pubescence of the calyx of glandular
hairs.
Sepals obtuse M. quinquefolia.
Sepals mostly long-acuminate M. cissoides.
Merremia aegyptia (L.) Urban, Symb. An till. 4: 505. 1910.
Ipomoea aegyptia L. Sp. PL 162. 1753. Convolvulus pentaphyllus L.
Sp. PL ed. 2. 223. 1762. /. pentaphylla Jacq. Coll. Bot. 2: 297. 1788.
M. pentaphylla Hallier f., Bot. Jahrb. 16: 552. 1893.
Moist or wet thickets, sometimes a weed in cornfields, 400 m. or
less; Zacapa; Chiquimula; Santa Rosa. Mexico; British Honduras
to Panama; West Indies; South America; Old World tropics.
A small or large herbaceous vine, the stems usually hirsute with very long,
spreading yellowish hairs; leaves long-petiolate, the 5 leaflets elliptic, entire, short-
petiolulate, acuminate or long-acuminate, acute or acuminate at the base, sparsely
hirsute on both surfaces; inflorescences long-pedunculate, 6-9-flowered, 15-30 cm.
long, hirsute; pedicels 1-3 cm. long; flower buds acute, the sepals oblong, acute,
2 cm. long, hirsute; corolla white, glabrous, 2-3 cm. long; capsule subglobose, 1 cm.
in diameter, 4-celled ; seeds 4, glabrous.
Merremia aturensis (HBK.) Hallier f., Bot. Jahrb. 16: 552.
1893. Convolvulus aturensis HBK. Nov. Gen. & Sp. 3: 96. 1818.
Ipomoea aturensis G. Don, Gen. Syst. 4: 226. 1838. /. aphylla
Standley, Field Mus. Bot. 11: 139. 1932 (type from Cornhouse Creek,
British Honduras, H. H. Bartlett 11316}.
In savannas or other open grassy places, little above sea level;
British Honduras; Mexico (Chiapas); Colombia to Brazil.
Plants erect or ascending, the stems often very numerous, slender, wiry, often
suffrutescent at the base, striate, glabrous; leaves squamiform, 1.5-2 mm. long;
flowers mostly solitary, axillary, the peduncles 1-2 cm. long; sepals oblong-lance-
olate, obtuse, mucronulate, 6-10 mm. long; corolla white, 2.5-3.5 cm. long; anthers
spirally twisted; ovary glabrous, 4-celled; capsule 4-valvate, seeds 4, glabrous;
flower buds acute.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 73
The small scale-like leaves distinguish this plant from all other
Central American members of the Convolvulaceae.
Merremia cissoides (Lam.) Hallier f., Bot. Jahrb. 16: 552. 1893.
Convolvulus cissoides Lam. Tabl. Encyl. 1: 462. 1791. Ipomoea cis-
soides Griseb. Fl. Brit. W. Ind. 473. 1861.
In thickets or waste ground, 1,800 m. or less; Santa Rosa; Suchi-
tepe"quez. Mexico; British Honduras; El Salvador and to Costa Rica;
West Indies; South America.
A large or small, herbaceous vine, the stems hirsute-pilose with rather short
hairs or sometimes glabrate; leaves long-petiolate, the 5 leaflets elliptic or lance-
olate, mostly 2-4.5 cm. long, short-petiolulate, acute or acuminate, shallowly
dentate or entire, viscid-puberulent or glabrate; flowers usually in few-flowered
cymes, rarely solitary, the cymes axillary, usually about equaling the leaves;
flower buds acute; sepals 10-15 mm. long, ovate or lance-ovate, long-acuminate,
foliaceous, viscid-pubescent; corolla white, 2-3 cm. long; ovary glabrous, 3-4-
celled, 3-4-ovulate; capsule 7-8 mm. broad, the seeds black, pubescent.
The Maya name of Yucatan is reported as "kixoloc"; "bejuco
coronal" (El Salvador).
Merremia discoidesperma (Donn.-Sm.) O'Donell, Lilloa 6: 495.
1941. Ipomoea discoidesperma Donn.-Sm. Bot. Gaz. 27. 1889.
Quiebra-cajete.
Wet or mixed forest, 2,600 m. or lower; Izabal; Alta Verapaz
(type from Pansamala, Tuerckheim 744)', Huehuetenango. Mexico
(Chiapas); Nicaragua (?); Cuba; Hispaniola.
Plants stout, twining, glabrous, the stems sometimes several cm. in diameter;
leaves on petioles 3-6 cm. long, oval-cordate, 10-15 cm. long, 3-10 cm. broad,
entire, coriaceous, acute or acuminate; flowers solitary or the peduncles 2-flowered;
sepals oblong, 2-3 cm. long; corolla yellow, 4-5.5 cm. long, glabrous; capsule very
large, 3-4 cm. in diameter, subtended by the accrescent and persistent sepals, these
in fruit 5-6 cm. long; capsule thin, irregularly dehiscent; seeds usually 1, depressed,
2-3 cm. in diameter, black, velutinous-pubescent.
Merremia dissecta (Jacq.) Hallier f., Bot. Jahrb. 16: 552. 1893.
Convolvulus dissectus Jacq. Obs. Bot. 2: 4, t. 28. 1767. Ipomoea
sinuata Ortega, Hort. Matr. Dec. 7: 84. 1798. Operculina dissecta
House, Bull. Torr. Bot. Club 33: 500. 1906. /. dissecta Pursh, Fl.
Amer. Sept. 145. 1814.
Dry to wet thickets, 400 m. or less; El Progreso; Chiquimula;
Zacapa. Southern United States; Mexico; Honduras and El Salvador
to Panama; West Indies; South America.
74 FIELDIANA: BOTANY, VOLUME 24
A small or large, herbaceous vine, the stems hirsute or glabrous; leaves palm-
ately divided almost to the base, the 7-9 segments coarsely sinuate dentate, usually
glabrous, sometimes sparsely hirsute; flowers solitary or in 2-4-flowered cymes, the
peduncles 5-7 cm. long; pedicels thickened at the apex; sepals 18-25 mm. long,
oblong, mucronate, glabrous; corolla white, 3-4.5 cm. long; anthers spirally
twisted; ovary glabrous, 2-celled, 4-ovulate; capsule 1.5 cm. in diameter, depressed-
globose, surrounded by the accrescent calyx, the sepals in fruit sometimes 3 cm.
long; seeds 4, black, glabrous, 8 mm. long.
Merremia quinquefolia (L.) Hallier f., Bot. Jahrb. 16: 552.
1893. Ipomoea quinquefolia L. Sp. PI. 162. 1753. Manecilla blanca
(fide Aguilar).
Moist or dry thickets or hedges, sometimes in open fields, 1,500
m. or less; Izabal; Zacapa; Chiquimula; El Progreso; Jutiapa; Santa
Rosa; Escuintla; Guatemala; Retalhuleu; San Marcos. Mexico;
British Honduras to El Salvador and Panama; West Indies; South
America.
Usually a small herbaceous vine, sometimes prostrate, the stems glabrous or
sparsely hirsute; leaves long-petiolate, the 5 leaflets subsessile, elliptic or lance-
olate, glabrous, acute or acuminate, sinuate-dentate, 2-7 cm. long; inflorescence a
simple 3-flowered, or rarely compound, dichasium or reduced to a single flower,
axillary and usually longer than the leaf, the peduncles glandular-pilose, the pedi-
cels 1-1.5 cm. long; sepals unequal, oblong, obtuse, glabrous, the outer ones 3-5
mm. long, the inner ones 4-7 mm. long; corolla white, 1.5-2.5 cm. long; capsule
glabrous, 7-8 mm. broad, subglobose, 4-celled; seeds 4, pubescent.
Called "white cowslip" in British Honduras.
Merremia tuberosa (L.) Rendle in Thist.-Dyer, Fl. Trop. Africa
4: 104. 1905. Ipomoea tuberosa L. Sp. PL 160. 1753. Operculina
tuberosa Meissn. in Mart. Fl. Bras. 7: 212. 1869. 7. nuda Peter in
Engler & Prentl, Pflanzenf. IV. 3a: 31. 1891, hyponym (based on
Guatemalan material). Quiebra-machete (Suchitepe"quez; probably an
erroneous rendering of quiebra-cajete); rosa de barranco; foco de luz;
bejuco de golondrina; quinamacal (Pete*n).
Moist or wet thickets, 1,200 m. or lower; Pete"n; Alta Verapaz;
Baja Verapaz; Izabal; Escuintla; Suchitepe"quez; Retalhuleu; Quiche";
Quezaltenango; San Marcos. Mexico; British Honduras to Panama;
West Indies; South America.
Usually a very large, herbaceous vine, often growing over good-sized trees, the
stems stout, glabrous or nearly so; leaves large, long-petiolate, 8-15 cm. long or
more, deeply 7-lobate, often almost to the base, the lobes lanceolate to elliptic,
acuminate, entire, glabrous; peduncles 10-20 cm. long, the pedicels 1-2 cm. long,
clavate, the flowers usually several in each inflorescence, sometimes solitary; sepals
unequal, the outer ones 2.5-3 cm. long in anthesis, oblong, obtuse, mucronate;
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 75
corolla bright yellow, 5-6 cm. long; anthers spirally twisted; capsule irregularly
dehiscent, with 4 or fewer seeds, 2-celled; sepals in fruit as much as 7 cm. long;
seeds to 17 mm. in diameter, covered with a very dense, short and black pube-
scence.
Called "seven-fingers" in British Honduras. In Honduras the
vine is called "mala hierba," it being claimed that the huge tuberous
roots are poisonous to pigs and horses. In the Canal Zone this is
known as "Tivoli vine," because it was formerly at least planted
about the Tivoli Hotel in Panama City. It is one of the more showy
of Central American morning-glories and under favorable conditions
attains a great size and is covered with the large blossoms. In Guate-
mala the dry capsules with their enveloping sepals are much used as
decorations in houses or on alters, either in their natural brown color
or embellished with silver or gold paint.
Merremia umbellata (L.) Hallier f., Bot. Jahrb. 16: 552. 1893.
Convolvulus umbellatus L. Sp. PL 155. 1753. Ipomoea polyanthes
Roem. & Schult. Syst. 4: 234. 1819. Cajete; quilamulillo (fide
Aguilar) .
Dry to wet thickets or hedges, often in second growth, or a weed
in cultivated ground, 1,500 m. or less; Pete"n; Alta Verapaz; Baja
Verapaz; Izabal; El Progreso; Zacapa; Jutiapa; Santa Rosa; Es-
cuintla; Guatemala; Solola; Suchitepe"quez; Retalhuleu; San Marcos.
Mexico; British Honduras to El Salvador and Panama; West Indies;
South America; Old World tropics.
A small or large, herbaceous vine, the stems slender, glabrate; leaves long-
petiolate, narrowly triangular to broadly ovate, usually long-acuminate, cordate or
hastate at the base, densely pubescent or almost glabrous, entire; inflorescence
umbelliform, usually many-flowered, peduncles 6-15 cm. long; flowers buds obtuse;
sepals equal, oblong, 6-8 mm. long, glabrous or pubescent, rounded at the apex,
scarious-margined ; corolla bright yellow, 3-3.5 cm. long, glabrous; anthers not
twisted; ovary glabrous, globose, 2-celled, 4-ovulate; capsule 8 mm. in diameter,
regularly 4-valvate; seeds 4 and 5-6 mm. long, densely pubescent.
Known in El Salvador as "cuelga-tabaco," "jicama cimarrona,"
and "cuajo de ule." The tough stems are used commonly for hanging
tobacco to dry, while the milky sap has been utilized for coagulating
Castilla rubber sap. The plant is a common weed through much of
Central America. The abundant yellow flowers, in head-like clusters,
are rather pretty, and they apparently remain open throughout the
day.
76- FIELDIANA: BOTANY, VOLUME 24
OPERCULINA Manso
Reference: H. D. House, Studies in the North American Con-
volvulaceae, II. The genus Operculina. Bull. Torr. Bot. Club 33:
495-503. 1906.
Large or small vines, herbaceous or somewhat woody, the stems, petioles, and
pedicels often winged; leaves entire or variously lobate, often cordate; flowers
large, axillary, in few-flowered cymes or solitary; sepals large, scarious, glabrous,
enlarged in fruit and becoming somewhat coriaceous, often irregularly erose on the
margins; disk small or none; corolla large, broadly funnelform or campanulate,
white, yellow, or salmon-red; stamens and style included, the anthers twisted, at
least in age; ovary glabrous, each cell with 2 ovules, style one, simple, filiform, in-
cluded, the stigma biglobose; capsule large, dry, epicarp circumscissle in or above
the middle, the upper part more or less fleshy and separating from the lower part
and from the endocarp.
Species perhaps 20 in the tropics of both hemispheres. One or
two additional species in Mexico, no others are known from Central
America.
Leaves entire, cordate; peduncles winged 0. pteripes.
Leaves digitate, lobate or dentate; peduncles not winged O. pinnatifida.
Operculina pinnatifida (HBK.) O'Donell, Lilloa 23: 432. 1950.
Convolvulus pinnatifida HBK. Nov. Gen. & Sp. PL 3: 85. 1819.
Ipomoea pinnatifida G. Don, Gen. Syst. 4: 280. 1838. Merremia
pinnatifida Hallier f. in Engler, Bot. Jahrb. 16: 552. 1893. Ipomoea
ornithopoda Robinson, Proc. Am. Acad. 19: 183. 1893. Operculina
ornithopoda House, Bot. Gaz. 44: 414. 1907. Operculina pectinata
House, Muhlenbergia 5: 69. 1909. Quiebra cajete.
A vine in thickets or growing over trees at 300 m. or less; Pete"n;
Huehuetanango. Texas; Mexico (widely distributed) .
A glabrous rampant vine, the internodes of the stems to 15 cm. long. Leaves
with petiole shorter than the blade, the blade ovate in outline, 2-12 cm. long and
about as broad, with 5-9 pinnate divisions or lobes, the segments linear to ovate
or rarely obovate and sometimes lobulate, acute or acuminate, the base cordate or
sub truncate; inflorescence a simple, axillary 2-3-flowered dischasium or cyme,
sometimes with a single flower, the peduncles 3-12 cm. long, sometimes subalate
above; sepals obovate, obtuse, glabrous, 15-16 mm. long and 10-13 mm. broad;
corolla white, subcampanulate, 3-5 cm. long and about as broad, the interpetaline
area sericeous; capsules subglobose-depressed, 1.5-2 cm. in diameter; seeds black,
6-7 mm. long.
Operculina pteripes (G. Don) O'Donell, Lilloa 23: 435. 1950.
Calonyction pteripes G. Don, Gen. Syst. 4: 264. 1838. Ipomoea
FIG. 11. Operculina pteripes. A, habit and bud, all natural size.
77
78 FIELDIANA: BOTANY, VOLUME 24
alatipes Hook. Bot. Mag. 88: t. 5330. 1862. Operculina alatipes
House, Bull. Torr. Bot. Club 33: 499. 1906. Chorreque (Jutiapa).
A vine in thickets, usually 950 m. or less; Chiquimula; Jutiapa;
Santa Rosa; Escuintla; perhaps in all lowland departments. Mexico
to El Salvador and Panama. South America.
A large herbaceous vine, growing over good-sized trees, glabrous throughout;
leaves long petiolate, ovate, mostly 3-8 cm. long, long-acuminate, cordate at the
base, entire; peduncles narrowly winged, 2-7-flowered, usually strongly curved, the
pedicels thickened; calyx 2.5 cm. long, the sepals acute or acuminate; corolla
almost salverform, 6-7 cm. long, salmon-colored or brick-red.
Known in El Salvador as "mechoacan," "quiebra-cajete," and
"campanilla chocolate." The large turnip-like roots often weigh
several pounds. Among all the related plants of Guatemala, this may
be recognized at once by the unusual color of the flowers, which are
attractive.
QUAMOCLIT Moench
References: H. D. House, Studies in North American Convolvu-
laceae— V. Quamoclit, Bull. Torr. Club 36: 595-603. 1909. C. A.
O'Donell, Las especies americanas de Ipomoea L., Sect. "Quamo-
clit" (Moench.) Griseb., Lilloa29: 19-86. 1959.
Herbaceous vines; leaves alternate, entire, lobate, or pinnately divided;
flowers small or large, often showy, usually red, yellow, or greenish, axillary, in
often dichotomous, few-many-flowered cymes; sepals membranaceous or herba-
ceous, equal or nearly so, often appendaged at the apex; corolla tube short or elon-
gate, the limb more or less salverform or funnelform, shallowly or rather deeply
lobate; stamens and style exserted, usually declinate, the filaments filiform; ovary
2-celled, 4-ovulate; stigma capitate; capsule globose or somewhat elongate, the
style often persistent; seeds glabrous.
About a dozen species, in tropical America. Only the following
are known from Central America. The genus perhaps is not distinct
from Ipomoea.
Leaves pinnately parted into linear segments; sepals not awned at apex.
Q. pennata.
Leaves entire or palmately lobed; sepals awned at apex.
Corolla limb suburceolate with very short tube; inflorescence a bifid cyme with
secund flowers Q. lobata.
Corolla limb campanulate, tube long or short; inflorescence cymose or corymbi-
form, the flowers not secund.
Limb of corolla deeply lobed, the lobes 0.7-1.5 cm. long Q. vitifolia.
Limb of corolla subentire or shallowly lobed, the lobes broad and short.
Branches of inflorescence short, corymbiform; corolla yellow or sometimes
reddish Q. lutea.
Branches of inflorescence long; corollas red or reddish.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 79
Leaves lobate Q. hederifolia.
Leaves not lobate, rarely dentate Q. cholulensis.
Quamoclit cholulensis (HBK.) G. Don, Gen. Hist. 4: 259.
1838. Ipomoea cholulensis HBK. Nov. Gen. & Sp. PL 3: 88. 1819;
O'Donell, Lilloa 29: 28. 1959. Q. coccinea var. pubescens (Schlecht.
& Cham.) G. Don, I.e. 258. Q. coccinea of authors, in part. Camp-
anita roja.
Vines growing in thickets or over other plants, often weedy,
1,000-1,500 m.; Alta Verapaz; Sacatepe"quez ; Guatemala; Chimal-
tenango; Santa Rosa. El Salvador; Honduras; Nicaragua; Costa
Rica. South America.
An annual or possibly more enduring, much branched, glabrous or tomentose.
Leaves ovate or lanceolate-ovate, cordate, entire or sometimes dentate, acute to
acuminate or caudate, tomentose to pilose or nearly glabrous, 2-12 cm. long and
1-7 cm. broad; inflorescence cymose, 2-8-flowered or rarely the flowers solitary,
peduncles 7-19 cm. long; pedicels 0.5-1.5 cm. long; exterior sepals oblong or ovate,
2.5-3.5 mm. long and 1.5-2.5 mm. broad, obtuse with a subterminal awn, glabrous
or pubescent; the interior sepals elliptic to ovate, 3-4.5 mm. long, 2-3 mm. broad,
awned; corolla hypercratiform, tube slightly spread at the apex, 1.5-2.5 cm. long
and 0.15-0.3 cm. in diameter, limb broadly campanulate, obscurely 5-lobate;
stamens barely exserted; ovary ovoid, 4-locular, 4-ovulate; stigma biglobose;
capsules subglobose.
This is one component of the complex that has been called Q.
coccinea (L.) Moench. Most Central American material has been
determined with that name. O'Donell, in his revision of this group,
limits Q. coccinea to the United States.
Quamoclit hederifolia (L.) G. Don, Gen. Hist. 4: 259. 1838.
Ipomoea hederifolia L. Syst. Nat. ed. 10. 925. 1759. Q. brevipedicellata
Hallier f. Bull. Herb. Boiss. 7: 416. 1899. Q. coccinea var. hederifolia
House, Ann. N. Y. Acad. Sci. 18: 262. 1908. Q. coccinea, of many
authors, in part. Campanita; xocxop (Huehuetenango).
Moist thickets or hedges, often a weed in cornfields, 1,800 m. or
less; Zacapa; Chiquimula; Jutiapa; Santa Rosa; Escuintla; Sacate-
pe"quez; Chimaltenango; Retalhuleu; Huehuetanango. Southern
United States; Mexico; British Honduras to Panama; West Indies;
South America; Malaysia; Africa.
Usually a small herbaceous vine, glabrous to laxly pubescent; leaves long-
petiolate, ovate or rounded-ovate, 4-10 cm. long; acute or acuminate, cordate at
the base, (rarely) entire, shallowly to deeply angulate-lobate; peduncles equaling
or longer than the leaves, few-many-flowered; sepals oblong, 4-6 mm. long, the
outer ones with subulate awns 2-5 mm. long arising on the outside below the apex;
FIG. 12. Quamoclit hederifolia. A, habit, X 1A; B, calyx, X 2; corolla and pistil,
natural size (after Flora Brasiliensis). Ipomoea pes-caprae. D, habit, X V4.
80
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 81
corolla scarlet, the tube 3-4 cm. long, slightly dilated near the apex, the almost
rotate limb 1.5-2 cm. broad, 5-angulate; capsule globose, 6-7 mm. long.
Called "Indian creeper" in British Honduras; "hiedra colorada"
(Yucatan). The plant is a weedy one, not very decorative, in spite
of its brilliantly colored flowers.
Quamoclit lobata (Llave & Lex.) House, Bull. Torr. Bot. Club
36: 602. 1909. Mina lobata Llave & Lex. Nov. Veg. Descr. 3. 1824.
Ipomoea versicolor Meissn. in Mart. Fl. Bras. 7: 220. 1869. Bandera
espanola; banderita de Espana; banderita espanola.
Cultivated occasionally for ornament. Perhaps native of Mexico.
A slender herbaceous vine; leaves slender-petiolate, ovate, deeply cordate at
the base, deeply 3-lobate, the middle lobe constricted near the base, the lateral
lobes often lobate, glabrous or nearly so; peduncles much longer than the leaves,
dichotomous, few-many-flowered, the flowers secund; sepals 3-4 mm. long, the
apical appendages recurved; corolla bright yellow, tinged above with red, about
2 cm. long, the tube proper very short, scarcely exceeding the calyx, the throat
much dilated, 1 cm. broad, the margin somewhat constricted, with 5 very small
and short lobes; stamens twice as long as the corolla.
A showy and handsome plant, deserving wide cultivation. The
plants wither when the rains stop, but if watered might remain in
bloom for a longer period. The name "bandera espanola" is given
because the corollas bear the colors of the flag of Spain.
Quamoclit lutea (Hemsl.) Hallier f. in Engler, Bot. Jahrb. 16:
557. 1893. Ipomoea lutea Hemsl. Diagn. PI. Nov. 34. 1878 (type from
Guatemala, Salvin & Godman); in Godman & Salvin, Biol. Cent. Am.
Bot. 2: 389, t. 60. 1882. /. lutea var. rubra O'Donell, Lilloa 29: 67.
1959.
Moist thickets or forest, 1,300-1,500 m.; Guatemala; Chimal-
tenango; Sacate"pequez; Quezaltenango; San Marcos; Suchitipe"quez.
Southern Mexico.
A large vine, glabrous throughout or nearly so, herbaceous; leaves long petio-
late, cordate or hastate at the base, sometimes broadly ovate and entire, usually
3-lobate, the lobes acute to long-acuminate, the outer ones often angulate; ped-
uncles greatly elongate, the flowers usually numerous, long-pedicellate, in small
compact cymes; sepals 5 mm. long, the slender appendages of equal or greater
length; corolla yellow or orange-yellow, 5 cm. long, the tube slightly curved, very
gradually dilated upward, 7-8 mm. broad in the throat, the lobes rounded, small,
scarcely more than 8 mm. long; stamens shortly exserted.
This species was much misunderstood by House, who treated it
as a variety of Q. coccinea, from which it is altogether distinct.
82 FIELDIANA: BOTANY, VOLUME 24
Quamoclit pennata (Desr.) Bojer, Hort. Maurit. 224. 1837.
Ipomoea Quamoclit L. Sp. PI. 159. 1753. Convolvulus pennatus Desr.
in Lam. Encycl. 3: 567. 1791. Q. vulgaris Choisy, Mem. Soc. Phys.
Hist. Nat. Geneve 6: 434. 1833. Clarintitto; clarin.
Probably native in tropical Asia, but planted in many temperate
and tropical regions for ornament; grown commonly in Guatemalan
gardens, at low and middle elevations; naturalized in many places,
usually in moist thickets or in hedges, 1,500 m. or less; Izabal;
Zacapa; Chiquimula; Jutiapa; Escuintla; Guatemala; Suchitepe"-
quez; Retalhuleu. British Honduras. Naturalized widely in trop-
ical America.
A slender glabrous vine, often several meters long; leaves 2-10 cm. long,
parted to the costa into very numerous linear segments; peduncles mostly longer
than the leaves and 1-2-flowered, the pedicels 2-3 cm. long, thickened in fruit;
sepals oblong, obtuse, mucronate, not appendaged, 3-5 mm. long; corolla scarlet,
2.5-3.5 cm. long, the tube slender, the limb rotate, with short ovate acute lobes;
capsule ovoid, 8-10 mm. long.
Known in El Salvador by the names "clavellina," "cundeamor,"
and "cambustera" (Yucatan). The plant is often grown in the
United States, where usually it is called "cypress vine." Plants with
white flowers sometimes are found in cultivation, but they have not
been noted in Central America.
The specific name of this plant is often spelled pinnata, which is
the form most commonly used in botanical Latin. The original,
however, uses the form we have followed which is preferred in classi-
cal Latin.
Quamoclit vitifolia (Cav.) G. Don, Gen. Syst. 4: 259. 1838.
Calboa vitifolia Cav. Icon. 5: 51, i. 4.76. 1799. Ipomoea pedun-
cularis Bertol. Fl. Guat. 408, t. 38. 1840 (type from Escuintla,
Velasquez) . Ipomoea hartwegii Meissn. in Mart. Fl. Bras. 7: 220. 1869
(syntype from Guatemala, Hartweg 603). Bejuco de mariposa.
Usually in moist or wet thickets, often in second growth, 1,500 m.
or less; Alta Verapaz; Zacapa; Jutiapa; Santa Rosa; Escuintla;
Guatemala; Sacatepe*quez ; Suchitepe"quez ; Retalhuleu; Quezalten-
ango; San Marcos. Mexico; El Salvador to Panama.
Usually a large coarse herbaceous vine with milky sap, often climbing over
medium-sized trees, glabrous; leaves long-petiolate, about as broad as long, cordate
or cordate-sagittate at the base, rarely subentire but usually deeply trilobate, the
lobes ovate to lanceolate, acute to long-acuminate; inflorescence few-many-
flowered, the flowers in dense or open cymes, peduncles usually much longer than
the leaves; sepals ovate or rounded, 4-5 mm. long, the terminal appendages of less
or equal length; corolla 2.5-3.5 cm. long, thick in texture, bronze-yellow, brownish
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 83
yellow or pale greenish yellow, the lobes and upper part of the tube usually tinged
with maroon or brown, the tube proper short and thick, the throat abruptly dilated
and funnelform, the limb deeply 5 lobate, the lobes ovate, obtuse or acutish,
ascending; stamens long-exserted ; capsule globose-ovoid, 7 mm. long.
Called "campanilla" and "ala de cucaracha" in El Salvador. This
plant is less conspicuous than other species of Quamoclit, the color of
the flowers being bizarre but rather dull.
This species has often been considered to include Q. globosa (Llave
& Lex.) G. Don or that is a distinct species. O'Donell considers Q.
globosa a synonym of Ipomoea hastigera HBK., a species of Quamoclit
as delimited here, limited to a small area in Mexico.
TURBINA Rafinesque
Woody vines, pubescent or glabrous; leaves petiolate, cordate and entire;
peduncles axillary, 1-many-flowered, often paniculate at the ends of the branches;
sepals ovate or lanceolate, the outer ones prominently sphacelate, often very un-
equal and accrescent in fruit; corolla usually large and funnelform; stamens and
style usually not exserted; ovary glabrous, with 2 cells; stigmas 2, globose; fruit dry,
usually ligneous, globose or ellipsoid, 1-celled and commonly 1 seeded; seeds
glabrous.
Perhaps a half dozen species in America. Only the following is
known from Central America.
S. J. van Ooststroom, Blumea 5: 355. 1943, points out that our
species does not belong in the genus Rivea Choisy and transferred it
to Legendrea, a genus more recent than Turbina, with the same
circumscription.
Turbina corymbosa (L.) Raf. Fl. Tell. 11: 81. 1836. Convolvulus
corymbosa L. Syst. Nat. ed. 10. 293. 1759. Rivea corymbosa Hallier f.
in Engler, Bot. Jahrb. 18: 157. 1893. Convolvulus sidaefolia HBK.
Nov. Gen. & Sp. 3: 99. 1818. Ipomoea sidaefolia Choisy, Mem. Soc.
Phys. Hist. Nat. Geneve 6: 459. 1833. Xtabentun (Pete"n, Maya).
Moist or wet thickets or hedges, often in second growth, 650 m.
or less; Pete"n; Alta Verapaz; Izabal; El Progreso; Zacapa; Santa
Rosa. Mexico; British Honduras to Panama; West Indies; South
America. Introduced in Phillipines.
A large or small, woody vine, often climbing over small trees, glabrous or
nearly so; leaves slender-petiolate, ovate, 4-10 cm. long, acute or acuminate,
cordate at the base, entire; peduncles axillary, equaling or longer than the leaves,
few-many-flowered, often forming large dense terminal panicles, the pedicels
slender; sepals oblong, persistent, the inner ones 8-12 mm. long, almost twice as
FIG. 13. Turbina corymbosa. A, habit, X 1A; B, dissected corolla showing
stamens and pistil, natural size; C, calyx, X 4.
84
GIBSON: FLORA OF GUATEMALA 85
long as the outer ones; corolla white, 2.5-3 cm. long; capsule ovoid-oblong, acute,
shorter than the inner sepals, 1-seeded.
Called "pascua" and "flor de pascua" in Yucatan, presumably in
allusion to the fact that it is in bloom there at Christmas time. In
Panama the latex is said to be employed for coagulating Castillo,
rubber latex.
Excluded:
RIVEA BERNOULLIANA (Peter) Hallier f. in Engler, Bot. Jahrb. 18:
158. 1893. Ipomoea bernoulliana Peter in Engler & Prantl, Pflanzenf.
IV. Abt. 3a: 30. 1891, nomen nudum.
This name is based upon material from Guatemala collected by
Bernoulli but no description of the plant has been published.
POLEMONIACEAE. Phlox Family
DOROTHY N. GIBSON
REFERENCES: Benth. in DC. Prodr. 9: 302-322. 1845. Benth. &
Hook., Gen. PL 2: 820-824. 1876. Brand in Engler, Pflanzenr. IV,
250: 1-203. 1907. Grant, Natural History of the Phlox Family 1-
1-273. 1959.
Annual or perennial herbs, shrubs or small trees, usually erect (scandent in
Cobaea), usually somewhat pubescent, often glandular; leaves alternate (lower ones
sometimes opposite), entire, serrate, or pinnately divided; inflorescences usually
cymose, the flowers rarely solitary and axillary; flowers bisexual, actinomorphic
or weakly zygomorphic; calyx 5-lobed, imbricate or valvate, persistent, usually
accrescent; corolla tubular (campanulate in Cobaea), 5-lobed, lobes contorted,
salverform to rotate; stamens 5, inserted on corolla tube at various, often unequal
heights, anthers versatile, bilocular, introrse; ovary superior, inserted on disc,
usually trilocular (rarely 2 or 5 carpels), placentation axile, ovules 1-many on
each placenta; style filiform, stigmatic branches 3 (rarely 2 or 5); fruit a capsule
dehiscing loculicidally (septicidally in Cobaea) or rarely indehiscent; seeds usually
small, sometimes winged, often mucilaginous when wetted; embryo spatulate; en-
dosperm usually firm, abundant.
The Polemoniaceae, containing 317 reported species, are naturally
distributed throughout North, Central and South America and Eura-
sia, occurring most frequently in temperate regions. A few occur as
86 FIELDIANA: BOTANY, VOLUME 24
weeds in Africa and Australia. Of the 15 to 18 genera (not always
very distinct) only three, Bonplandia, Loeselia, and Cobaea are known
from Central America. Although Gilia capitata Sims and Phlox
Drummondii Hook., are cultivated occasionally for ornament and
sometimes sold in the markets, neither has yet been reported as an
escape.
Plants scandent by tendrils Cobaea.
Plants not scandent, without tendrils.
Calyx foliaceous; leaves lyrate lobate or pinnately lobed Bonplandia.
Calyx membranaceous between costae; leaves dentate or serrate Loeselia.
BONPLANDIA Cav.
REFERENCE: A. Brand in Engler, Pflanzenr. IV, 250: 179-181,
/. 39. 1907.
Annual herbs, usually erect, sometimes becoming bushy, more or less glandu-
lar-pubescent throughout; leaves alternate, simple, pinnately divided, entire or
with serrate margins, sessile or petiolate; inflorescence usually loose, terminal
panicles composed of axillary, short-pedicellate flowers usually in pairs (rarely 1
or 3) or flowers may occur in axils without becoming paniculate; calyx persistent,
herbaceous, tubular, slightly zygomorphic; corolla blue to purple, bilabiate, the
tube gibbous, the lobes unequal, about equalling or shorter than the tube; stamens
glabrous, inserted at different levels on upper part of tube, about equalling the
corolla or shortly exserted; ovary broadly ellipsoid to ovoid, superior, on small disc;
mature capsule elliptic-ovoid, glabrous, much shorter than the accrescent calyx,
dehiscing loculicidally, each locule containing one small, oblong, narrowly winged
seed.
Only one of the two known species is in Guatemala.
Bonplandia geminiflora Cav. Anal. Hist. Nat. Madrid 2: 131.
1800. Caldasia heterophylla Willd. Hort. Berol. t. 71, 1807; Benth.
in DC. Prodr. 9: 320. 1845.
Known in Guatemala only from Azacualapa, Jutiapa, 400 m.,
Heyde & Lux 6223. Widely distributed in Mexico, 270-3,150 m., in
wet, shady places, sometimes growing in water, or in rocky canyons.
Plants 20-60 cm. tall, the stems simple or branched, densely glandular-pilose
above; leaves petiolate, usually ovate but may be oblanceolate, spatulate, lyrate-
ovate, obate or pinnate-lobate below, usually coarsely and irregularly serrate,
acuminate, narrowed to the base, thin, sparsely or densely villous or glandular-
villous; flowers on short, densely pilose pedicels; calyx ca. 1 cm. long in maturity,
densely glandular-pilose, prominently nerved, the triangular lobes much shorter
than the tube; corolla purple, glabrous, 1.5-2 cm. long; mature capsule ca. 0.5 cm.
long.
FIG. 14. Bonplandia geminiflora. A, habit, }/% natural size; B, corolla dis-
sected to show stamens and pistil, X 2; C, fruiting calyx, X 3.
87
88 FIELDIANA: BOTANY, VOLUME 24
The leaves are highly variable; some which are pinnate-lobate
with long, almost linear lobes, suggest those of some Cruciferae.
COBAEA Cav.
REFERENCE: Paul C. Standley, A revision of the genus Cobaea,
Contr. U. S. Nat. Herb. 17: 448-458, t. 26-31. 1914.
Herbs or sub-shrubs, probably always annual (fide Standley), scandent by
coiled tendrils, somewhat pubescent; leaves alternate, pinnately compound, sessile
to short-petiolate, the 4-6 leaflets entire, usually short-petiolulate, the terminal
leaflet usually modified as a tendril; flowers axillary, usually solitary, occasionally
2-3 on long peduncles; calyx persistent, herbaceous, the 5 sepals broad to narrow,
usually divided almost to base but may be united for one-third to more than one-
half their length, regular; corolla green, greenish-white, greenish-yellow, yellow,
pale pink or lavender to purplish-green, sometimes striped, campanulate, usually
exceeding the calyx, lobes orbicular to ovate or broadly triangular to acuminate,
sometimes caudate; stamens equally inserted near base of corolla tube, usually
exserted, filaments slender, wooly at point of insertion with pubescence sometimes
extending down onto tube, sometimes pubescent to wooly near apex, anthers ver-
satile; style usually exserted, stigma 3-lobed or branched; ovary superior on a large,
5-lobate disc, glabrous, elliptic-ovoid, tri-carpellate, tri-locular; mature capsule
exceeding calyx, ellipsoid or elliptic-ovoid, coriaceous, dehiscing septicidally, each
locule containing 2 to several large, flat, winged seeds.
Of the 19 species described, six are known from Guatemala. An
American genus of humid, montane forests and thickets, sometimes
persisting in hedges along cleared fields and slopes, it ranges from
Mexico throughout Central America and in the Andes from Vene-
zuela to Colombia and northern Chile.
Cobaea scandens has long been cultivated as an ornamental in
Europe as well as the Americas.
Sepals broadly rounded at apex C. scandens.
Sepals attenuate to acute.
Sepals ovate to broadly ovate C. skutchii.
Sepals oblong-lanceolate to linear-lanceolate.
Corolla lobes acuminate C. lutea.
Corolla lobes short, rounded, ovate-obtuse or broadly triangular.
Corollas 2.5-4 cm. long.
Corollas 2.5 cm. long C. steyermarkii.
Corollas 3-4 cm. long C. triflora.
Corollas 5-6 cm. long C. pachysepala.
FIG. 15. Cobaea lutea. A, habit, J/£ natural size; B, pistil, with disc and calyx,
itural size; C, corolla opened to show stamens, natural size.
89
90 FIELDIANA: BOTANY, VOLUME 24
Cobaea lutea Don, Edinburgh Phil. Journ. 10: 112. 1824.
C. macrostema Pavon ex Don, I.e. as synonym; Hook. Bot. Mag.
66: pi. 3780. 1840. C. acuminata DC. ex Hook. I.e. C. macrostoma
DC. Prodr. 9: 322. 1845. Rosenbergia macrostoma House, Muhlen-
bergia 4: 24. 1908. Campana morada (Guatemala).
Usually in damp thickets but has been collected on dry hillsides,
open, gravelly slopes, in hedges and as a weed in cornfields, 300-
2,666 m.; San Marcos; Quezaltenango; Retalhuleu; Escuintla; Sacate-
pe"quez; Guatemala; Santa Rosa; Jalapa; Jutiapa; Chiquimula. Mex-
ico (Hinton 11671), San Antonio, Dist. Montes de Oca, Gro.
A small or large vine, sometimes densely covering shrubs or small trees, the
typical form glabrous almost throughout; leaflets thin when dry but somewhat
succulent when fresh (fide Standley), essentially glabrous but often villous ciliate
at base, short-petiolulate, oblong to obovate, sometimes pandurate, 3-8 cm. long,
acute to short-acuminate, mucronulate, rounded or subcordate at base, often un-
equal; flowers usually solitary but may be 2 or 3 on peduncles 15-25 cm. long;
sepals divided almost to base, 2-3.5 cm. long, lanceolate or linear-lanceolate, usu-
ally long-attenuate, essentially glabrous but those of the typical form ciliate with
long, scattered, slender hairs; corolla pale green or yellowish-green, rarely tinged
with brown or light purple, 3.5-6 cm. long, the lobes acuminate to long-acuminate,
usually minutely puberulent outside; stamens very long-exserted, purplish; style
long-exserted ; capsule 3-5.5 cm. long.
The specific name is inappropriate as the corolla is nearly always
described by collectors as "pale green"; only one label of the 39 col-
lections studied reads "yellowish-green."
C. lutea f. villosa (Standl.) Gibson, Fieldiana: Botany 31: 354.
1968. C. vittosa Standl. Contr. U. S. Nat. Herb. 17: 454. 1914. Ala
de cucaracha and Granadina (El Salvador).
Differs from C. lutea only by villous nodes; sometimes also villous
or puberulent on petioles.
Although it has not been reported from Guatemala, it may be
expected there, as one collection, Padilla 415, was made in Ahua-
chapan, near the Guatemalan border.
C. lutea f. viorna (Standl.) Gibson. Fieldiana: Botany 31: 354.
1968. C. viorna Standl. Contr. U. S. Nat. Herb. 17: 453. 1914.
Colchito and Rosanillo (Jutiapa).
GIBSON: FLORA OF GUATEMALA 91
Differs from C. lutea only in the absence of long cilia on sepals
and leaf bases.
Gobaea pachysepala Standl. Contr. U. S. Nat. Herb. 17: 456,
t. 29. 1914.
Damp forests and thickets, 1,500-3,000 m. Guatemala; Sacate-
pe"quez; Chimaltenango.
A small or large vine, the stems glabrous except villous at nodes; leaflets essen-
tially glabrous, sometimes villous-ciliate, narrowly oblong to lance-oblong, 5-10
cm. long, 2.3-3 cm. wide, acuminate, mucronate, rounded or sub-cordate at base,
petiolules short-villous; flowers 1-3 on peduncles 12-22 cm. long; sepals divided
almost to base, 1.5-3.8 cm. long, thick and leathery, lanceolate, acuminate, puberu-
lent outside near base, the typical form finely tomentulose only inside along mar-
gins; corolla pale green or yellowish-green, prominently nerved, narrowed rather
abruptly near base, 5.5-6 cm. long, essentially glabrous but sparsely villous near
base, the lobes rounded ovate, obtuse, 12-20 mm. long, glabrous but minutely cili-
ate; stamens about equalling corolla or surpassing it by ca. 1 cm., filaments villous
to wooly, more densely so at base; style surpassing corolla by only 8-12 mm., stig-
mas thick, 2.5-3 mm. long; capsule 5-6.5 cm. long.
G. pachysepala f. tomentulosa (Standl.) Gibson. Fieldiana:
Botany 31: 354. 1968. C. tomentulosa Standl. Contr. U. S. Nat.
Herb. 17: 457. 1914. Flor de campana (Quezaltenango) .
Differs from C. pachysepala only in more dense vesture: the stems
may be obscurely tomentulose to densely villous, the nodes often
densely white-villous, the petiolules short-villous to wooly, the leaf-
lets sparsely pubescent to villous beneath, especially along veins, the
sepals usually densely tomentulose on the outside as well as along
inside margins, and the corollas may have more scattered puberu-
lence on the lobes.
Cobaea scandens Cav. Icon. PL 1: 11, t. 16, 17. 1791; Benth.
in DC. Prodr. 9: 322. 1845; Brand in Engler, Pflanzenr. IV, 250: 24.
1907. Rosenbergia scandens House, Muhlenbergia 3: 23. 1908. Cam-
pana, campana morada (fide Standley), bejuca campana (Dept. Gua-
temala, fide Aguilar) .
Cultivated at various places, 1,500-3,800 m. ; cultivated specimens
seen from Huehuetenango, San Marcos and Guatemala; often per-
sisting in hedges (fide Standley) . Mexico and western South Amer-
ica, Venezuela to northern Chile.
A large vine with only scattered pubescence on stem and petioles; mature leaf-
lets usually 7-8 cm. long, 3-3.5 cm. wide but may attain 10 by 5 cm., glabrous
(rarely minutely ciliate), acuminate, the upper 2 pairs on short-petiolules, elliptic
92 FIELDIANA: BOTANY, VOLUME 24
to obovate, oblique at base, the lower pair almost panduriform, usually auriculate,
sessile or subsessile; flowers solitary on peduncles 15-22 cm. long; calyx broadly
campanulate, outer surface glaucous but minutely short-pubescent on the inside,
the sepals united for more than one-half their length, 2.5-3.5 cm. long, 2-3 cm.
wide, oval or suborbicular, rounded at apex, undulate, mucronate; corolla usually
greenish-lavender or purplish or variously striped but may be green or greenish-
white, purplish inside, 5-6 cm. long, the lobes orbicular to obovate, sometimes with
sparse, minute puberulence on the outside; stamens exserted; style slightly exceed-
ing or about equalling corolla tube; capsule 5-7 cm. long, seated on disc of deeply
bilobate lobes.
Cobaea skutchii I. M. Johnston, Journ. Arn. Arb. 19: 128. 1938.
Wet, mixed forests or thickets, 1,200-2,650 m.; Quezaltenango;
Sacatepe"quez; Escuintla.
A slender vine, almost entirely glabrous but may be sparsely and minutely
villous at nodes; leaflets slender-petiolulate, membranaceous, glabrous, lance-
oblong to oblong-ovate, usually 6-11 cm. long, acuminate, mucronate, truncate
to subcordate at base, often unequal; flowers solitary on peduncles averaging 20 cm.
in length; sepals united for one-fourth to one-third their length, 1.3-1.6 cm. long,
0.8-1.1 cm. wide, glabrous, ovate, acuminate, mucronate; corolla pale green, ca.
2.5 cm. long, the lobes obtuse or rounded, minutely puberulent on the outside;
stamens shortly exserted (usually not more than 0.5 cm.); style exserted by 1-2
cm.; capsule ca. 4 cm. long.
Cobaea steyermarkii Standl. Field Mus. Bot. 22: 271. 1940.
Known only from the type, San Marcos, Loma Corona, 9 miles
northwest of El Porvenir, southwestern slopes of Volcan Tajumulco,
1,300-2,000 m., Steyermark 37759.
A very slender vine, essentially glabrous, the stems sparsely and minutely
villous only at the nodes; leaflets thin, glabrous, slender-petiolulate, oblong-ovate
to lance-oblong, 2.5-6 cm. long, acute to acuminate, mucronate, truncate to sub-
cordate at base, often unequal; flowers solitary on peduncles to 22 cm. long, sepals
divided almost to base, ca. 2 cm. long, glabrous, linear-lanceolate, narrowly long-
acuminate, mucronate; corolla pale green at first, becoming yellow, ca. 2 cm. long,
the lobes broadly triangular, subobtuse, minutely puberulent on the outside; sta-
mens long-exserted (fide Standley); style exserted, capsule 4.5 cm. long.
As this type specimen is a unicate collection, dissection was not
made. The small size of the corolla may be misleading, as the flower
may not have been completely developed. If the stamens were long-
exserted as Standley says, they are now broken and gone. In other
details, this collection is much like C. triflora.
Cobaea triflora Donn.-Smith, Bot. Gaz. 13: 75. 1888. C. macro-
stoma var. triflora Brand, Pflanzenr. IV, 250: 26. 1907. Rosenbergia
triflora House, Muhlenbergia 4: 25. 1908.
GIBSON: FLORA OF GUATEMALA 93
Wet forests or thickets, Alta Verapaz and possibly Baja Verapaz,
900-1,650 m.
A small or large vine, usually glabrous except for some minute pubescence on
stems and petioles; leaflets thin, glabrous, slender-petiolulate, oblong-ovate to
lance-oblong, 2.6-8 cm. long, acuminate, mucronate, somewhat narrowed to the
truncate or subcordate and often oblique base; flowers 1-3 on peduncles 15-22 cm.
long; sepals divided almost to base, 2-3 cm. long, essentially glabrous, sometimes
minutely villous at base and on margins, linear-lanceolate to lanceolate, long-
attenuate; corolla purplish-green, or pale yellow tinged with reddish-purple, 3-4
cm. long, the lobes short, rounded to broadly triangular, minutely puberulent with-
out; stamens shortly exserted, exceeding corolla by not more than 0.5 cm.; style
usually shortly exserted but may surpass corolla by 1-2 cm.; capsule 4-5 cm. long.
Although the specific name alludes to the fact that there are
sometimes three flowers at a node, this character is variable; three
flowers have been observed in the nodes of several collections of
C. lutea and C. pachysepala and may occur in other species. The
corollas of some specimens of C. lutea occasionally display purple or
brownish tinges, but C. triflora is readily separated from C. lutea by
its smaller corollas with short lobes, and the stamens only shortly
exserted. The corolla lobes of C. pachysepala are similar in shape but
the corolla tubes of C. triflora are much smaller, and the filaments of
the stamens are never wooly near the apex as in C. pachysepala.
LOESELIA L.
REFERENCE: Brand in Engler, Pflanzenr. IV, 250: 172-179. 1907.
Annuals or perennials, sometimes low shrubs; leaves sessile to short-petiolate,
mostly alternate, occasionally sub-opposite, entire or dentate; flowers 1-3 in leaf
axils or glomerate at ends of upper branches, subtended by imbricate bracts; calyx
tubular to tubular-campanulate, slightly zygomorphic, the membranous sinuses
hyaline, rupturing in age, the lobes herbaceous; corolla pale yellow, white, lavender
or purple, salverform, bilabiate, the tube slender or gibbous, the lobes equalling
or shorter than the tube; stamens glabrous, inserted at the same height just below
the throat of the corolla, exserted; style exserted; ovary elliptic-ovoid, seated on a
small disc; mature capsule ellipsoid to elliptic-obovoid, glabrous, enclosed in the
accrescent calyx, about equalling the calyx tube, dehiscing loculicidally, each locule
containing 1-several small, narrowly winged seeds.
Of the ten reported species, only three are known from Guatemala.
Bracts of inflorescence broadly ovate to reniform, ciliate with long, stiff bristles;
corolla lobes pale yellow L. ciliata.
Bracts of inflorescence linear or lanceolate, usually dentate but not setose-ciliate;
corolla lobes purple.
Inflorescence of several flowers in usually dense, long pedunculate, headlike
clusters L. glandulosa.
Inflorescence of only 1 or 2 flowers, almost sessile or very short pedicillate.
L. ramosissima.
94 FIELDIANA: BOTANY, VOLUME 24
Loeselia ciliata L. Sp. PI. 628. 1753. Benth. in DC. Prodr. 9:
319. 1845. L. aristata G. Don, Gen. Hist. Dichl. PI. 4: 248. 1838.
L. involucrata G. Don, I.e. L. ciliata var. echinophylla Brand in
Engler, Pflanzenr. IV, 250: 178. 1907. Hoitzia aristata HBK. Nova
Gen. & Sp. 3: 164. 1818. H. loeselia Spreng. Syst. 1: 626. 1825.
H. lupulina Hook. & Arn. Bot. Beech. Voy. Suppl. 441. 1841. Ca-
narios (Guatemala).
Usually in dry thickets or brushy slopes but occasionally in wet
thickets, 75-1,800 m.; Baja Verapaz; Chimaltenango; Chiquimula;
Escuintla; Guatemala; Jutiapa; Retalhuleu; Santa Rosa; Sacatepe*-
quez; Zacapa. Mexico to Panama.
Annuals, usually much branched, erect (rarely prostrate), 0.45-1.2 m. tall
(rarely to 2 m.) the stems and branches slender and brittle, usually puberulent but
may be glabrate, especially in age; leaves short-petiolate, oblong-ovate to broadly
ovate, dentate, acute, truncate to attenuate at base, puberulous or glabrate; in-
florescences usually long pedunculate, with few to many flowers, congested and
bracteate, each flower subtended by a pair of papery, awned, glume-like bracteoles
in addition to the 3 or more large, green bracts which are broadly ovate to reni-
form, sessile and clasping, and coarsely dentate, each tooth ending in a long, stiff,
pale bristle; calyx lobes narrow, scarious and dentate, the terminal tooth usually
prolonged; corolla ca. 1 cm. long, pale yellow, or the tube white with straw-colored
lobes, the margins of the lobes usually minutely tomentose; mature capsule ca.
4 mm. long, each locule usually containing 1 (rarely 2) narrowly winged seed.
Loeselia glandulosa (Cav.) G. Don, Hist. Dichl. PI. 4: 248. 1837.
L. glandulosa (Cav.) G. Don ssp. cervantesii (HBK.) Brand and var.
nepetifolia (Cham. & Schlecht.) Brand in Engler, Pflanzenr. IV, 250:
176-177. 1907. L. glandulosa (Cav.) G. Don ssp. conglomerata (HBK.)
Brand and varieties hirsuta Brand and scabra (Mart. & Gal.) Brand,
I.e. L. nepetifolia G. Don, Gen. Syst. Dichl. PI. 4: 248. 1837. L. con-
glomerata G. Don, I.e. L. scabra Walpers, Rep. Bot. 6: 527. 1846^47.
Hoitzia glandulosa Cav. Icon. 4: 45, t. 367. 1797. H. cervantesii HBK.
Nova Gen. & Sp. 3: 128. 1819. H. conglomerata HBK. I.e. H. capi-
tata Willd. in Roem. & Schult. Syst. Veg. 4: 370. 1819. H. spicata
Willd. I.e. H. nepetifolia Cham. & Schlecht. Linnaea 6: 385. 1831.
H. elata Hook. & Arn. Bot. Beech. Voy. 441. 1841. H. scabra Mart.
& Gal. Bull. Acad. Brux. 12, 2: 274. 1845. Clarincillo silvestre (Hue-
huetenango) ; salvia virgen (Escuintla) ; orozuz amargo, salvia mo-
rada (Dept. Guatemala, fide Aguilar); Verbena (Chiapas, Mexico,
fide Standley).
Usually in dry places, rocky, bushy hillsides and slopes, along
roadsides, in pine-oak forests; occasionally in Cupressus groves and
GIBSON: FLORA OF GUATEMALA 95
damp thickets, 350-2,950 meters; Alta Verapaz; Baja Verapaz; Chi-
maltenango; Chiquimula; Escuintla; Guatemala; Huehuetenango;
Jalapa; Jutiapa; El Quiche"; Sacatepe"quez; Santa Rosa; Solola;
Zacapa. A heterogeneous species extending from southern Arizona
through Mexico and Central America; Colombia and Venezuela.
Annuals, but often suffrutescent, usually 40-80 cm. tall, occasionally only
15-20 cm. tall, the stems often much branched, slender, brittle, usually glandular-
puberulent, occasionally glabrate, rarely eglandular; leaves short-petiolate, lanceo-
late to broadly ovate, acute to acuminate, sharply serrate, contracted at base and
decurrent, usually pubescent but may be glabrate; flowers sessile or subsessile,
usually in dense, headlike clusters, these few to many-flowered, on short or much
elongated peduncles, the bracts linear to lanceolate, appressed, usually coarsely
dentate, often glandular; calyx lobes narrow, dentate, aristate, usually puberulent;
corolla pink, blue, lavender or purple, 1-2 cm. long, glabrous, the lobes about
equalling the tube; mature capsule 2-3.5 mm. long, each locule containing 3 flat,
narrowly winged seeds; the seeds very sticky when wetted.
Loeselia glandulosa sens. lat. is a most interesting genetic complex.
For instance, it was possible to readily separate all of the material in
the species into two groups —those with corollas only 8-10 mm. long,
and those with corollas 15-20 mm. long. It was also observed that
the shorter corollas are often sharply constricted above the base of
the tube, and that the peduncles of plants with large corollas tend to
be longer and considerably farther apart on the stems than those with
short corollas. However, none of these characters is constant; plants
with small corollas sometimes have internodal distances of 5-8 cm.,
very short peduncles, and the corolla tubes are not always constricted
at the base. Leaf shape and pubescence also vary, occasionally an
eglandular form is seen, and a few plants, all from Huehuetenango
(Skutch 1930, Standley 65651 and 82092 and Van Hook 1), were ob-
served to have bracts which, although short ciliate and strigose, lack
dentation or serration. All of the varieties separated by Brand ap-
pear to be only variously pubescent forms, with the single exception
of his var. ramosissima. This is Loeselia ramosissima (Mart. & Gal.)
Walpers, incorrectly reduced by Brand to varietal status.
Loeselia ramosissima (Mart. & Gal.) Walp. Repert. Bot. 6:
526. 1846-47. L. glandulosa var. ramosissima Brand in Engler, Pflan-
zenr. IV, 250: 177. 1907. Hoitzia ramosissima Mart. & Gal. Bull.
Acad. Brux. 12(2): 273. 1845.
Usually in dry places, along roads in oak or oak-pine forests or
on open slopes, occasionally a weed in cornfields, 1,600-2,800 m.;
Huehuetenango; Quezaltenango (fide Standley) ; El Quiche". Western
and southern Mexico.
96 FIELDIANA: BOTANY, VOLUME 24
Annuals, usually branched from the base, erect, low, 8-30 cm. tall, the stems
glandular-pilose; leaves short-petiolate, lanceolate or ovate, acute, serrate, usually
pubescent (rarely glabrate), the lower leaves soon withering; flowers solitary or
geminate, short-pedicillate or almost sessile, the bracts lanceolate or linear-lanceo-
late, dentate, unequally trilobate (the two opposite lateral lobes in no way equal-
ling the terminal lobe); calyx lobes narrow, scarious, usually dentate; corolla violet
or purple, 7-10 mm. long, glabrous, the lobes about equalling the tube; mature
capsule 2-3 mm. long, each locule containing only one plump, broadly winged seed.
Although reduced by Brand to varietal status under L. glandu-
losa, this species, well represented by numerous collections from Mex-
ico and Guatemala, may be separated at a glance from L. glandulosa
by the quite different appearance of the long, narrow, flowering
branches. Further, the obvious enlargment of the two lateral lobes
of the bracts does not occur in L. glandulosa, and L. ramosissima pro-
duces only one broadly winged seed in each locule, whereas three nar-
rowly winged seeds are usually found in each locule of L. glandulosa.
LENNOAECAE
DOROTHY N. GIBSON
REFERENCES: 0. Drude in Engler & Prantl, Pflanzenf. IV(1) : 12-
15. 1891. P. A. Rydberg, Lennoaceae, N. Amer. Fl. 29: 19-20. 1914.
Small parasitic herbs, without chlorophyll, the stems fleshy, the leaves reduced
to small scales; inflorescences cymose- paniculate, cymose-thyrsoid, spicate, or with
flowers arranged on a peltate disk; flowers small, perfect; sepals 5-10, linear, dis-
tinct or nearly so; corolla gamopetalous, tubular or narrowly funnelform or salver-
form, 5-8-lobate; stamens 5-10, usually the same number as and alternate with the
corolla lobes, the filaments almost entirely adnate to the corolla; anthers 2-celled,
introrse, dehiscent by longitudinal slits; gynoecium of 6-14 wholly united carpels,
the ovary divided by false dorsal partitions into twice as many cells, these uniovu-
late; style simple, the stigma capitate or peltate, crenate; ovules anatropous,
almost horizontal; fruit drupaceous-capsular, finally irregularly circumscissile,
containing 12-28 one-seeded nutlets; seeds with endosperm, the embryo globose.
Three genera, confined to the extreme southwestern United States,
western and southern Mexico, and Guatemala. Only one genus is
known to be represented south of Mexico.
LENNOA Lexarza
Low, succulent parasitic herbs, glabrous or somewhat pubescent, the stems
simple or branched, bearing numerous more or less dentate bracts; inflorescence
cymose-thyrsoid, the flowers densely crowded, bracteate; sepals 5-8, distinct al-
most to the base, pubescent or puberulent, linear; corolla funnelform to salverform,
the limb plicate, the 8 lobes emarginate, the tube ventricose at the base, persistent;
stamens 8, in 2 whorls, included, the linear filaments adnate to the corolla for most
of their length, anther cells ellipsoid, coherent at the apex, divergent below; ovary
8-carpellate, with 16 locules in a ring, the 22 ovules horizontal, anatropous; fruit
somewhat drupaceous.
FIG. 16. Loeselia ramosissima. A, habit, natural size; B, flower with bracts,
X 5; C, corolla opened to show stamens and pistil, X 5.
97
98
FIELDIANA: BOTANY, VOLUME 24
FIG. 17. Lennoa madrepor aides. A, habit of plant, natural size; B, corolla
opened to show stamens, X 5; C, flower, X 5; D, pistil, X 10.
Two species or perhaps one species with two varieties, one con-
fined to Mexico, one extending southward.
Lennoa madreporoides Lexarza in La Llave & Lex. Nov. Veg.
Descr. 1: 13. 1824. L. madrepoides Steud. Norn. Bot. ed. 2. 2: 22.
1841. L. madreporoides subsp. Australia Steyerm. Acta Bot. Venez.
3: 230. 1969.
Dry banks, especially in cinders, 300 meters or less; Zacapa;
El Progreso. Mexico. Reported from Colombia.
GIBSON: FLORA OF GUATEMALA 99
Plants somewhat succulent, yellowish or brownish, the thick stems 5-17 cm.
high, simple up to the inflorescence or sometimes branched below; leaf scales ellip-
tic or lanceolate, acute, dentate; flowers sessile or subsessile, very numerous and
crowded, forming a broad dense inflorescence; sepals 8, linear, hispidulous-puberu-
lent, shorter than the corolla; corolla pale violet to bluish white, sometimes yellow-
ish within throat, funnelform to salverform, minutely puberulent outside, some-
what papillose within, 5-8 mm. long, the limb 3-6 mm. broad, sinuate-lobate.
This curious plant has been collected but twice in Guatemala,
and may be rare. In both cases it was found growing on railway
embankments, among cinders, most of the stems being buried and
only the upper branches with their small flowers appearing above
the surface of the ground.
HYDROPHYLLACEAE. Waterleaf Family
DOROTHY N. GIBSON
REFERENCES: Choisy, Me'm. Soc. Phys. Gen. 6: 106. 1833 and in
DC. Prodr. 10: 179. 1846. A. DC., Prodr. 9: 287 and 564. 1845.
Benth. & Hook., Gen. PI. 2: 825. 1876. Peter in Engl. & Prantl,
Pflanzenf. IV(3a): 54-71. 1892. Brand in Engler, Pflanzenr. IV
251: 1-210. 1913.
Annual or perennial herbs or suffruticose plants, often pubescent, scabrid,
glandular, and sometimes armed; leaves alternate or opposite, sometimes rosulate,
entire or pinnately (rarely palmately) divided; inflorescences usually cymose,
often helicoid, or may be reduced to a single flower; flowers bisexual, actinomor-
phic; calyx 5-lobed, imbricate; corolla 5-lobed, imbricate (rarely contorted), rotate,
campanulate, or funnelform; stamens usually 5, inserted on corolla tube near its
base, alternate with corolla lobes, anthers bilocular, dehiscing longitudinally, fila-
ments often basally dilated or subtended by appendages; ovary superior to half-
inferior, usually unilocular with 2 parietal placentae meeting in the center, style 1
or 2 (rarely 3 or 5), ovules 4 to many; fruit of 2 carpels, each 2 to many-seeded
(occasionally 1-seeded due to abortion), dehiscing loculicidally, septicidally or
rarely indehiscent; seeds variously rugose, reticulate or muricate.
The Hydrophyllaceae are of almost world-wide distribution, hav-
ing been reported from all continents except possibly Australia.
Most of the 20 genera are highly variable; four of them occur in
Guatemala.
)vary bilocular Hydrolea.
Dvary unilocular.
Style 1 . .Phacelia.
Styles 2.
Plant seldom more than 30 cm. tall; herbaceous to partly woody Nama.
Plant quite large, to several meters tall; suffruticose to arborescent.
Wigandia.
100 FIELDIANA: BOTANY, VOLUME 24
HYDROLEA L.
REFERENCE: Brand in Engler, Pflanzenr. IV, 251: 174-185. 1913.
Annuals or perennials from taproots, usually erect, sometimes decumbent,
herbaceous to partly woody, usually glandular-pubescent, sometimes spiny in leaf-
axils; leaves alternate, entire, elliptical-lanceolate to obovate; flowers usually few
in lateral and/or terminal cymes; calyx divided to base with lobes overlapping at
base, persistent, accrescent; corolla usually blue, sometimes white, broadly cam-
panulate, usually exceeding the calyx; stamens included, inserted on tube, partly
adnate at base, filaments dilated at base, anthers sagittate; styles 2 (rarely 3 or 5),
elongating, with clavate-capitate stigmas; ovary superior, globose to nearly elliptic,
bilocular, with two fleshy placentae containing numerous ovules; capsule globose
to nearly elliptic, dehiscing irregularly, containing numerous, minute seeds.
A variable genus of several species, ranging from the southern
United States through Mexico, Central and South America, the
East and West Indies, and in Asia and Africa, it is represented in
Guatemala by only one species.
Hydrolea spinosa L. Sp. PI. ed. 2: 328. 1762; Aubl. Hist. PL
Guian. 1: 281. 1775; HBK. Nova Gen. & Sp. 3: 98. 1818; Edwards'
Bot. Reg. 7: t. 566. 1821; Choisy in M<§m. Soc. Phys. Geneve 6: 110.
1833 and in DC. Prodr. 10: 181. 1846; Brand in Engler, Pflanzenr. IV,
251: 180. 1913. H. cervantesii Brand and var. maior Brand, I.e. 184.
H. trigyna Swartz, Nov. Gen. & Sp. Ind. Occ. 54. 1788. Sagonea
palustris Aubl. Hist. PL Guiana 1: 285. 1775. Campanula americana
hirsuta Houst. ex Swartz, Nov. Gen. & Sp. Ind. Occ. 54. 1788. Reich- \
elia guianensis Spreng. Syst. 1: 940. 1825. R. palustris Billberg ex
Beurl. in Vet. Acad. Handl. Stockholm 141. 1856. Espina de agua
(Dept. Guatemala).
In marshes, wet thickets, meadows, and in sand along streams,
rivers and lakes, 45-1,600 m.; Baja Verapaz; Chimaltenango; Chiqui- j
mula; Escuintla; Guatemala; Isabal; Jalapa; Jutiapa; Pete"n; El Pro-
greso; El Quiche"; Retalhuleu; San Marcos; Santa Rosa; Saca-
tepe"quez; Zacapa. Southwestern United States through Mexico to
Panama.
Annuals or perennials, usually erect, simple or much branched, 15-130 cm.j
tall, the stems and branches pilose and glandular; leaves subsessile to short petioJ
late, variable, usually elliptic to elliptic-ovate, occasionally narrowly elliptic, rarely
obovate, averaging 2-12 cm. long; flowers usually short-pedunculate, few in axill
lary and/or terminal cymes, often congested; sepals linear-lanceolate to lanceolate,
acute, 5-7 mm. long, glandular-pilose; corolla open-campanulate, pale blue to pur-
plish, 7-12 mm. across, 5-12 mm. long; stamens less than or surpassing or about I
equalling the corolla, adnate near base of tube, filaments dilated at base; ovarji
ovoid to globose, largely glabrous but minutely glandular near top; styles glandu-
lar at base, elongating; mature capsule 4-5 mm. long, ovoid to globose; seeds
numerous, minute, light brown, longitudinally ridged.
C JTV8' Hyfrolea;sPi^sa. A, habit, Y2 natural size; B, fruiting calyx x 3-
C, corolla opened to show stamens, X 2.
101
102 FIELDIANA: BOTANY, VOLUME 24
Brand, in his description of H. cervantesii, stated, "Species basi
staminum a ceteris speciebus generis valde diversa." However, sev-
eral dissections of material cited by him revealed no appreciable or
constant differences.
NAMA L.
REFERENCES: Brand in Engler, Pflanzenr. IV, 251: 142. 1913;
Hitchcock, A Taxonomic Study of the Genus Nama, I and II, Am.
Journ. Bot. 20: 415-430 and 518-534. 1933.
Erect to prostrate, simple or branched, herbaceous to partly woody, pubescent,
often glandular annuals or perennials from slender taproots; leaves mostly alter-
nate and entire, rarely dentate; flowers borne singly or in pairs in axils of upper
leaves or several in reduced lateral or terminal cymes; calyx divided nearly to base,
the lobes linear-lanceolate to linear-spatulate, subequal, accrescent; corolla whit*
to purple, sometimes partly puberulent, tubular to funnelform, exceeding the calyx;
stamens usually included, all basally adnate and unequally inserted on the corolla
tube or may be unequal in length, filaments usually glabrous, their bases dilate
or sometimes appendaged; styles 2, elongating, usually free but sometimes partially
united with small capitate stigmas; ovary superior to semi-inferior, usually s«
what puberulent, unilocular but appearing bilocular due to the ingrowing parietal
placentae; mature capsule usually ovoid to oblong but may be globose, cartilagi-
nous, dehiscing loculicidally or septicidally, containing numerous brown, varu
pitted, reticulate, or smooth seeds.
A variable genus ranging from the western United States through
Mexico, Central America, the West Indies, western South America
and the Hawaiian Islands.
Plants erect.
Leaves linear-spatulate to oblong-elliptic . . • - N. dichotomum.
Leaves oblong-lanceolate, obovate, or nearly ovate.
Calyx lobes broadly spatulate,
Calyx lobes linear to linear-spatulate,
Plants ascending, trailing, or prostrate.
Plants ascending or trailing; leaves obovate to elliptic-lanceolate or el lip tie,
petioles not winged ................. • •«• hirsu
Plants prostrate; leaves broadly ovate to spatulate, narrowed towuijed
petioles ......
Nama dichotomum (R. & P.) Choisy, M^m. Soc. Phys. Geneve
6: 113. 1833; Gray, Proc. Am. Acad. 5: 338. 1861; Brand in Engler,
Pflanzenr. IV, 251: 150. 1913. Hydrolea dichotoma Ruiz & Pav. Fl.
Peruv. 3 : 22, t. 2U, fig- &• 1802. N. tetrandra Pa von ex Choisy, Me"m.
Soc. Phys. Geneve 6: 113. 1833, in synon. N. stricta Phil. Fl. Atac.
37. 1860. Marilaunidium strictum (Phil.) Kuntze, Rev. Gen. PI. 2,
GIBSON: FLORA OF GUATEMALA
103
434. 1891. A7. dichotomum, subsp. eu-dichofomum Brand in Eneler-
Pflanzenr. IV, 251: 151. 1913 (in part). N. dichotomum subsp eu-
dichotomum f . stricta (Phil.) Brand, I.e. 151, fig. 28. N dichotomum
subsp. eu-dichotomum f. amplifolia Brand, I.e. N. dichotomum var
angustifolium Gray, Proc. Am. Acad. 8: 284. 1870. N. angustifolium
(Gray) A. Nels., Coult. and Nels. New Man. Rocky Mt. Bot 410
909. N. dichotomum subsp. angustifolium (Gray) Brand in Engler
Pflanzenr. IV, 251: 151. 1913. N. dichotomum ft pauciflora Choisy
m f « ProcoAm. Acad. 8: 284. 1870. Conanthus angustifolius
(Gray) Heller, Bull. Torr. Bot. Club 24: 479. 1897. Marilaunidium
dichotomum (R. & P.) Kuntze, Rev. Gen. PI. 2: 434. 1891. Marilau-
mdium tenue Woot. & Standl. Contr. U. S. Nat. Herb. 16: 162 1913
N. dichotomum var. amplifolium (Brand) Hitchc. Am. Journ. Bot.
£0 : 5oO. 19o3.
A common weed in gardens, fields and pastures, on open hillsides,
sandy stream banks and in open oak forests, 1,600-4,000 m. ; Chimal-
tenango; Guatemala; Huehuetenango; Quezaltenango; El' Quiche*-
Sacatape"quez. Mexico.
Annual, slender, erect, simple or more or less dichotomously branched herbs
4-30 cm. tall, hirsute and glandular; leaves linear-spatulate to oblong-elliptic 4-
10 mm. long, 1-10 mm. wide, tapering to short petioles; flowers usually in pair's in
leaf axils, sessile or on short pedicels; calyx lobes linear-spatulate, 2-5 mm in
flower, to 10 mm. in fruit, hirsute and ciliate; corolla white to pale blue or lavender
>r white with blue or lavender lines, narrowly campanulate, to 5 mm. long; stamens
unequally placed about 1 mm. from base of corolla, the glabrous filaments widened
slightly just above the adnate base and with free margins along the adnate portions-
styles to 2 mm. long; capsules more or less puberulent, ovoid to oblong, containing
JO-60 brown, minutely reticulate seeds.
N. dichotomum var. chasmogamum Brand in Engler Pflan-
zenr. IV, 251: 151. 1913.
More robust and more profusely branched than N. dichotomum;
leaves usually larger and broader, to 40 mm. long and 15 mm. wide,
oblong-lanceolate to nearly ovate; flowers more numerous, 1-3 at
nodes and in axillary and terminal lax cymes, pedicels longer, to 6
mm.; cilia of calyx lobes usually longer; styles often longer, to 6 mm.
N. dichotomum var. pueblense (Rob. & Greenm.) Macbride
Contr. Gray Herb. 49: 45. 1917. N. pueblense Rob. & Greenm Proc'
Am. Acad. 32: 39. 1896.
Leaves oblong-ovate to obovate; calyx and leaves greyish-hirsute.
The only collection seen is from Huehuetenango, Williams, Mo-
lina & Williams 22352, apparently a new record for Guatemala.
FIG. 19. Nama dichotomum var. chasmogamum. A, habit of plant, Jxj natural
size; B, corolla opened to show stamens and pistil, X 5; C, capsule with calyx, X 5.
104
GIBSON: FLORA OF GUATEMALA 105
Nama hirsutum Mart. & Gal. Bull. Acad. Sci. Brux. 12(2): 277
1845; Gray, Proc. Am. Acad. 5: 339. 1861; Brand in Engler, Pflan-
zenr. IV, 251 : 148. 1913. Marilaunidium hirsutum (Mart. & Gal )
Kuntze, Rev. Gen. PI. 2: 434. 1891.
Open slopes, 1,800-3,200 m.; Chimaltenango; Quezaltenango; So-
lola and Totonicapan. Mexico.
A slender perennial, hispid to strigose-hirsute, ascending or with trailing some-
what woody older branches, 8-60 cm. long; leaves obovate to elliptic-lanceolate or
elliptic, 0.8-3 cm. long, 0.5-1.2 cm. broad, with short petioles on older branches
and extreme forms, but may attain 8 cm. by 3 cm. on new shoots and stems with
petioles 1-2 cm. long; flowers in pairs or 3-10 in lax cymes; calyx lobes narrowly
oblanceolate-spatulate to spatulate, 3-6 mm. long in flower, 4-8 mm. in fruit, hir-
sute and ciliate; corolla white to pale lilac or white veined with purple, funnel-
shaped, 6-11 mm. long; stamens unequally placed 1-2 mm. from base of corolla
free filaments flattened, much widened directly above point of insertion and
throughout adnate bases, except with margins of adnate bases free; styles 2-5 mm
long; mature capsules more or less puberulent, especially near base of styles, usu-
ally ovoid but occasionally nearly globose, containing less than 20 (usually 4-12)
granular, minutely alveolate, brown seeds.
It should be noted that all of the Guatemalan collections studied
which had been previously identified as N. prostratum Brand (John-
ston 1915, Skutch 597, Standley 80138a, Steyermark 47212, Williams
et al. 22571 and 23029) are now determined to be N. hirsutum, despite
minor differences. Although the leaves are smaller and all the styles
(with the single exception of Skutch 597) are about 2 mm. shorter
than those of the typical N. hirsutum, several dissections disclosed
the same number of seeds (never more than 12) and showed the na-
ture of the adnate portions of the filaments to match exactly those
of N. hirsutum. Brand's description of N. prostratum leaves much
to be desired, and as I was unable to examine the type fragment,
Ehrenberg 326 (P) from Mexico, my knowledge of this species is lim-
ited to Hitchcock's description and his drawing of the opened corolla
(Am. Journ. Bot. 20: t. 27, fig. 28, 1933). Our specimens in no way
resemble that drawing. The creation of a new variety does not seem
justifiable at this point.
Nama jamaicense L. Syst. Nat. ed. 10, 2: 950. 1759; Choisy in
DC. Prodr. 10: 182. 1846; Gray in Hemsl. Biol. Cent. Am. Bot. 2:
565. 1882; Brand in Engler, Pflanzenr. IV, 251 : 155. 1913. N. jamai-
tense var. gracile Brand, I.e. 156. N. redinata villosa P. Browne,
^iv. & Nat. Hist. Jam. 185, t. 18, fig. 2, 1756, fide Brand. Hydrolea
amaicense Roenschel, Norn. ed. 3, 76. 1797. H. decurrens Moc. ex
"hoisy in DC. Prodr. 10: 181. 1846. Conanthus jamaicensis (L.)
106 FIELDIANA: BOTANY, VOLUME 24
Heller, Cat. N. Am. PI. 6. 1898. Marilaunidium jamaicense (L.)
Kuntze, Rev. Gen. PL 2: 434, t. 27, fig. 19. 1891. Colchita (Jutiapa) ;
paletoria and parietaria (Huehuetenango) fide Steyermark.
A common weed of fields, clearings, dry hillsides, wet banks and
sand-bars, 85-2,000 m.; Alta Verapaz; Chimaltenango; Guatemala;
Huehuetenango; Isabal; Jalapa; Jutiapa; Pete*n; Quezaltenango; Sa-
catepe"quez; Santa Rosa. Mexico, West Indies, South America.
A slender annual, prostrate, strigose-hirsute, with branches 10-50 cm. long;
leaves broadly ovate to spatulate or broadly obovate-spatulate, 1-8 cm. long,
0.5-3.5 cm. broad, narrowed to distinct, winged petioles a fifth as long as the
blades; flowers few, solitary or in 2's, 3's or 5's in leaf axils, nearly sessile or on
pedicels to 1.5 cm. long; calyx lobes usually linear and acute, but may be narrowly
spatulate, from ca. 5 mm. long in flower to 12 mm. long in fruit, often clasping the
ovary rather tightly; corolla white, sometimes tinged with lavender, almost tubu-
lar, 6-7 mm. long; stamens unequally inserted on tube 1-2 mm. from base of
corolla, filaments terete above, but flattened and greatly expanded, ca. 1 mm.
above point of insertion, the adnate portion expanded, with free margins; styles
1.5-2 mm. long, usually distinct but sometimes united J^ to J^ their length; cap-
sule usually puberulent near base of styles, elongated, containing 50-70 light brown,
minutely alveolate seeds.
Nama jamaicense closely resembles and intergrades with N. bi-
florum Choisy of Mexico. However, I agree with Hitchcock that
both species should be maintained, as N. biflorum exhibits several
differing characters which appear to be constant, i.e., smaller leaves,
larger corollas (to 11 mm. long), and the flowers are borne on longer
pedicels (1.5-4 cm.).
PHACELIA Juss.
REFERENCES: A. Brand in Engler, Pflanzenr. IV, 251: 60-130.
1913; Lincoln Constance, A Revision of Phacelia subgenus Cosman-
thus, Contr. Gray Herb. 168: 16. 1949.
Herbaceous, branching, erect to decumbent annuals or perennials from tap-
roots or creeping rootstocks, usually pubescent and often glandular; cauline leaves
usually alternate, entire, lobed, pinnatifid or pinnately dissected; basal leaves
often rosulate, flowers few to numerous in usually helicoid, raceme-like terminal
cymes, long pedunculate to sessile; corolla blue, purple, pale lilac or white, campan-
ulate; calyx divided nearly to base, persistent, often accrescent; stamens usually
exserted, all adnate to and equidistant from the base of the corolla tube, a pair of
scales or glands adnate to tube sometimes present at base of each filament; style
bifid; ovary unilocular with 2 linear placentae adherent to walls; capsule ovoid to
globose, loculicidally dehiscent, containing 2 to many, minutely rugose-aerolate
seeds.
FIG. 20. Phacelia platycarpa. A, habit of plant, ^ natural size; B, flower, X 5.
107
108 FIELDIANA: BOTANY, VOLUME 24
A variable American genus, widely distributed, especially through-
out the western United States, it is well represented in Mexico and
in the Andes, but only one species is known from Guatemala and
none has been reported from other Central American countries. The
subgenus Cosmanthus, to which our single species belongs, contains
14 species; ours is apparently limited to the highlands of Mexico and
Guatemala.
Phacelia platycarpa (Cav.) Spreng. Syst. 1: 584. 1825. Con-
volvulus platycarpos Cav. Icon. 5: 155, t. ^82. 1799. Polemonium
pimpinelloides Willd. ex Roem. & Schult. Syst. 4: 793. 1819. P. achil-
laeifolium Willd. I.e. Eutoca pimpinelloides Spreng. Syst. 1: 569.
1825. E. mexicana Benth., Trans. Linn. Soc. 17: 277. 1834. E. an-
drieuxii A. DC. Prodr. 9: 294. 1845. E. gracilis Mart. & Gal. Bull.
Acad. Brux. 12(2) : 276. 1845. E. ortgiesiana Heer ex Regel, Gartenfl.
10: 309, t. 337. 1861. Cosmanthus mexicanus A. DC. Prodr. 9: 297.
1845. Nemophila ortgiesiana Roezl ex Regel, Gartenfl. 10: 309. 1861.
Phacelia pimpinelloides A. Gray, Proc. Am. Acad. 10: 321. 1875.
P. pubescens Peter in Engler & Prantl, Pflanzenf. 4, 3a: 64. 1893.
P. patuliflora var. mexicana Brand in Engler, Pflanzenr. IV, 251: 66.
1913.
In dry or moist meadows and thickets, and along roadsides, corn-
fields and rivers, 1,500-3,700 m.; Huehuetenango; Quezaltenango; So-
lola; Chimaletenango. A variable species, ranging throughout
montane Mexico to Guatemala.
Perennial from a stout taproot, 5-40 cm. tall, the branches decumbent to
weakly ascending from a basal rosette, pubescent; basal leaves petiolate, linear to
oblanceolate or oblong, 3-30 cm. long, 0.8-6 cm. broad, pinnate with 4-8 pairs of
oblong to obovate, entire to pinnatified leaflets, or the terminal one confluent;
cauline leaves like the basal but reduced upward, toothed or pinnate, usually
somewhat strigulose to hirsute on both surfaces but especially so underneath along
mid vein; inflorescence a loose cyme of few to many flowers on pedicels 5-30 mm.
long; calyx lobes linear-lanceolate to ovate-lanceolate, 3-6 mm. long, unequal,
acute, more or less puberulent; corolla pink to pale blue or white tinged with lav-
ender, rotate-campanulate, 6-10 mm. long, 7-15 mm. broad, the lobes rounded,
somewhat pilose on the back, puberulent gland flaps free at tips; stamens about
equalling or less than the corolla, filaments villous at base and for at least one-half
their length; ovary at least partly hirsute, containing 4-10 ovules on each placenta,
style hirsute on lower half; mature capsule globose, 3-6 mm. in diameter, contain-
ing 9-15 brown, trigonate, rugose-aerolate seeds.
WIGANDIA HBK.
REFERENCE: Brand in Engler, Pflanzenr. IV, 251: 134. 1913.
GIBSON: FLORA OF GUATEMALA 109
Large, suffruticose to arborescent perennials, variously pubescent (including
stinging hairs) and glandular throughout; leaves alternate, often quite large but
variable in size from 5-60 cm. long, 3-42 cm. broad, coriaceous, ovate, obtuse to
sub-acute at apex, cordate to truncate at base, irregularly twice-dentate; flowers
few to many, sessile to sub-sessile, often appearing densely spicate, in scorpioid,
terminal cymes or panicles; calyx persistent, divided nearly to base, the lobes sub-
equal, linear-lanceolate to lanceolate, acute, accrescent, hispid-hirsute to tomen-
tose and ciliate; corolla greenish-white, yellow, or lavender to purple, usually
exeeding the calyx, often partly pilose to tomentose without, the lobes rounded;
stamens exserted, partly adnate to the base of the corolla tube, filaments at least
partly pubescent, anthers linear-oblong, dorsifixed; styles 2, elongated, with capi-
tate-clavate, depressed stigmas; ovary superior, oblong, pubescent, unilocular, with
2 laminiform placentae, the transverse portions of each meeting in the center,
with numerous ovules borne chiefly along their revolute edges; mature capsule
ovate-oblong, pubescent, dehiscing loculicidally or septicidally, and containing
numerous minute, reticulate-rugose, brown seeds.
A widely distributed, weedy, variable genus, ranging from Mexico
throughout Central America, to Argentina.
Plants setose-hispid, at least on stems, petioles, and midveins of leaves . . W. urens.
Plants without setae on stems, petioles, or leaves W. urens var. caracasana.
Wigandia urens (R. & P.) HBK. Nova Gen. & Sp. 3 : 127. 1819;
Choisy in DC. Prodr. 10: 184. 1846, incorrectly as "W. urens (R. &
P.) Choisy." W. kunthii Choisy, Mern. Soc. Phys. Geneve 6: 116.
11833 and in DC. Prodr. 10: 184. 1846. W. peruviana W. Miller,
[Bailey Cyclop. Am. Hort. 1975. 1902. W. urens var. eu-kunthii
Brand, Pflanzenr. IV, 251: 137. 1913. Hydrolea urens Ruiz & Pav.
|F1. Peruv. 3: 21. t. 2^3. 1802. Ernstamra urens (R. & P.) 0. Ktze.
|Rev. Gen. PI. 2: 434. 1891. Chocon (Sacatepe"quez) ; la Aurora (Gua-
jtemala); chichicaste (Guatemala, Honduras, El Salvador); quema-
\dora (Mexico); ortiga (Mexico and Costa Rica).
Frequent along roadsides, in thickets, damp meadows or sandy
(fields, dry slopes, oak and oak-pine forests, and in waste places, 70-
i3,000 m.; Huehuetenango; San Marcos; Quezaltenango; Sacatepe"-
i|quez; Chimaltenango; Guatemala; Alta Verapaz; Santa Rosa. Mex-
ico to Panama and in Peru and Ecuador.
Suffruticose plants, occasionally becoming small trees to 5 m. tall, usually
iidensely sordid with hispid to bristly-hirsute, sometimes glandular-viscid, and often
istinging hairs; leaves alternate, 5-60 cm. long, 3-42 cm. wide, ovate, obtuse to sub-
lacute at apex, cordate to truncate at base, irregularly biserrate, appressed pilose
'on upper surface and usually white-tomentose underneath, with bristles at least on
imidvein and petiole; flowers sessile to subsessile in dense, scorpioid terminal cymes
or panicles; calyx persistent, the lobes subequal, acutely linear-lanceolate, setose-
hispid to tomentose and hispid-ciliate, about equalling or slightly surpassing the
FIG. 21. Wigandia urens. A, flowering branch, H natural size; B, pistil, with
calyx, X 3; C, corolla opened to show stamens, X 2.
110
GIBSON: FLORA OF GUATEMALA 111
mature capsule; corolla pale violet to purple, lighter in throat, silky pilose to to-
mentose without, to 20 mm. long, equalling or slightly exceeding the calyx ; stamens
included, subequal, filaments retrorsely hispid from base to half or nearly two-
thirds their length, basally adnate to corolla tube; styles at an thesis 6-10 mm.
long, often becoming 10-15 mm. long in maturity, pilose for more than one-third
their length; stigmas clavate-capitate; ovary ovoid-oblong, silky to hispid-hirsute;
mature capsule oblong-conical, partly hoary to hispid-hirsute; seeds numerous,
minute, reticulate-rugose, brown.
A widepsread and variable species, especially in vesture.
One collection from Panama, Davidson 972, has leaves with the
soft appearance of var. caracasana, but with stems and calyces cov-
ered with long, soft, shaggy hairs and only a few scattered hard
bristles. This type of vesture was also seen on peduncles and calyces
of a few specimens of the var. caracasana from Mexico and Costa Rica.
W. urens var. caracasana (HBK.) Gibson. Fieldiana: Botany
31:353. 1968.
W. caracasana HBK. Nova Gen. & Sp. 3: 128. 1819. W. mac-
rophylla Cham. & Schlecht. Linnaea 6: 382. 1831. W. scorpioides
Choisy, Ann. Sc. Nat. Se*r. 1, 30: 249. 1833. W. caracasana var. ma-
crophylla (Cham. & Schlect.) Brand, Pflanzenr. IV, 251: 136. 1913.
W. caracasana var. calycina Brand, I.e. W. caracasana var. viscosa
(Donn.-Sm.) Macbride, Contr. Gray Herb. New ser. 49: 42. 1917.
W. kunthii var. intermedia Brand, Pflanzenr. IV, 251: 137. 1913.
W. kunthii var. viscosa (Donn.-Sm.) Brand, I.e. Chocon, chichicaste
(Guatemala, El Salvador and Honduras); tabacon (Jalapa), taba-
quillo (Costa Rica).
Mexico to Costa Rica; Colombia and Venezuela.
Differs from W. urens only in its lack of bristles on leaves and
stems. The leaves usually appear velvety, without bristles on the
mid vein. The calyx lobes and capsule, however, may be hispid to
bristly-hirsute, as in W. urens.
BORAGINACEAE. Borage Family
DOROTHY N. GIBSON
Annual or perennial herbs, shrubs, trees, or woody vines, usually with scabrous,
setose, or hispid indument, rarely tomentose or glabrous; stipules none; leaves alter-
nate or rarely opposite or verticillate, entire or dentate; inflorescence normally
dichotomous-cymose but sometimes greatly modified, the flowers often secund,
sessile or pedicellate, arranged on helicoid or scorpioid cymes; flowers small or large,
112 FIELDIANA: BOTANY, VOLUME 24
usually blue, purple, or white, sometimes yellow, regular, perfect or rarely by abor-
tion polygamous; calyx usually persistent and often accrescent in fruit, the tube
when present usually campanulate, the teeth, lobes, or segments usually 5, rarely
fewer or 6-8, slightly imbricate or often open in bud; corolla gamopetalous, funnel-
form, tubular, salverform, campanulate, or subrotate, the limb usually 5-lobate,
imbricate or rarely contorted; stamens as many as the corolla lobes and alternate
with them, inserted in the tube or throat of the corolla, equal or slightly unequal;
filaments short or long, sometimes dilated at the base, the anthers ovate, oblong,
or linear, obtuse or appendaged, dorsifixed above the base, shallowly or deeply
bilobate at the base, the cells distinct, parallel, introrsely or laterally dehiscent by
longitudinal slits; disk annular, entire or 5-lobate, usually inconspicuous and con-
tinuous with the ovary, or obsolete; ovary superior, sessile and confluent at the
base with the disk, usually composed of 2 carpels, entire or 4-lobate, the cells 1-2-
ovulate; style in the lobate ovary inserted between the lobes, in the entire ovary
terminal, filiform or short, entire or 1-2-bifid or rarely divided into 2 styles; stigma
entire and capitate, or bilobate, or annular; ovules erect, oblique, or subhorizontal;
fruit drupaceous, with a hard 2-4-celled endocarp, or the fruit separating into 2-4
nutlets, or by abortion 1-celled and consisting of a single nutlet, the exocarp fleshy
and juicy, or when the fruit composed of 4 or by abortion fewer nutlets, these
usually hard and distinct, indehiscent, often echinate or variously roughened;
seeds straight or incurved; endosperm carnose, abundant, scant, or none; embryo
straight or curved; cotyledons flat, plano-convex, or thick and fleshy, the radicle
short.
About 90 genera, widely distributed in both hemispheres, fre-
quently weedy. Excepting two, which are native to Europe but
frequently grown in Guatemalan gardens (Borago officinalis L. and
Myosotis scorpioides L.) and an Asian species (Cynoglossum amabile
Stapf & Drumm., which is becoming naturalized in Guatemala),
only the following are known from Central America.
It is often difficult to delimit certain genera, especially if fruiting
material is not available. Some shrubby species of Heliotropium are
much like Tournefortia; flowering material of Hackelia, Cynoglossum,
and Myosotis is often confused in herbaria; monographic work would
be most helpful.
Style inserted between the usually 4 distinct lobes of the ovary; plants herbaceous.
Nutlets echinate; corolla blue.
Nutlets erect, the spines of at least some of them always more than 1 mm.
long (1.5-4 mm.); gynobase pyramidal; upper cauline leaves more or less
narrowed to the base Hackelia.
Nutlets widely spreading, the spines less than 1 mm. long; gynobase de-
pressed; upper cauline leaves rounded at the base and more or less
clasping Cynoglossum.
Nutlets not echinate; corolla white or yellow.
Stamens exserted Macromeria.
Stamens included Lithospermum.
Style apical on the ovary; plants trees, shrubs, or vines, except in Heliotropium.
Style twice bifid Cordia .
GIBSON: FLORA OF GUATEMALA 113
Style entire or only bifid.
Fruits dry, separating into 2-4 bony nutlets; herbs or low shrubs.
Heliotr opium.
Fruits drupaceous; shrubs, trees, or vines.
Leaves in fascicles Rochefortia.
Leaves not in fascicles.
Stigma annular or conic, topped by a sterile, usually bifid appendage.
Tournefortia.
Stigmas 2, truncate, capitate, or clavate.
Calyx closed in bud, valvate, irregularly 2-5 lobate at anthesis.
Bourreria.
Calyx not closed in bud, the 5 lobes distinct, often imbricate.
Ehretia.
BOURRERIA P. Browne
References: P. Browne, Civ. Nat. Hist. Jamaica 168. 1756; M.
Glirke in Engler & Prantl, Pflanzenf. IV, 3a: 86-87. 1897.
Trees or shrubs, glabrous or pubescent, often tomentulose; leaves mostly alter-
nate, usually petiolate, smooth or scabrous, the margins entire, dentate or serrate;
inflorescences terminal, of dichotomous, corymbose cymes, usually many-flowered,
the flowers white, small or large, often fragrant; calyx in bud globose or ovoid,
closed, at anthesis cleft into 2-5 valvate teeth or lobes; corolla 5-lobate, the
lobes imbricate in bud, the tube short or elongated, the throat often dilated,
the lobes broad and spreading; stamens 5, attached to the tube, included or ex-
serted, the filaments glabrous or villous near the base, the anthers ovate or oblong,
2-3 mm. long; ovary at anthesis 4-celled, the style terminal, bifid, the stigmas
truncate, capitate or clavate; ovules attached laterally above or below the middle;
fruit drupaceous, subglobose or ovoid, separating into 4 hard, often triquetrous
nutlets, the central column 4-parted; seeds attached laterally, the endosperm car-
nose; cotyledons flat.
Perhaps 15 or 20 species, from Mexico to the West Indies and
northern South America. The generic name has been written Beu-
reria, but the name Bourreria is conserved.
Filaments of stamens villous near base.
Upper surface of leaf blades smooth; calyx glabrous outside B. huanita.
Upper surface of leaf blades scabrous; calyx pubescent to velvety or tomentose
outside B. andrieuxii.
Filaments of stamens glabrous.
Lower surface of leaf blades glabrous, or puberulent only along veins.
B. oxyphylla.
Lower surface of leaf blades densely and finely grayish-pubescent B. mollis.
Bourreria andrieuxii (A. DC.) Hemsl. Biol. Cent. Am. Bot. 2:
369. 1882. Ehretia andrieuxii DC. Prodr. 9: 510. 1845. Crematomia
andrieuxii Miers, Contr. Bot. 2: 254. 1869.
114 FIELDIANA: BOTANY, VOLUME 24
At 800-900 meters, Huehuetenango. Mexico, Honduras, Nica-
ragua.
Shrubs or small trees, to 4.5 m. tall; leaves on stout petioles 1.5-3 cm. long,
the blades coriaceous, with 7-9 pairs of lateral veins, ovate to broadly ovate,
6-12 cm. long, 3-9 cm. wide, acute to almost rounded at apex, acute to subtrun-
cate at base, entire, upper surface scabrous, lower surface finely and softly pubes-
cent; inflorescence cymose, terminal, ca. 8 cm. long and wide, many-flowered, the
stems and peduncles puberulent to velvety or tomentose, flowers nearly sessile or
on pedicels 1-4 mm. long (rarely to 10 mm. long) ; calyx campanulate, often some-
what contracted at base, 4-5 mm. long, the 5 lobes lanceolate, acute, about as long
as the tube, sparsely or densely pubescent, often velvety outside, white tomentose
within; corolla white, sometimes pinkish in age, 1-1.5 cm. long, the limb 1-1.5 cm.
broad, the rounded lobes ca. 6 mm. long, spreading; stamens exserted, filaments
villous near base, anthers 2 mm. long; fruits broadly ovoid, pointed, 7mm. long,
5 mm. broad (perhaps immature).
The single collection from Guatemala, Steyermark 514-02, super-
ficially resembles B. pulchra Millsp. (Yucatan) but the leaf surfaces
of B. pulchra are never scabrid; its calyces are 5-6 mm. long, glabrous
outside, and its anthers 3 mm. long. It has been confused with
Ehretia latifolia A. DC.
Bourreria huanita (Llave & Lex.) Hemsl. Biol. Cent. Am. Bot.
2: 370. 1882. Morelosia huanita Llave & Lex., Nov. Veg. Descr. 1:
1. 1824. Ehretia formosa A. DC. Prodr. 9: 510. 1845. B. grandiflora
Bertol. Fl. Guat. 410. 1840 (type said to be from Escuintla). B. for-
mosa Hemsl. Diagn. PI. Nov. 34. 1879. Esquinsucha, esquinsuchil
(Guatemala) ; oreja de Ie6n (Quezaltenango) .
Damp forest, sea level to 2,100 m.; often planted for ornament;
Alta Verapaz; El Quiche"; Guatemala; Izabal; Jutiapa; Quezalte-
nango; Sacatepe"quez. Southern Mexico; Honduras; El Salvador;
Costa Rica.
Trees, essentially glabrous throughout; leaves on petioles usually 1-3 cm. long,
the blades glabrous or sometimes with scattered short hairs on costae and petioles,
membranaceous, usually obovate to elliptic-oblong, rarely ovate, mostly 6-12 cm
long, 3-8 cm. wide, obtuse to rounded or acuminate at the apex, broadly rounded
to acute and sometimes oblique at the base, margins entire, lateral nerves 7-9 pairs;
cymes usually large, ca. 8 cm. long and as broad, usually many-flowered, the flowers
sessile or on pedicels 1-6 mm. long; calyx campanulate, 6-8 mm. long, apiculate
in bud, glabrous outside, white with short tomentum within, the 5 lobes unequal,
1-2 mm. long, triangular, acute or subacute, usually rupturing into 3 unequal seg-
ments in fruit; corolla white, ca. 2 cm. long, the tube only slightly exceeding the
calyx, the limb 2-3 cm. broad, glabrous outside but sometimes with a tuft of ap-
pressed sericeous hairs within the lobes near the apex; stamens long-exserted, the
filaments villous near the base; fruits large, ovoid, when dry ca. 12 mm. long and
17 mm. broad.
FIG. 22. Bourreria oxyphylla. A, habit of plant, >2 natural size; B, a larger
leaf, j/2 natural size; C, flower, X 3; E, calyx, style, and stigmas, X 3.
115
116 FIELDIANA: BOTANY, VOLUME 24
Bourreria mollis Standl. Trop Woods 8: 5. 1926. Opay; roble;
black fiddlewood.
Known only from British Honduras, at or a little above sea level.
Trees, the branches densely velutinous-pubescent; leaves on petioles 8-15 mm.
long, the blades elliptic or ovate-elliptic, 3-8.5 cm. long, 1.5-5.5 cm. wide, acute or
obtuse, obtuse or broadly cuneate at the base, glabrate above, densely grayish-
pubescent to velutinous beneath; cymes few-many-flowered, pedunculate, 4 cm.
broad, the flowers short-pedicellate, peduncles and pedicels densely tomentose,
calyx narrowly campanulate, 5-6 mm. long, densely tomentulose, the lobes usually
3, triangular or rounded-ovate, obtuse or subacute, shorter than the tube; corolla
white, the tube slightly longer than the calyx, the lobes 5 mm. long, rounded at the
apex, puberulent outside; stamens exserted, the filaments glabrous, anthers 2 mm
long. Fruits unknown.
Bourreria oxyphylla Standl. Trop. Woods 16: 40. 1928 (type
from British Honduras, Aitken 4)- B. wagneri Standl. in Yuncker,
Field Mus. Bot. 9: 328. 1940 (as Beureria). Laurel, roble, sac-bay-
eck, wild craboo (British Honduras).
Damp or wet, mixed forest, 800 m. or less; Pete'n. Chiapas,
Mexico; British Honduras, Honduras.
Trees, to 18 m., the trunk almost a meter in diameter, sometimes fluted, the
young branchlets sparsely hirtellous or glabrate; leaves on slender, somewhat pu-
bescent or glabrate petioles, 8-12 mm. long, the blades oblong-elliptic or oblong,
4-11.5 cm. long, 2.5-5.5 cm. wide, acute or short-acuminate, acute or obtuse at the
base, firm-membranous, with 6-8 pairs of lateral nerves, entire, essentially gla-
brous, sometimes puberulent beneath with scattered hairs along the nerves; cymes
many-flowered, to 7.5 cm. long and broad, pedunculate, grayish-puberulent, the
pedicels 2-4 mm. long (rarely to 10 mm.); calyx narrowly campanulate, 5-6 mm.
long, obtuse and abruptly contracted at the base, more or less hirtellous outside,
often densely so, at least near the base, white tomentose or sericeous within, the
lobes usually 3-4, ovate to lance-ovate, shorter than the tube; corolla 8-10 mm.
long, the limb 8-12 mm. across, the tube equalling the calyx, the lobes broadly obo-
vate, rounded, more or less sericeous outside, spreading; stamens exserted, fila-
ments glabrous, anthers 2-2.5 mm. long; fruits subglobose, ca. 15 mm. long, 12 mm.
in diameter, first orange-red, but black when mature.
GORDIA L.
References: Ivan M. Johnston, Observations on the species of
Cordia and Tournefortia known from Brazil, Paraguay, Uruguay,
and Argentina, Contr. Gray Herb. 92: 3-65. 1930; Studies in Bora-
ginaceae, X, The Boraginaceae of Northeastern South America,
Journ. Arn. Arb. 16: 1-46. 1935; Notes on some Mexican and Cen-
tral American species of Cordia, I.e. 21: 336-355. 1940; Studies in
GIBSON: FLORA OF GUATEMALA 117
the Boraginaceae, XVII, A. Cordia section Varronia in Mexico and
Central America, I.e. 30: 85-104. 1949; Studies in the Boraginaceae,
XIX, B. Cordia section Gerascanthus in Mexico and Central America,
I.e. 31: 179-187. 1950.
Trees or shrubs, often with very rough pubescence, the hairs simple, stellate,
or branched; leaves all or mostly alternate, petiolate, the margins entire or den-
tate; inflorescences paniculate, cymose, spicate or capitate; flowers small or large,
sessile or pedicellate; calyx tubular to campanulate, sulcate, striate, or smooth,
2-5 lobate, accrescent; corolla most often white, funnelform, salverform, or cam-
panulate, the lobes or divisions usually 5, sometimes more numerous, the lobes in
bud plicate or plane, imbricate or subcontorted ; stamens as many as the corolla
lobes, the filaments equally or unequally inserted on the corolla tube, exserted or
included; anthers ovate, oblong or linear, sagittate or hastate; ovary 4-celled, the
style elongated, twice bifid, the stigmas capitate or clavate; ovules erect, laterally
attached at or below the middle or at the base; fruits drupaceous, the stone very
hard, the cells 4 and 1-seeded or by abortion fewer; seeds without endosperm.
About 250 species in the tropics of both hemispheres, most numer-
ous in America. In addition to the 22 species occurring in Guate-
mala, a number of others are found in Central America.
Inflorescences cymose or paniculate.
Inflorescences of dichotomous cymes.
Cymes lax, open; corollas funnelform; leaves broadly ovate, truncate or
rounded at base C. truncatifolia.
Cymes much reduced, often somewhat headlike; corollas tubular-campanu-
late; leaves elliptic-oblong or ovate-oblong, cuneate at base. . .C. foliosa.
Inflorescences paniculate.
Calyces at anthesis tubular, tubular-campanulate, or short-cylindrical.
Calyx tubes conspicuously sulcate.
Indument of stellate hairs C. alliodora.
Indument of simple hairs.
Corollas 15-25 mm. long, marcescent C. gerascanthus.
Corollas 5-11 mm. long, not marcescent.
Leaves rounded cordate or ovate C. cordiformis.
Leaves lanceolate, lance-oblong or oblanceolate.
Leaf blades nearly glabrous, margins conspicuously serrate-den-
tate C. skutchii.
Leaf blades hirsute and scabrous above, sometimes pilose beneath;
margins entire or only somewhat serrate near apex.
C. diver si folia.
Calyx tubes striate or obsoletely sulcate or smooth.
Corollas bright orange or vermillion.
Calyces at anthesis truncate at apex, later becoming urn-shaped, en-
closing the fruit, the orifice inconspicuous; corolla limb 12-16-
lobate C. dodecandra.
Calyces at anthesis 2-3-lobate, later becoming flask-shaped, enclosing
the fruit, the orifice narrow but evident; corolla limb 5-6-lobate.
C. sebestena.
Corollas white.
118 FIELDIANA: BOTANY, VOLUME 24
Calyces 10-14 mm. long, the lobes 2, unequally rupturing, spreading.
C. megalantha.
Calyces 3.5-5 mm. long, the lobes 5, triangular, erect C. bicolor.
Calyces at anthesis campanulate to broadly campanulate.
Calyx lobes 5, sharply triangular, in fruit spreading and distinctly star-
shaped C. stellifera.
Calyx lobes usually 3 (rarely 4 or 5), irregularly torn or broadly and un-
evenly triangular or rounded, never star-shaped in fruit.
Corollas 5-6 mm. longer than calyces, the lobes broader than long, not
reflexed; leaf blades more or less toothed C. dentata.
Corollas only 2-3 mm. longer than calyces, the lobes longer than broad,
reflexed; leaf blades entire or nearly so.
Calyces densely fulvous-strigose; inflorescences developing with or be-
fore the new leaves C. collococca.
Calyces only partly and minutely strigose; inflorescences developing
after the leaves C. prunifolia.
Inflorescences spicate or capitate.
Inflorescences spicate.
Spikes long and slender, often interrupted, 3-15 cm. long.
Inflorescences chiefly lateral; leaves ovate, lance-ovate or oblong-ovate
2-6 cm. broad, bases rounded and abruptly decurrent. . .C. spinescens.
Inflorescences terminal, never arising from leaf axils; leaves lanceolate,
lance-oblong or narrowly oblanceolate, 1-3 cm. broad, bases acute or
attenuate C. curassavica.
Spikes very dense, thick, ellipsoidal, 2-2.8 cm. long C. coyucana.
Inflorescences capitate-globose.
Calyx lobes caudate, the tips 2-7 mm. long.
Indument of stellate hairs C. pringlei.
Indument of simple hairs.
Peduncles 0.5-2 cm. long, hairs usually appressed; heads 1.5-2 cm. in
diameter; leaves more or less lanceolate C. globosa.
Peduncles 3-7 cm. long, hairs usually erect or spreading, not appressed;
heads 1-1.5 cm. in diameter; leaves usually ovate or oblong-ovate.
C. bullata.
Calyx lobes short-subulate, the tips less than 1 mm. long.
Leaf bases acute to long-attenuate; petioles 4-15 mm. long; peduncles
mostly internodal; indument usually of appressed hairs C. inermis.
Leaf bases rounded or cuneate; petioles 1-3 mm. long; peduncles mostly
arising from leaf axils; indument usually of spreading hairs. . . .C. lineata.
Cordia alliodora (R. & P.) Oken, All. Naturgeschichte, Bot.
2(2): 1098. 1841; DC. Prodr. 9: 472. 1845 (incorrectly as C. alliodora
(R. & P.) Cham.); Johnston, Journ. Am. Arb. 31: 181. 1950. C.
gerascanthus Jacq. Stirp. Amer. 43. 1763, not L. 1759. Cerdana allio-
dora R. & P. Fl. Peruv. 2: 47, 1. 184. 1799. Laurel and bojon (Pet&i) ;
laurel bianco (Sacatepe"quez) .
Usually in dry forests or thickets but sometimes in wet, mixed
forest or along roadsides, sea level to 1,300 m.; Alta Verapaz; Chi-
quimula; El Progreso; Escuintla; Guatemala; Huehuetenango ; Iza-
GIBSON: FLORA OF GUATEMALA
119
FIG. 23. Cordia alliodora. A, habit of plant, ]/% natural size; B, large leaf
from another collection, y^ natural size; C, corolla dissected to show stamens and
pistil, X 3; D, calyx, X 3; E, pubescence from under side of leaf, much enlarged.
bal; Jutiapa; Pete"n; Quezaltenango; Retalhuleu; Sacatepe"quez ; San
Marcos; Santa Rosa; Zacapa. Mexico; Honduras to Panama; West
Indies; through South America southward to northern Argentina
and western Brazil.
Trees to 25 m. high with trunks to 50 cm. in diameter, sometimes with small
buttresses, the bark grayish to dark brown, the branchlets densely or finely stellate-
120 FIELDIANA: BOTANY, VOLUME 24
pubescent; leaves on petioles 1-3 cm. long, the blades usually 7-15 cm. long but
may attain 25 cm., oblong to elliptic, oblong lanceolate, or elliptic-obovate, acute
or acuminate, usually attenuate to base, sometimes obtuse, entire, green and gla-
brate on upper surface, pale beneath and usually densely and minutely stellate-
tomentulose; inflorescences paniculate, the panicles sometimes as much as 30 cm.
broad, usually dense with numerous flowers, the flowers sessile, glomerate; calyx
tubular, 4-7 mm. long, stellate-tomentulose, conspicuously 10-costate, every other
rib prolonged into a short tooth; corolla marcescent, white, at least twice as long
as the calyx, glabrous outside, the lobes oblong-spatulate, broadly rounded or
truncate at the apex; stamens equalling or slightly exceeding the corolla, the fila-
ments inserted in throat of tube, dilated and pubescent at base, just above point of
insertion; ovary elongated, narrowly ovoid; style included or only shortly exserted,
about as long as the ovary; mature fruits ellipsoidal, acute (crowned by the per-
sistent style base), thin-walled and dry, containing one seed.
This is one of the most common trees of the Central American
Pacific lowlands and is often used for lumber in Central America.
The branchlets at the base of the inflorescence are often thickened
and inhabited by small ants.
Cordia bicolor A. DC. in DC. Prodr. 9: 485. 1845; Johnston,
Journ. Am. Arb. 16: 23. 1935. C. trichostyle Pittier, Contr. U. S.
Nat. Herb. 18: 252, /. 102. 1917. C. belizensis Lundell, Am. Midi.
Nat. 29: 488. 1943. Sombra de ternero (Izabal).
Wet, mixed forest, thickets, or pastures, sea level to 550 meters;
Alta Verapaz, Izabal. Mexico to Panama, northern South America.
Trees, 6-12 m. tall, the slender branchlets hispidulous or hirtellous; leaves on
petioles to 1 cm. long, the blades lance-oblong to ovate-oblong, 8-18 cm. long,
3-7.5 cm. wide, gradually or abruptly acuminate, usually rounded to subacute at
base, rarely subcordate, the margins essentially entire, often undulate or crenate,
rarely dentate toward apex, scabrous on upper surface, densely and finely veluti-
nous pubescent beneath or grayish with minute, appressed hairs; inflorescences
terminal, cymose-paniculate, usually large, lax, many-flowered; calyx short-cylin-
drical to tubular-campanulate, obovoid in bud, 3.5-5 mm. long, densely strigose or
strigillose, the 5 lobes triangular, subequal; corolla white, the tube equalling the
calyx, the lobes 2-3 mm. long, reflexed; stamens exserted, the filaments pilose near
the base; fruit inequilaterally ovoid, ca. 1 cm. long, minutely strigillose, seated in
the shallow, cupular, somewhat accrescent calyx.
The only character separating C. belizensis Lundell from C. bicolor
is that of indument of the lower leaf surface, and it appears unstable.
The hairs of Lundell's plant (Gentle 40^5 from Honduras) are erect
or suberect, rather than appressed. However, Turckheim 7963 has
appressed hairs on the lower leaf surface while the hairs of the mid-
rib and petiole are erect and the other costae exhibit both types of
hairs. Lundell also noted that leaves on his plant were entire while
GIBSON: FLORA OF GUATEMALA 121
those of Gentle 3099 which he cited as C. belizensis were dentate. A
rew leaves of Standley 24-990 are also somewhat dentate toward the
apex, and the leaf margins of most collections seen are undulate
and/or crenate.
Cordia bullata (L.) Roem. & Schult. Syst. Veg. 4: 462. 1819;
Varronia bullata L. Syst. ed. 10: 916. 1759. C. bullata var. angustata
DC. Prodr. 9: 496. 1845. C. asperrima DC. I.e. 498. Varronia asper-
rima (DC.) Friesen, Bull. Soc. Bot. Geneve, se>. 2, 24: 155. t. l,f. 5.
1933. Cambray (Chiquimula and Jalapa).
Damp or dry, brushy, sometimes rocky thickets and slopes, occa-
sionally in fields, 500-1,400 meters; Chiquimula, Escuintla; Guate-
mala; Jalapa; Pete"n. Mexico to Nicaragua; West Indies.
Erect shrubs, 1-3 m. tall, the young branches fairly stout, hispid or hispidu-
ous; leaves short-petiolate, the blades broadly ovate to oblong-ovate, 2-6 cm. long,
thick, acute at apex, acute to rounded at base, sometimes abruptly contracted and
decurrent, margins serrate to serrulate, rugose to bullate and hispid on upper sur-
'ace, the setae usually arising from bulbous bases, strigose or hispid beneath with
erect or ascending hairs, veins conspicuous; inflorescences capitate, the heads glo-
3ose, 1-1.5 cm. in diameter, many-flowered, the solitary peduncles 3-7 cm. long;
calyx campanulate, hispid on upper portion, the lobes triangular and caudate, the
iliform tips 1-2 mm. long and hirsute; corolla white, 4-7 mm. long, stamens in-
serted just above the ring of hairs within the corolla throat; ovary globose to ovoid,
the style included; fruits red, 3-4 mm. long, nearly two-thirds enclosed in the ac-
crescent calyx.
Obviously very closely related to C. globosa (Jacq.) HBK. and
perhaps not deserving specific status. Although Guatemalan plants
are readily separable by length of peduncle and type of indument,
some Mexican collections exhibit intergradation between these two
torms.
Cordia collococca L. Sp. PI. ed. 2: 274. 1762. C. micrantha Sw.
Prodr. 47. 1788 and Fl. Ind. Occ. 1: 460. 1797.
Near sea level to 1,400 meters, fide Standley; Escuintla? Guate-
mala? Pete"n. Southern Mexico; El Salvador to Costa Rica; West
Indies; Venezuela.
Dioecious trees, 5-15 m. tall, with spreading branches, the young branchlets
and stems of inflorescences hirtellous or hispidulous; leaves on petioles 3-13 mm.
long, the blades chartaceous, elliptic to obovate or oblong-obovate, deciduous
during the dry season, mostly 7-15 cm. long, obtuse or short-acuminate to acumi-
nate at the apex, usually cuneate, sometimes obtuse at base, entire or nearly so,
glabrous above or nearly so, sparsely hirtellous or hispidulous beneath, at least
along the costa, the hairs mostly erect; inflorescences cymose-paniculate, appearing
122 FIELDIANA: BOTANY, VOLUME 24
with or even before the new leaves, 4-8 cm. broad, dense or open, many-flowered,
the flowers sessile, glomerate; calyx densely fulvous-strigose or strigillose, globose
in bud, broadly campanulate at anthesis, ca. 2 mm. long, unevenly 3-5 dentate,
separating into 3-5 uneven lobes in fruit; corolla white, ca. 5 mm. long; filaments
of stamens villous below; fruits cherry-red, ca. 8 mm. in diameter.
This species has often been called C. glabra L., but that name is
a synonym of Bourreria succulenta Jacq. (see I. M. Johnston, Journ.
Am. Arb. 21:345. 1940).
Cordia collococca is easily confused with C. eriostigma Pittier of
El Salvador and Costa Rica, which also has a three-parted, strigose
calyx, but produces perfect flowers. According to Johnston, it can
be separated by examining the hairs of the lower leaf surfaces, as
those of C. eriostigma are closely appressed rather than erect. How-
ever, two of the Guatemalan specimens in question (Molina 12484,
Dept. Escuintla, and Aguilar 371, Dept. Guatemala) have old and
glabrate leaves, are fruiting and bear no flowers; the third specimen,
Lundell 2743, Pete"n, has only very young leaves. I have placed them
in C. collococca, as it is known to have the wider range and is to be
expected in Guatemala.
Cordia cordiformis I. M. Johnston, Journ. Am. Arb. 18: 10.
1937.
At 800 meters; Guatemala; Santa Rosa. Mexico, 3,200 meters.
Dioecious trees of 6 m. or more with rufous or fulvous pubescence, the young
branches hirsute with stiff, spreading hairs; leaves on stout petioles, the blades
rounded-cordate or ovate, mostly 4-15 cm. long and wide, obtuse or abruptly
short-acute at the apex, cordate or rounded at the base and abruptly contracted
into a petiole 1-5 cm. long, repand-denticulate, strigose on the upper surface, paler
beneath, densely tomentose with long, soft, often tangled hairs, the lateral nerves
4-6 pairs; inflorescences cymose-paniculate, terminal, short-pedunculate, 7-15 cm.
broad; calyx 3-4 mm. long, 10-costate, obovoid in bud, short-cylindrical to tubular
at anthesis (shallowly cupular in fruit), pubescent with unequal subappressed
hairs; corolla of staminate flowers pinkish, 6-7 mm. long, the tube ca. 3 mm., the
lobes oblong, recurved; corolla of pistillate flowers not seen; ovary glabrous, glo-
bose; dry fruits 7 mm. long.
Cordia coyucana I. M. Johnston, Journ. Arn. Arb. 29: 227.
1948 and 30: 94. 1949.
The only Guatemalan collection seen is Standley 76595, rocky oak
forest, 1,360 meters, Jalapa. Mexico, Honduras and Costa Rica.
Erect shrubs to 3 m. tall, the young branches short-hirsute to hispidulous;
leaves nearly sessile or on petioles to 9 mm. long, the blades lanceolate to lance- !
ovate, 4-8 cm. long, 1-3 cm. wide, acute, acute or obtuse at base, often somewhat
GIBSON: FLORA OF GUATEMALA 123
ique, upper surface usually green and subvelutinous, sometimes scabrid, or
papillose without hairs except when young, thinly grey-tomentose beneath with
soft, slender hairs 1-2 mm. long, the longer hairs on costae and near the base,
margins dentate, at least on upper portions; inflorescences terminal, the very dense
spikes ellipsoidal-capitate, 2-2.8 cm. long, 1-1.4 cm. thick, on peduncles 3-5 cm.
long; calyx at an thesis 3-5 mm. long, somewhat striate, usually more or less his-
pidulous, sometimes hirsute with appressed hairs, the lobes broadly triangular;
corolla white or greenish-white, 4-6 mm. long, the lobes irregularly sinuate to erose
or emarginate; stamens about equalling or somewhat exceeding the sinuses, the
glabrous filaments 2-3 mm. long, inserted just above the ring of hairs within the
corolla throat; fruits partially enclosed in the accrescent calyx.
These plants, as Johnston pointed out, despite their nearly capi-
tate inflorescences, appear to be very closely related to C. curassavica
(Jacq.) Roem. & Schult., and perhaps may not really be distinct.
The Guatemalan collection and others from Honduras and Costa
Rica were originally distributed with an unpublished name provided
by Johnston.
Cordia curassavica (Jacq.) Roem. & Schult. Syst. Veg. 4: 460.
1819; Johnston, Journ. Arn. Arb. 30: 99. 1949. Varronia curassavica
Jacq. Enum. PI. Carib. 14. 1760. V. macrostachya Jacq. I.e. C. macro-
stachia (Jacq.) Roem. & Shult. Syst. Veg. 4: 461. 1819. C. brevispi-
cata Mart. & Gal. Bull. Acad. Brux. 11 (2): 331. 1844. C. hispida
Benth. Bot. Sulphur 139. 1845. C. peruviana var. mexicana DC.
Prodr. 9: 491. 1845. C. cylindrostachya var. graveolens (HBK.)
Griseb. Fl. Brit. W. I. 480. 1861. C. cylindrostachya var. interrupta
(DC.) Griseb. I.e. C. brevispicata var. hypomalaca Greenm. Field
Mus. Bot. 2: 338. 1912. C. imparilis Macbride, Contr. Gray Herb.
1 49: 16. 1917. C. chepensis Pitter, Contr. U. S. Nat. Herb. 18: 253.
1917. C. littoralis Pittier, I.e. C. mollis Pittier, I.e. 294. Lantana
bullata L. Sp. PI. 627. 1753, not Cordia bullata (L.) Roem. & Schult.
Chincura, moco de chumpe, morellita (Zacapa).
Damp or dry thickets, hillsides, sometimes in oak forests, sea
level to 1,360 meters; Chiquimula; Guatemala; Jalapa; Jutiapa;
Retalhuleu; Zacapa. Southern Mexico to Panama; West Indies;
northern South America.
Erect shrubs to 3 m. high, the young branches resinous glandular or sparsely
or densely hispidulous, short-hirsute, or strigose; leaves nearly sessile or on short
petioles to 8 mm. long, the blades thick or thin, lanceolate to oblong- ovate, or
narrowly elliptic or oblanceolate, 3-12 cm. long, usually 1-3 cm. broad, but to
7 cm. in one form, acute to acuminate or obtuse, acute or attenuate to the base,
usually serrate-dentate, sometimes entire or nearly so, pilose or hirsute above, often
scabrid, sometimes papillose and without hairs except when young, somewhat paler
124 FIELDIANA: BOTANY, VOLUME 24
beneath and puberulent, sometimes glandular, the veins conspicuous; inflorescences
spicate, terminal (in that they never arise from leaf axils), the spikes 3-10 cm. long,
dense or interrupted; calyx campanulate, 2-3.5 mm. long, granulate and strigillose
or only somewhat granulate and hispidulous with spreading hairs; corolla white or
greenish-white, 3.5-6 mm. long, the lobes 1-1.5 mm. long, broadly rounded and
reflexed, or irregularly sinuate to erose or emarginate; stamens about equalling or
somewhat exceeding the sinuses, the short, glabrous filaments inserted just above
the ring of hairs within the corolla throat; fruits red, partially enclosed in the ac-
crescent calyx, the mature drupe broadly ovoid, 4-5 mm. long.
A complex species of several diverse forms, some of which, as
Johnston observed, may eventually be treated as separate species.
The two most diverse forms, both in Guatemala, are those plants
with elongated, interrupted inflorescence spikes and hispiduous ca-
lyces with spreading hairs, and those with shorter, more dense spikes
and granulate, strigillose calyces with short, appressed hairs. Co-
rollas of the elongated, interrupted spikes tend to have more broadly
rounded lobes, eventually reflexed, while corolla lobes from the
shorter, dense spikes are not as often reflexed and are more apt to be
irregularly sinuate or erose.
Because the few Guatemalan collections can be readily separated
into these two different forms, one is tempted to maintain them sep-
arately. However, when the extremely variable collections from
Mexico, Honduras, Costa Rica, and Panama are examined, separa-
tion becomes most difficult. The shorter, dense spikes of the second
group of plants sometimes elongate and become interrupted with age;
collections from Honduras and Panama have calyces that are granu-
late on the lower part, appressed strigillose above, with a few long,
spreading hairs on the lobes. Therefore, until further study of the
intergradation of these and related plants from the West Indies and
South America can be made, they are treated here collectively as
C. curassavica (Jacq.) Roem. & Schult., sens. lat.
Cordia dentata Poir. Encycl. 7: 48. 1806; Vahl, Eel. Amer. 3: 5,
t. 22. 1807; Johnston, Journ. Am. Arb. 21: 347. 1940. C. calyptrata
Bertero ex Sprengel, Syst. 1: 649. 1825. C. tenuifolia Bertol. Rendic.
Sess. Accad. Bologna 1860-61: 73. 1860 and Mem. Accad. Sci. Bo-
logna 11: 199, 1. 11. 1861. C. leptopoda Krause, Bot. Jahrb. 37: 628.
1906. C. ovata Brandegee, Pub. Univ. Calif. Bot. 10: 187. 1922.
Varronia calyptrata (Bert.) DC. Prodr. 9: 469. 1845; Delessert, Icon.
5: 41, t. 97. 1846. Upay, supay, tiguilote (Jutiapa, Santa Rosa,
Zacapa); Upayol (Chiquimula).
GIBSON: FLORA OF GUATEMALA 125
Dry, sandy or rocky slopes and hills, damp or dry thickets or
mixed forest; grassy plains and pastures, sea level to 900 meters;
Chiquimula; El Progreso; Escuintla; Guatemala; Izabal; Jalapa;
Jutiapa; Santa Rosa; Zacapa. Mexico; Honduras to Panama; West
Indies; Colombia and Venezuela.
Shrubs or trees to 15m. tall, the trunk usually short, often crooked, the bark
light brown and scaly, the branchlets slender, puberulent or short-pilose; leaves on
petioles 1-3 cm. long, the blades 3-10 cm. long, elliptic to ovate or obovate-ellip-
tic, rounded or acute at apex, rounded or cuneate at base, subentire, undulate, or
coarsely dentate, somewhat scabrous on the upper surface, sparsely pilose beneath,
usually along the costa, or almost glabrous, lateral nerves usually 3 to 6 pairs,
oblique; inflorescence cymose-paniculate, often lax, to 20 cm. broad, many-flow-
:ered, the flowers subsessile; calyx 2.5-4.5 mm. long, striate or ribbed, strigose, obo-
|void in bud, open-campanulate at anthesis; corolla 5-lobate, 8-9 mm. long, funnel-
jform or broadly campanulate, with lobes broader than long, white, cream, or
jyellow; stamens included, the short, pubescent filaments attached to tube about
2 mm. above its base; ovary subglobose; style usually somewhat exserted; mature
fruits white, the flesh mucilaginous, somewhat translucent, 6-10 mm. long (said
to attain 16 mm. in the West Indies).
In various publications on the flora of Mexico, Central America,
and the West Indies, this tree has been called Cordia alba (Jacq.)
Roem. & Schult., a name probably to be applied to a plant of an-
other family.
Cordia diversifolia Pavon ex A. DC. in DC. Prodr. 9: 474. 1845.
\C.johnstoni Cufodontis, Arch. Bot. 10: 41. 1934. C. petenensis Lun-
dell, Wrightia 4: 49. 1968 (type from Pete"n, Contreras 894). Upay
; (Izabal); chischis (British Honduras).
Wet thickets or forest, along rivers, in pastures and plains, sea
|level to 450 meters; Alta Verapaz; Escuintla; Izabal; Pete"n; Retalhu-
leu; Santa Rosa. Mexico; British Honduras to Panama.
Dioecious trees, to 10 m. tall, the young branches hirsute or hispid with spread-
ing brown hairs; leaves on petioles 5-15 mm. long, the blades lanceolate, lance-
oblong, or oblanceolate to broadly elliptic, 3-20 cm. long, obtuse to acuminate,
obtuse or acute at base, membranaceous, entire or somewhat serrate near the apex,
[hirsute and scabrous on the upper surface, paler and short-hirsute or pilose beneath,
often densely so; inflorescences cymose-paniculate, the panicles usually small, dense
|or open, few or many-flowered, the branches hirsute; calyx at anthesis tubular
:hispidulous, the hairs underlain by papillae, costa te, 4-7 mm. long, with minute
[teeth, but separating into 3 uneven lobes in age; corolla creamy white, the tube
6-7 mm. long, the lobes 1-3 mm. long, reflexed; stamens (staminate flowers) well
exserted, the filaments villous at base; style in female flowers exserted; absent in
male flowers; drupes ovoid, glabrous, ca. 7 mm. long, white at maturity.
126 FIELDIANA: BOTANY, VOLUME 24
The concept C. petenensis Lundell is a small-leaved form of this
species.
Cordia dodecandra A. DC. in DC. Prodr. 9: 478. 1845. C. angi-
ocarpa A. Rich, in Sagra, Fl. Cub. Fanerog. 2: 110, t. 60. 1853.
Plethostephia angiocarpa (Rich.) Miers, Trans. Linn. Soc. Ser. II, 1:
32, t. 8. 1875. Cocopera (Jutiapa). Copte, siricote (Pete"n, fide
Lundell) .
Sea level to 900 meters; planted in various places in Jutiapa;
probably native in Alta Verapaz and Pete"n. Mexico; British Hon-
duras; Honduras; Cuba.
Trees, 8-10 m. tall, usually with a broad crown and short trunk, the young
branches stout, hispidulous and brown-tomentose; leaves on short or elongated
petioles, the blades oblong or suborbicular, 6-15 cm. long, 4-8 cm. broad, rounded
or obtuse at apex and at base, entire or somewhat sinuate, very scabrous on both
surfaces, hispidulous beneath; inflorescences cymose-paniculate, the cymes few-
flowered, the panicles terminal and fairly compact; calyx 1-1.3 cm. long, tubular-
campanulate, truncate at apex, glabrous or somewhat tomentulose with a few
scattered, straight, appressed hairs, ferruginous-tomentulose near apex and on
lobes; corolla funnelform, bright orange, 3-5 cm. long, striate with fairly heavy
tomentum along the lines and on margins of lobes, within as well as outside, the
limb 2.5-3.2 cm. across, 12-16 lobate; stamens as many as the corolla lobes, in-
cluded, the puberulent filaments inserted unevenly on tube at a point about 1.5 cm.
above its base; ovary ovoid, seated on a large disc; style elongated, included; ma-
ture fruits greenish-yellow, ovoid-oblong, acute at apex, ca. 2 cm. long, the drupe
enclosed in a fleshy, urn-shaped calyx.
The very sweet fruit is said to be highly esteemed for making
preserves and other dulces.
Cordia foliosa Mart. & Gal. Bull. Acad. Brux. 11(2): 330. 1844.
C. chiapensis Fernald, Proc. Am. Acad. 40: 52. 1904. Salvia santa
(Huehuetenango) .
At 1,300-2,000 meters; Guatemala; Huehuetenango. Southern
Mexico.
Shrubs to 3 m. tall, the slender branches hispidulous or glabrate; leaves on
petioles 1-7 mm. long, the blades membranaceous, elliptic-oblong to lance-oblong,
usually 3-6 cm. long, 1-2.5 cm. wide, acute or subobtuse at apex, cuneate at base,
the margins serrate, upper surface scabrous, lower surface hispidulous; inflores-
cences cymose, on peduncles 2-5 cm. long, the cymes dichotomous, usually much
reduced, somewhat head-like, ca. 2 cm. broad, the flowers subsessile, secund; calyx
campanulate (turbinate and apiculate in bud), 3-5 mm. long, scabrous or strigose,
the triangular lobes subulate, the tips usually ca. 1 mm. long; corolla campanulate,
white, 4-7 mm. long; stamens equalling or exceeding the corolla; fruits ovoid, par-
tially enclosed in the cupular, somewhat accrescent calyx.
GIBSON: FLORA OF GUATEMALA 127
Cordia gerascanthus L. Syst. ed. 10. 936. 1759; Johnston,
Contr. Gray Herb. 73: 77. 1924. C. gerascanthoides HBK. Nova
Gen. & Sp. 3: 69. 1818. C. bracteata DC. Prodr. 9: 472. 1845. C.
langlassei Loes. in Fedde, Rep. Spec. Nov. 12: 240. 1913. C. roth-
schuhlii Loes. Bot. Jahrb. 60: 368. 1926. Cerdana gerascanthus (L.)
Moldenke, Phytologia 1: 16. 1933. Gerascanthus lanceolatus J. S.
Presl, Wseob. Rostl. 2: 1103. 1846, in part. Palo de asta (Zacapa).
Laurel negro (Honduras).
Lowlands; the only Guatemalan collections are from Gualan,
along railroad, transition from desert to wet land, Zacapa, and from
Cadenas, on rocky hill, Pete"n. Mexico; Honduras; Nicaragua; Costa
Rica; West Indies; Colombia.
Trees, usually 10-15 m. tall, but may attain 30 m., with a trunk to 30 cm. in
diameter, the branches glabrate; leaves slender-petiolate, lanceolate, lanceolate-
ovate, or elliptic-oblong, entire, smooth or nearly so, 5-12 cm. long, acute or acumi-
nate, acute at the base; flowers sessile in dense cymes, the branches covered with a
dense, fine pubescence; calyx 7-10 mm. long, tubular, 10-costate, usually thinly
hirsute with short hairs or glabrescent, the 5 lobes deltoid; corolla marcescent,
funnelform, white, becoming lavender with age and finally brown, 1.5-2.5 cm.
long, the cylindrical tube equalling the calyx, the 5 lobes obovate, truncate to
retuse, separating about midway of the limb, the sinuses to 1 mm. wide; stamens
usually exserted, filaments inserted in tube just below the throat, more or less
pubescent, wooly at base, anthers linear, sagittate; ovary short-cylindrical, style
shortly exserted, the branches deeply bifid; fruit ellipsoidal, the fibrous walls en-
closing a single seed; mature fruits remaining enclosed by the persistent corolla
tube and the calyx, which it somewhat exceeds.
Although heterostyly is common in the species, the flowers of the
Guatemalan specimen display long stamens, surpassing considerably
the only shortly exserted style.
In Mexico and in the West Indies the wood of the tree is used in
carpentry and construction work.
Cordia globosa (Jacq.) HBK. Nova Gen. & Sp. 3: 76. 1818.
Varronia globosa Jacq. Enum. PL Carib. 14. 1760. V. humilis Jacq.
I.e. Cordia humilis (Jacq.) D. Don, Gen. Syst. 4: 383. 1838. C. jac-
meliana Krause, Beih. Bot. Centralb. 32(2): 344. 1914. C. globosa
var. humilis (Jacq.) I. M. Johnston, Journ. Am. Arb. 30: 98. 1949.
Varronia jacmeliana (Krause) Friesen, Bull. Soc. Bot. Geneve, se>. 2,
24: 177. 1933. V. humilis var. mexicana Friesen, I.e. 162, t. 1, f. 4-
V. mexicana Friesen, I.e. Diente de perro, diente de chucho (Zacapa) ;
palo negro (Huehuetenango) .
128 FIELDIANA: BOTANY, VOLUME 24
Damp or dry, brushy, sometimes rocky plains and hillsides,
coastal thickets, sand dunes, sea level to 1,600 meters; Chiquimula;
Huehuetenango; Retalhuleu; San Marcos; Zacapa. Mexico to Pan-
ama; West Indies; northern South America.
Erect shrubs, 1-3 m. tall, the young branches strigose with appressed hairs,
leaves short-petiolate, the blades lanceolate, lance-ovate or ovate-oblong, 1.5-7 cm.
long, acute or subacute at apex, acute to rounded at base, sometimes abruptly con-
tracted and decurrent, the margins crenate-dentate to dentate, somewhat rugose
and strigose with appressed hairs on upper surface, more densely appressed strigose
on lower surface, the veins usually elevated and prominent; inflorescences capitate,
the heads globose, 1.5-2 cm. in diameter, many-flowered, on peduncles 0.5-2 cm.
long; calyx campanulate, hispid, the lobes triangular and caudate, the filiform tips
1-2 mm. long and hirsute; corolla white, 5-9 mm. long; stamens inserted just above
the ring of hairs within the corolla throat; ovary globose to ovoid, the style in-
cluded; fruits red, ovoid, partially enclosed in the accrescent calyx.
Cordia inermis (Mill.) I. M. Johnston, Journ. Arn. Arb. 30: 95.
1949. Lantana inerma Miller, Card. Diet. 1768. Cordia cana Mart.
& Gal. Bull. Acad. Brux. 11(2): 331. 1844. C. insularis Greenm.,
Proc. Am. Acad. 33: 482. 1898. Cuajatinta (Guatemala).
Damp or dry lowland forest or on brushy, rocky slopes, near sea
level to 1,200 meters; Alta Verapaz; Chiquimula; Escuintla; Guate-
mala; Santa Rosa; Zacapa. Mexico to Panama. Northern Colombia.
Shrubs to 3 m. tall, usually densely branched, the slender branches strigose
with short, whitish appressed hairs; leaves on petioles 4-15 mm. long, the blades
membranaceous, lanceolate to oblong, ovate or elliptic, 2.5-7.5 cm. long, usually
1-2.5 cm. broad (rarely to 5 cm. broad), long-acuminate to subobtuse at apex, acute
to long-attenuate at base, the margins serrate or subentire, the upper surface green
and somewhat scabrous, paler and often densely strigose beneath; inflorescences
capitate, the heads globose, 5-10 mm. in diameter, usually with 20 or more flowers,
the peduncles 1-4 cm. long, terminal or axillary (mostly internodal), their indument
usually of appressed hairs; calyx campanulate, 1.5-3 mm. long, strigose, the tri-
angular lobes short-subulate, 0.5-1 mm. long; corolla white or greenish white,
3-5 mm. long, tubular; stamens about equalling or slightly exceeding the corolla;
ovary globose to ovoid, the style equalling the corolla or somewhat exserted; fruits
red, 4-5 mm. long, about three-fourths enclosed by the accrescent calyx.
Cordia lineata (L.) Roem. & Schult. Syst. Veg. 4: 464. 1819;
Johnston, Journ. Arn. Arb. 30: 91. 1949. Varronia lineata L. Syst.
ed. 10: 916. 1759. Cordia ulmifolia var. lineata (L.) DC. Prodr. 9:
495. 1845. Lantana corymbosa L. Sp. PI. 628. 1753.
Damp thickets, brushy slopes, sea level to 225 meters; Izabal.
Southern Mexico to Panama; West Indies; possibly northern South
America.
GIBSON: FLORA OF GUATEMALA 129
Slender, erect shrubs of 1-5 meters; the stems hirtellous or pilose with short
and long, spreading hairs; leaves on petioles 1-3 mm. long, the blades membrana-
ceous, lanceolate to ovate-lanceolate, 3.5-10 cm. long, 1-3 cm. broad, acute to
long-acuminate at apex, rounded or cuneate at base, green above, scabrous or
somewhat tuberculate and strigose, densely and finely pubescent beneath, usually
somewhat velutinous, margins serrate or subentire; inflorescences capitate, globose,
the heads usually 5-10 mm. in diameter, usually 6-12-flowered, the peduncles
2-3 cm. long, the majority arising from leaf-axils, their indument usually of spread-
ing hairs; calyx campanulate, 2-3 mm. long, strigose, the triangular lobes short-
subulate, ca. 0.5 mm. long; corolla white or greenish white, 3-5 mm. long, tubular;
stamens about equalling or slightly exceeding the corolla; ovary globose to ovoid,
the style equalling the corolla or somewhat exserted; fruits red, 4-5 mm. long,
nearly three-fourths enclosed by the accrescent calyx.
Reported from Mexico and Central America as C. corymbosa (L.)
Don and as C. ulmifolia Juss., both untenable names (Johnston,
Journ. Arn. Arb. 16: 33. 1935). Also reported as C. polycephala
(Lam.) I. M. Johnston, which according to Johnston is a separate
species of the West Indies and northern South America. However,
further studies may well prove it to be synonymous with C. lineata.
Cordia megalantha Blake, Proc. Biol. Soc. Wash. 36: 200. 1923.
C. macrantha Blake, Contr. U. S. Nat. Herb. 24: 19. 1922, not Chodat.
Laurel bianco (Zacapa) .
Open or forested slopes, at or a little above sea level; Izabal;
Zacapa. Honduras.
Trees, 10-25 meters tall, the trunk to 66 cm. in diameter, the branchlets gla-
brous or nearly so; leaves on slender petioles 1-4 cm. long, the blades chartaceous,
elliptic-oblong to broadly obovate, or ovate, mostly 7-15 cm. long, 3-8 cm. wide,
obtuse or acute or broadly rounded and abruptly pointed at apex, cuneate to al-
most rounded at base, entire or practically so, glabrous; inflorescences cymose-
paniculate, terminal, 6-12 cm. long, the branches pubescent and glandular, the
pedicels 2-3 mm. long; calyx broadly tubular, striate, glabrous or minutely puberu-
lent, 10-14 mm. long, with 2 unequal, broadly triangular, ciliolate, spreading lobes;
corolla white, marcescent, 2-3 cm. long, the limb to 4 cm. broad, the 5 lobes tri-
angular-ovate, obtuse or acute; stamens about equalling the sinuses, the filaments
inserted in tube below the throat, somewhat hirsute, densely pubescent at and
above point of attachment; ovary glabrous, short-cylindrical to oblong; style in-
cluded ; fruits enclosed in the persistent corolla.
Cordia pringlei Robins. Proc. Am. Acad. 26: 169. 1891. Var-
ronia pringlei (Robins.) Friesen, Bull. Soc. Bot. Geneve, ser. 2, 24:
168. 1933. C. pringlei var. altatensis Brandegee, Zoe 5: 219. 1904.
C. asterothrix Killip, Journ. Wash. Acad. 17: 330. 1927. C. storkii
Standley, Field Mus. Bot. 18: 984. 1938. Cambray, chiligua, col-
millo de perro, diente de perro (Jutiapa) .
130 FIELDIANA: BOTANY, VOLUME 24
Dry, brushy, often rocky plains and hillsides, 200-900 meters;
Chiquimula; El Progreso; Jutiapa; Zacapa. Mexico to Costa Rica.
Erect shrubs to 3 m. tall, the older branches dark and glabrate, the younger
ones densely stellate-torn entose; leaves on stout petioles 0.5-2 cm. long, the blades
thick, oblong-ovate to broadly ovate, 3-10 cm. long, 2-7 cm. broad, acute or sub-
acute at apex, rounded or truncate at base, often abruptly decurrent, sometimes
cordate, coarsely and unevenly crenate-dentate, densely and minutely stellate-
pubescent on the upper surface, densely and softly stellate-tomentose beneath,
pale; inflorescences capitate, on terminal peduncles 2-10 cm. long, the heads glo-
bose, dense, many-flowered, 2-2.5 cm. in diameter; calyx stellate-pubescent, the
tube narrowly campanulate (turbinate in bud), 6-7 mm. long, the triangular lobes
caudate, the filiform tips 6-7 mm. long; corolla white, funnelform, more than twice
as long as the calyx, the limb 1-1.5 cm. broad; stamens exceeding the tube; ovary
ovoid, the style included; fruits ovoid or obovoid, partly enclosed in the somewhat
accrescent calyx.
Cordia stellata Greenm. of Mexico closely resembles C. pringlei,
differing only in the calyx lobes which lack the filiform tips and its
usually slightly smaller flowering heads. C. stellata may not deserve
specific rank.
Cordia prunifolia I. M. Johnston, Journ. Arn. Arb. 21: 353.
1940.
In Guatemala known only from the type, Skutch 298, coffee plan-
tation, evidently a relict of the forest, Sacatepe"quez, 1,080 meters.
Mexico, Costa Rica, Panama.
A tree of 21 m., the trunk 40 cm. in diameter, the bark brown, finely checkered,
young branchlets almost glabrous; leaves on petioles to 1 cm. long, the blades stiff-
membranaceous, oblong-ovate to elliptic-oblong, mostly 10-12 cm. long and 3.5-
5.5 cm. wide, abruptly acute or short-acuminate, obtuse or acute at the base, en-
tire or nearly so, lustrous above and glabrous or nearly so, slightly paler beneath,
with sparse appressed hairs or practically glabrous; inflorescences terminal and
lateral, cymose-paniculate, short-pedunculate, 5-12 cm. broad, lax, many-flow-
ered, the slender branches brownish-puberulent; calyx obovoid- globose in bud and
strigillose, campanulate at an thesis and becoming almost glabrous, 3-3.5 mm. long,
irregularly lobate, the lobes usually 3, usually rounded but occasionally broadly
triangular; corolla white, 5-5.5 mm. long, the lobes oblong, somewhat recurved;
stamens exserted, the filaments villous below; ovary glabrous; fruits unknown.
This species is much like C. nitida Vahl of the West Indies, and
it is quite possible that this plant, as well as C. lasiocalyx Pittier and
C. lucidula Johnston of Costa Rica and Panama should properly be
referred to C. nitida.
Cordia sebestena L. Sp. PI. 190. 1753.
Near sea level; British Honduras. Mexico (Yucatan and Quin-
tana Roo) ; Honduras. West Indies to northern Venezuela.
GIBSON: FLORA OF GUATEMALA 131
Shrubs or small trees, usually not more than 8 m. tall, the branches stout;
leaves on stout petioles 1-3 cm. long, the blades ovate to ovate-elliptic, somewhat
undulate, acute, rounded or almost truncate at base, entire or somewhat dentate
near apex, scabrous above, somewhat scabrous and pubescent beneath, mostly
6-18 cm. long, 5-14 cm. wide, with 5-9 pairs of primary lateral veins; inflorescences
pedunculate, the panicles composed of dichotomous, few-flowered cymes, the flow-
ers on pedicels usually 4-6 mm. long but may be 2-15 mm. long; calyx tubular-
campanulate, rupturing irregularly but usually 2-3-lobed at anthesis, usually pu-
bescent with appre^sed hairs inside and out, ca. 1.5 cm. long; corolla funnelform,
bright orange to vermilion, 3-4 cm. long, the limb 3-3.5 cm. across, 5-6 lobate;
stamens included or slightly exserted, attached to tube at a point 2-3 mm. below
throat, the anthers 3-3.5 mm. long; style included, elongated to ca. 20 mm., per-
sistent; mature fruits hard, dry, ovoid, 2-2.5 cm. long, completely enclosed in the
accrescent, flask-shaped calyx which becomes white and fleshy.
Although not reported from Guatemala, these wide ranging plants
might be expected along the east coast. They are very ornamental
and are widely cultivated in the tropics.
Cordia skutchii I. M. Johnston, Journ. Arn. Arb. 21: 339. 1940.
Canela.
Wet, mixed forest, 1,150-1,500 meters; endemic; Quezaltenango
(types: pistillate, Skutch 1426', staminate, Skutch 1425).
Dioecious trees to 8 m. tall, the branchlets slender, hispidulous with spreading
or ascending hairs; leaves on petioles 1-2 cm. long, the blades membranaceous,
oblanceolate, 10-30 cm. long, 2.5-9 cm. wide, long- acuminate, long-attenuate to
the base, serrate-dentate, with scattered hairs on the upper surface, glabrate be-
neath but with scattered hairs on veins; inflorescences terminal, on long, slender
peduncles, cymose-paniculate, lax and open, 7-16 cm. broad, many-flowered; calyx
tubular at anthesis, 3-5 mm. long, costate or striate, minutely tuberculate or mu-
riculate and hispidulous, irregularly 3-5 dentate, separating into 3 uneven lobes;
corolla white, pubescent within throat, 9-11 mm. long when staminate, 5 mm. long
when pistillate, the lobes ca. 2-4 mm. long, reflexed; stamens of staminate flowers
well exserted, pubescent near point of insertion in throat; mature fruits white,
ellipsoid-ovoid, to 1 cm. long, much exceeding the shallowly cupular fruiting calyx.
As Johnston has pointed out, C. skutchii closely resembles C. di-
versifolia Pavon but is more slender, more loosely branched, less
hirsute, with thinner, nearly glabrous leaves which are always obvi-
ously dentate. Johnston felt that C. skutchii may be more closely
related to the broader-leaved C. salvadorensis Standley, which is not
jas hirsute, with hairs of calyx and herbage much more closely ap-
pressed. Although the three plants can be readily separated by leaf
form and margin, the flowers of the three are so much alike that the
i validity of supposed specific difference may be questioned.
132 FIELDIANA: BOTANY, VOLUME 24
Cordia spinescens L. Mant. 2: 206. 1771; I. M. Johnston, Journ.
Arn Arb. 30: 103. 1949. Varronia ferruginea Lam. Illust. 1: 418.
1791. Cordia riparia HBK. Nova Gen. & Sp. 3: 71, t. 207. 1818.
C. ferruginea (Lam.) Roem. & Schult. Syst. Veg. 4: 468. 1819.
C. crenulata A. DC. in DC. Prodr. 9: 492. 1845. Bejuco negro (Izabal
and Sacatepe"quez) ; kekxej (Alta Verapaz) .
Usually in damp thickets, ravines, along rivers or swamps, occa-
sionally in mixed or pine forests, sea level to 1,650 meters; Alta Vera- \
paz, Chiquimula, Escuintla, Guatemala, Huehuetenango, Izabal,
Peten, Quezaltenango, Retalhuleu, Sacatepequez, San Marcos, Santa
Rosa, Zacapa. Western and southern Mexico, British Honduras to
Panama; Colombia.
Erect or somewhat scandent shrubs, 2-4 m. tall, usually with recurved
branches, the stems and branches tomentose, hispidulous, or glabrate; leaves on
petioles 0 5-2 cm. long, the blades membranaceous, usually broadly ovate, some-
times oblong-ovate or lance-ovate, rarely lanceolate, 4-15 cm. long, acute or acumi-
nate rounded and abruptly decurrent at the base, usually serrate to subentire,
rarely entire, dark green above, scabrous or glabrate, paler beneath and sparsely
or densely ferruginous-pubescent, the hairs usually erect, rarely appressed; inflores-
cences spicate, the spikes solitary, chiefly axillary, usually as long as or longer than
the leaves, dense or interrupted, many-flowered; calyx campanulate, 2-4 mm. long,
5-6 dentate, usually not costate, or the costae obsolete, usually hispidulous, tl
hairs often underlain by minute papillae; corolla 2-4 mm. long, usually exceeding
the calyx by only 1 mm., rarely by 2-3 mm., greenish-white or cream, nearly ti
cate or the ruffled or laciniate margin giving the appearance of numerous, small,
uneven lobes or teeth; stamens exserted, the filaments inserted in throat of corolla,
glabrous to point of insertion and there surrounded by shaggy hairs which ring the
corolla throat; ovary glabrous; style included; fruits red, the drupes partially o
completely enclosed in the accrescent calyx.
Cordia stellifera I. M. Johnston, Journ. Arn. Arb. 21: 350. 1940.
Usually in wet forests, fields, thickets, or on slopes near rivers,
occasionally in open pine forest, 50-225 meters ; Alta Verapaz ; Izabal ;
Pet£n. Southern Mexico; British Honduras; Honduras; Costa Rica.
Trees, 6-12 m. tall, the young branches slender, minutely pubescent with
mostly appressed hairs or almost glabrous; leaves on petioles 0.5-1.5 cm. long, the
blades membranaceous, elliptic-oblong to broadly oblanceolate, usually 8-.
cm long 4-7.5 cm. wide, obtuse to acuminate, acute at the base, entire or nearly
so lustrous above, almost glabrous and smooth to the touch, glabrous beneath or
with scattered, minute, appressed hairs; inflorescences lax, long pedunculate, o
12 cm broad, the branches strigillose or glabrate; calyx subglobose in bud, densely
strigillose, open-campanulate at anthesis, 3 mm. long, the 4-5 lobes triangular,
acute- fruiting calyx spreading and stelliform, 6-9 mm. broad; corolla white
cream, 7 mm. long, the lobes broadly ovate; stamens villous near base; ovary
glabrous, style 3-5 mm. long; fruits depressed transverse-ovoid.
GIBSON: FLORA OF GUATEMALA 133
It may be that this species should be referred to C. laevitata Lam.
of the West Indies, which differs from C. nitida Vahl only in its
smaller leaves and puberulous calyx. This group of Cordias, includ-
ing C. eriostigma Pittier (Costa Rica and El Salvador), C. lasiocalyx
Pittier, C. lucidula Johnston, and C. prunifolia Johnston, are cer-
tainly very closely related and monographic study is needed.
Cordia truncatifolia Bartlett, Proc. Am. Acad. 43: 58. 1907.
C. microsebestena Loes. Verh. Bot. Ver Brandenb. 55: 187. 1913.
C. gualanensis Donn.-Sm. Bot. Gaz. 55: 438. 1913.
Dry hills and thickets, 100-650 meters; Chiquimula, El Progreso;
Zacapa. Mexico (Oaxaca and Guerrero) .
Shrubs or small trees, to 8 m. tall, the branches rather stout, puberulent or
glabrate; leaves membranaceous, on petioles 3-10 mm. long, the blades broadly
ovate, 2-7.5 cm. long and 2-5 cm. broad, abruptly acute at apex, rounded or trun-
cate at the base, entire, subentire, or coarsely crenate-dentate, scabrous on the
upper surface, softly pubescent or subtomentose beneath; inflorescences cymose,
the cymes dichotomous, usually 6-12-flowered, pedunculate, the pedicels 2-7 mm.
long; flowers of two forms: calyces of the small-flowered form ca. 6 mm. long at
anthesis, broadly campanulate, brownish-sericeous, the 5 costae obscured by indu-
I ment, the 5 (sometimes 6) lobes subulate, calyces of the large-flowered form 10 mm.
! long, campanulate, sericeous but with longer hairs along costae and near base of
j tube, the 5 lobes long-attenuate or laciniate, 4-5 mm. long, both forms apiculate
Jin bud and accrescent; corollas funnelform, white or yellowish white, puberulent,
I those of the small form 15-16 mm. long, those of the larger form 20-22 mm. long,
I the lobes of both obovate, ca. one-third as long as the tube; stamens included, the
(glabrous filaments arising near the base of the tube and adnate to it for almost
half their length; ovary ovoid, style elongating but always included, those of the
small form ca. 3-6 mm. long, the larger ones 8-11 mm. long; fruits subglobose,
the drupes orange-yellow, glabrous, ca. 1 cm. long, finally rupturing the calyx.
It is not yet known whether both forms are fertile, as the calyx
becomes greatly enlarged in fruit. The ovaries of both forms bear
minute ovules.
Both forms have been confused with C. seleriana Fernald, which
they closely resemble; however, the calyx of C. seleriana has only
three lobes, although two of these lobes may each bear two minute
teeth. Further, the calyx in bud is never apiculate, and the style
may be 15 mm. long.
CYNOGLOSSUM L.
Reference: M. Gtirke in Engler & Prantl, Pflanzenf. IV, 3a: 102.
1893.
134 FIELDIANA: BOTANY, VOLUME 24
Perennial or biennial herbs, usually branched, pubescent; leaves alternate, the
basal leaves usually long-petiolate, the cauline ones usually sessile; inflorescences
racemose, the cincinni short and densely flowered at first, usually elongating and
becoming lax and remotely flowered, mostly ebracteate, sparsely branched or form-
ing broad panicles; the flowers pedicellate or subsessile, the pedicels often recurved
in fruit; calyx deeply 5-lobed, little accrescent in fruit, spreading or reflexed; co-
rolla blue, violet, or rarely white, the tube short, the throat closed by small append-
ages, the 5 lobes imbricate, obtuse, spreading; stamens 5, included, inserted on the
corolla tube, the filaments short; anthers ovoid or short-oblong, obtuse; ovary
4-lobed, the style short or elongated, the small stigma plane or subcapitate; ovules
horizontal; nutlets 4, depressed, widely spreading in fruit, adnate to the gynobase
by almost the whole lower surface, dorsally convex, flat, or surrounded by a raised
border, glochidiate-muricate.
About 60 species, in temperate and subtropical regions of both
hemispheres, mostly in the Old World. Only a few are native in
North America; only one is known from Mexico (C. pringlei Greenm.),
and none from Guatemala; the following species is treated here as it
has been collected as an escape from cultivation.
Cynoglossum amabile Stapf & Drumm. Kew Bull. 1906: 202
1906.
Native of China; cultivated frequently for ornament in Guate-
mala, in parks and gardens; naturalized in Solold and Sacatepe"quez
Also naturalized in Mexico and in scattered localities throughoui
Central America, Ecuador and Colombia.
Branched, erect, perennial herbs to about 50 cm. tall, rather finely grayish-
pubescent throughout and somewhat hirsute; basal leaves petiolate, the blades
lance-elliptic, acute at each end, 5-10 cm. long, 2-3.5 cm. wide, the cauline leaves
sessile, often clasping, oblong or lanceolate, obtuse or acute, entire or undulate, the
lower ones to 15 cm. long, decreasing in size upward to 2 or 3 cm. long; the cymes
of inflorescence forming a large panicle, the pedicels in flower scarcely 2 mm. long,
in age 4 mm. long and reflexed; calyx 2-3 mm. long, the segments ovate, subacute;
corolla usually blue, sometimes white, the limb 5-10 mm. broad; nutlets ovate,
depressed dorsally, 3 mm. long when mature, whitish, densely glochidiate-muricate,
the spines very short, usually less than 1 mm. long.
Because of its easily dispersed nutlets which attach themselves
readily to passing objects, this plant, already plentiful in some of the
cafetales about Antigua, Sacatepe"quez, may well become a natural-
ized weed in the temperate regions of Guatemala.
EHRETIA L.
References: Linnaeus, Syst. ed. 10: 936. 1759; M. Giirke in Engler
& Prantl, Pflanzenf. IV, 3a: 87-88. 1897.
GIBSON: FLORA OF GUATEMALA 135
Trees or shrubs, glabrous or pubescent; leaves alternate, petiolate, the blades
frequently coriaceous, often scabrous, the margins entire, dentate, or serrate; in-
florescences panicles or corymbose cymes, terminal; calyx small, 5-parted, the seg-
ments imbricate or open in bud; corolla tube short or cylindric, the throat some-
times ampliate, the 5 lobes imbricate, obtuse, spreading or recurved; stamens 5,
usually exserted, the filaments inserted on the corolla tube, the anthers ovoid or
oblong; ovary 2-celled with each cell imperfectly 2-locellate, or the ovary 4-celled,
style terminal, bifid, the 2 stigmas terminal, clavate or capitate; ovules attached
laterally; fruit small, drupaceous, generally globose or subglobose, pale yellow,
orange, red, or purple, at maturity breaking into two 2-seeded or four 1-seeded
pyrenes; seeds usually straight; endosperm scant, the cotyledons ovate, not plicate.
Perhaps 50 species, in the tropics of both hemispheres, chiefly in
the Old World. Only two species are known to occur in Guatemala
with three more in Mexico and two others reported from southern
Central America.
Leaf blades entire; panicles usually longer than the leaves E. tinifolia.
Leaf blades dentate or serrate; panicles usually shorter than the leaves . . E, luxiana.
Ehretia luxiana Donn.-Sm., Bot. Gaz. 18: 5. 1893. Pepenance
(El Quiche").
Damp, mixed or pine forest, wet thickets, or brushy hillsides,
1,000-2,500 meters; Alta Verapaz; Guatemala; Quezaltenango; El
Quiche" ; Sacatepe"quez ; San Marcos. Southern Mexico; El Salvador.
Trees, usually 6-9 (rarely to 15) m. high, the young branches ferruginous, gla-
brous or sometimes with scattered hairs; leaves on petioles 1-2.5 cm. long, the
blades chartaceous to subcoriaceous, ovate to lance-oblong or elliptic-oblong,
mostly 6-15 cm. long, acute or acuminate, acute or rounded at the base, the mar-
gin serrate or dentate, essentially glabrous on both surfaces or sparsely hispidulous
beneath with some scattered, stiff hairs, especially along the costa; older leaves
sometimes becoming scabrous; panicles terminal, usually shorter than the leaves,
the stems nearly glabrous or puberulent with appressed hairs, densely flowered,
the flowers sessile or short-pedicellate; calyx 2-2.5 mm. long, glabrous or obscurely
puberulent, the lobes obtuse, ciliate; corolla white, 4-5 mm. long, glabrous, the
broad lobes spreading or reflexed; stamens long-exserted; dry fruits subglobose, to
1 cm. long, glabrous; fresh fruits juicy, white, oblong-ovoid, ca. 13 mm. long.
Ehretia latifolia A. DC. of central and southern Mexico, may be
separated from E. luxiana by its lanceolate, acute sepals, and its
usually smaller leaves (mostly 4-8 cm. long) which are always very
scabrous on the upper surface, and finely pubescent beneath.
Ehretia tinifolia L. Syst. Nat. ed. 10: 936. 1759.
Twice collected in the Department of Guatemala, Steyermark
b8083a, cultivated, and Aguilar 530, possibly also cultivated. Mex-
ico; Honduras; West Indies.
136
FIELDIANA: BOTANY, VOLUME 24
FIG. 24. Ehretia tinifolia. A, habit, Y^ natural size; B, flower, X 5; C, fruit ,
X 4; D, calyx with style and stigma, X 5; E, dissection showing pistil, X 5.
Trees or shrubs, essentially glabrous throughout, said to attain a height of
25 m. in some regions; leaves short-petiolate, the blades chartaceous, usually lus-
trous, oblong or elliptic-oblong to ovate, 4-13 cm. long, 2.5-5 cm. wide, usually
rounded, sometimes acute at the apex, obtuse or rounded at the base, margins
entire; panicles terminal, many-flowered, usually longer than the leaves, dense or
open; calyx 1.5-2 mm. long, glabrous, the lobes rounded, ciliate; corolla white,
3-5 mm. long, the lobes oblong, reflexed; stamens and style long-exserted ; fruits
subglobose to ovoid, 5-6 mm. in diameter, red or purple.
GIBSON: FLORA OF GUATEMALA 137
HACKELIA Opiz
Reference: A. Brand, Pflanzenreich IV. 252: 117-135. 1931.
Herbs, usually perennial, rarely biennial or annual, pubescent; leaves alter-
nate, entire or nearly so, at least the lower ones conspicuously petiolate; flowers
small, usually blue, the raceme-like inflorescences terminal and axillary, usually
forming a panicle, calyx deeply parted; corolla cylindric or funnelform to campan-
ulate or subrotate, the throat usually closed by large appendages; stamens included,
the short filaments attached to corolla tube below each sinus; style and stigma
simple; gynobase pyramidal, with 4 large triangular pits; nutlets 4, aculeate, with
a large central scar, often dissimilar.
About 40 species, mostly in temperate North America, two or
three in South America, one in Europe, a few in Asia. Only the
following are known in Central America.
Lower leaf surfaces very densely strigose or sericeous with closely appressed hairs;
spines of armed nutlets ca. 1.5 mm. long H. skutchii.
Lower leaf surfaces pubescent with spreading hairs or rather sparse, short hairs;
spines of armed nutlets 3-4 mm. long H. mexicana.
Hackelia mexicana (Schlecht. & Cham.) I. M. Johnston, Contr.
Gray Herb. 68: 46. 1923. Cynoglossum mexicanum Schlecht. &
Cham. Linnaea 5: 114. 1830. Echinospermum mexicanum Hemsl.
Biol. Cent. Am. Bot. 2: 377. 1882. Lappula mexicanum Greene, Pit-
tonia 2: 182. 1891. L. costaricensis Brand in Fedde, Rep. Sp. Nov.
18: 310. 1922. L. guatemalensis Brand, I.e. 311 (type from Huehue-
tenango, Seler & Seler 31 44) • Hackelia costaricensis (Brand) I. M.
Johnston, Contr. Gray Herb. 68: 46. 1923. H. guatemalensis Brand,
Pflanzenr. IV. 252: 120. 1931. H. stricta I. M. Johnston, Journ.
Arn. Arb. 29: 238. 1948. Mozote.
Damp or wet meadows, thickets, or mixed forest, sometimes in
pine or Cupressus forest, 1,400-3,500 meters; Chimaltenango; Hue-
huetenango; Jalapa; Quezaltenango; El Quiche"; San Marcos. Mex-
ico; Costa Rica; Panama; Ecuador and Venezuela.
Erect, usually coarse, annual, biennial or perennial herbs to 75 cm. tall,
branched, from a thick, sometimes repent rootstock, the stems sparsely or densely
pilose with spreading white hairs; basal leaves on very long, slender petioles to
23 cm. long, the blades broadly ovate, ovate, lance-ovate, or elliptic, 5-14 cm. long
and 2-5.5 cm. wide, the cauline leaves short-petiolate or the uppermost sessile,
sometimes clasping, the blades thin, elliptic to oblong-lanceolate, 2-12 cm. long,
acute, when petiolate narrowed below into the broad petiole, rather sparsely hir-
sute with short, mostly spreading whitish hairs; cincinni many-flowered, lax, brac-
teate, usually forming a large panicle, the flowers pedicellate, becoming remote in
age, the pedicels reflexed in age, longer than the calyx, almost 1 cm. long in fruit;
calyx at anthesis 1-2 mm. long, lobate for one-half to two-thirds its length, accres-
138 FIELDIANA: BOTANY, VOLUME 24
cent, 2-3 mm. long in fruit, pilosulous, the lobes ovate, subacute; corolla blue to
lavender, the tube 1-2 mm. long, the broad, rounded lobes 2-3 mm. long, the throat
yellow; style ca. 1 mm. long; nutlets ca. 3 mm. long (excluding the spines), usually
2 or 3 nutlets covered with glochidiate spines to 4 mm. long, the remaining ones
with much shorter spines, or rarely, one nutlet muricate, lacking glochidia, and
occasionally all 4 nutlets with spines essentially the same length.
It should be noted that fruits with dissimilar nutlets and fruits
with nutlets alike sometimes occur on the same plant, and, in fact,
the fruits on the type specimen of Lappula guatemalensis Brand
(Seler & Seler 3144} are variable, some fruits with nutlets that are
all alike, others with only three nutlets maturing, two of these alike
and the third much smaller with shorter spines.
Hackelia skutchii I. M. Johnston, Journ. Am. Arb. 29: 238.
1948.
In meadows and on rocky slopes in open coniferous forests, 2,800-
3,500 meters; Huehutenango. Apparently confined to the subalpine
regions of the Sierra de los Cuchumatanes.
Erect, perennial herbs to 55 cm. tall, the stems appressed-strigose; leaves ap-
pressed-strigose to strigillose on upper surface, appressed-strigillose to sericeous
beneath, usually densely so, the midrib prominent, basal leaves oblanceolate,
usually 6-14 cm. long, 0.5-1 cm. wide, acute, mucronulate, attenuate to the base
and often extending onto the petiole; cauline leaves narrowly oblanceolate to lan-
ceolate, acute, mucronulate, sessile and rounded at the base, 1-6 cm. long, 0.2-
0.6 mm. wide, decreasing in size upward; cincinni many-flowered, lax, bracteate,
usually forming an open panicle, the flowers pedicellate, the pedicels reflexing and
becoming remote in age, longer than the calyx, ca. 1 cm. long in fruit; calyx strigil-
lose, ca. 1 mm. long at anthesis, 2 mm. long in fruit, the lobes ovate; corolla blue,
tube ca. 2 mm. long, limb 7-10 mm. in diameter, the obovate lobes spreading;
style ca. 0.5 mm. long; nutlets 4, grey, dissimilar, 2-2.5 mm. long, usually 3 (2)
nutlets muricate-papillate, the protuberances usually less than 0.3 mm. long, glo-
chidiate or not, and the remaining one (or 2) with glochidiate spines to 1.5 mm.
long.
The nutlets of H. skutchii are very similar to those of the Mexican
H. leonotis I. M. Johnston. However, H. leonotis has a much smaller
corolla (tube 1 mm. long, limb 2.5-4 mm. in diameter) and the indu-
ment of the stem and surface of the lower leaves is loose and some-
what shaggy.
HELIOTROPIUM L.
References: Ivan M. Johnston, The South American species of
Heliotropium, Contr. Gray Herb. 81: 3-73. 1928; The Boragina-
ceae of northeastern South America, Journ. Arn. Arb. 16: 1-64.
1935; Studies in the Boraginaceae, XII, Journ. Arn. Arb. 18: 1-
FIG. 25. Hackelia skutchii. A, habit, H natural size; B, dissection of calyx
showing pistil, X 5; C, accrescent calyx with mature nutlets, X 5; D, corolla dis-
sected to show stamens and faucal appendages, X 5; E, pubescence from stem
and leaves, much enlarged.
139
140 FIELDIANA: BOTANY, VOLUME 24
25. 1937; Studies in the Boraginaceae, XVIII, Journ. Arn. Arb. 30:
133-138. 1949.
Annual or perennial herbs, or sometimes low shrubs, usually pubescent, some-
times glabrous; leaves simple, alternate to subopposite; inflorescences usually of
solitary, geminate or ternate bostryces, occasionally dichotomous, or the flow-
ers solitary, cauline and internodal; calyx usually persistent but sometimes de-
ciduous, with 5 teeth or lobes, the sepals often unequal in length and/or width;
corolla small, usually white, sometimes blue, lavender or purple, rarely yellow,
salverform or funnelform, 5-lobed, the tube cylindric, the limb spreading; stamens
5, borne on the corolla tube, included, the filaments short; anthers ovate to oblong
or lanceolate, obtuse, mucronate, or short-appendaged; ovary 4-celled, often 4-
lobate, the style terminal, the stigma sessile or borne on a distinct style, annular
and peltate, turbinate, umbonate, or conic, the apex usually bearing a conic or
cylindric or elongated sterile appendage, this often bifid; fruits dry, greenish to
brown, sometimes lobate, at maturity separating into 2-4 bony nutlets; nutlets
1-2-seeded, often with 1-2 sterile cavities; endosperm thin; cotyledons flat.
A widely dispersed and diversified genus with perhaps 150 species
in both hemispheres, chiefly in tropical regions, a few in temperate
areas. In addition to the 14 species native to Guatemala, a Peruvian
species, H. arborescens L., is cultivated in Guatemalan gardens at
almost all elevations. It is grown because of the fragrance of its
small, purple flowers, rather than for its value as an ornamental.
It is sometimes difficult to separate suffruticose plants of Helio-
tropium with furcate inflorescences (H. macrostachyum (DC.) Hemsl.
and H. rufipilum (Benth.) I. M. Johnston) from Tournefortia, when
fruiting material is not available.
Plants glabrous, the leaves succulent H. curassavicum.
Plants pubescent, the leaves not succulent.
Flowers borne on ebracteate bostryces, these often dichotomous.
Leaves densely white-strigose or sericeous beneath, the. hairs all appressed.
Plants shrubby or suffruticose; veins strongly impressed on upper leaf
surfaces; leaf blades usually acute; corolla glabrous in throat . . . H. fallax.
Plants herbaceous; veins not impressed on upper leaf surfaces; leaf blades
usually obtuse; corolla villous in throat H. procumbens.
Leaves green beneath, pubescent with spreading hairs or glabrate.
Leaf blades broadly ovate or deltoid-ovate, conspicuously undulate.
Corollas white, 9-10 mm. long; inflorescence forked; fruits separating
into 2 nutlets . . . H. macrostachyum.
Corollas normally pale blue or lavender, 4-5 mm. long; inflorescence
simple; fruits separating into 4 nutlets H. indicum.
Leaf blades lanceolate to elliptic-oblong, not undulate.
Inflorescence of 2-3 times dichotomous cymes; calyx lobes accrescent,
enclosing the fruits H. rufipilum.
Inflorescence of solitary or geminate, elongating cincinni; calyx lobes not
accrescent, never enclosing the fruits.
GIBSON: FLORA OF GUATEMALA 141
Calyx 1-2 mm. long; indument of spreading, unequal hairs; fruits
glabrous, minutely tuberculate H. angiospermum.
Calyx 3-4 mm. long; indument of largely malpighiaceous hairs; fruits
puberulent, never tuberculate H. transalpinum.
Flowers borne in leaf axils, or if on bostryces, these bracteate, simple, solitary,
never dichotomous.
Plants with stems prostrate or decumbent.
Flowers borne in or above the leaf axils.
Leaf blades 2 mm. wide or less; fruiting calyx 2.5 mm. long or less; fruits
beaked, smooth H. lagoense.
Leaf blades 3-7.5 mm. wide; fruiting calyx 4-5 mm. long; fruits not
beaked, minutely reticulate- veined H. pringlei.
Flowers borne on terminal, short, usually dense cincinni . . H. foliosissimum.
Plants with stems erect or ascending.
Bracts of inflorescence large and leaf-like H. fruticosum.
Bracts of inflorescence small, inconspicuous, often subulate.
Calyx 3-4 mm. long H. ternatum.
Calyx 0.8-1.5 mm. long H. filiforme.
Heliotropium angiospermum Murray, Prodr. Stirp. Goett.
217. 1770. H. parviflorum L. Mant. PL 2: 201. 1771. Heliophytum
parviflorum DC. Prodr. 9: 553. 1845. Cola de alacran (Guatemala);
cola de mico (Pete"n).
Wet or dry thickets, brushy fields, hillsides, sandbars, hedgerows
or waste ground, 50-1,140 meters; Chimaltenango; Chiquimula;
Guatemala; Izabal; Jalapa; Pete"n; Zacapa. Southern Mexico; Brit-
ish Honduras to El Salvador and Costa Rica; West Indies; South
America.
Erect herbs, annual or perennial, to 1.5 m. tall, branched, the branches slender,
more or less hirsute with spreading, unequal hairs; leaves on slender petioles 5-
15 mm. long, often subopposite, the blades usually oblong-lanceolate or oblong-
elliptic, sometimes ovate to broadly lanceolate, 2-10 cm. long, acute or short
acuminate, usually acute at the base, sometimes rounded and decurrent, entire,
usually thinly hirsute on both surfaces with 7-8 pairs of lateral veins; inflorescence
of slender, pedunculate, ebracteate, solitary or geminate bostryces 6-22 cm. long,
the flowers sessile, usually crowded; sepals lanceolate to oblong, obtuse or acute,
1-2 mm. long; corolla white, 2 mm. long, barbate within throat; stamens sub-
sessile, inserted on tube at a point just above the middle of the tube, anthers
0.8 mm. long; stigma sessile, umbonate; ovary 0.2 mm. long; fruits didymous, de-
pressed, 1-1.5 mm. long, minutely tuberculate, the nutlets united in pairs.
This is a common, weedy plant in many parts of the Central
American lowlands.
Heliotropium curassavicum L. Sp. PL 130. 1753; Johnston,
Contr. Gray Herb. 81: 14. 1928.
142 FIELDIANA: BOTANY, VOLUME 24
Saline or alkaline flats, plains and meadows, usually along sea-
shores, sea level to 600 meters; Escuintla; Guatemala; Jutiapa;
Retalhuleu; San Marcos. Southern United States; Mexico to Pan-
ama; West Indies; northern and western South America.
Plants annual or perennial, glabrous, succulent, bluish green and often glau-
cous, usually diffusely branched, the stems prostrate or decumbent, sometimes to
50 cm. long or more; leaves sessile, the blades linear, linear- oblong, narrowly ob-
lanceolate or spathulate, entire, obtuse, attenuate to the base, 1-5 cm. long; inflo-
rescence of ebracteate bostryces, 1-10 cm. long, solitary or geminate; the small
flowers subsessile or on pedicels 1-2 mm. long; calyx 1-2 mm. long, the lobes tri-
angular-ovate or oblong, acute or obtuse, usually shorter than the mature fruits;
corolla white, 1-3 mm. long, the tube about equalling the sepals; anthers sagittate,
0.5-1 mm. long, sessile on tube; stigma broad, cap-like, sessile; fruits subglobose,
separating into 4 oblong nutlets.
In Guatemala the plant occurs principally along the Pacific coast,
where it is often abundant.
Heliotropium fallax I. M. Johnston, Journ. Arn. Arb. 18: 14.
1937. Hierba de toro (Huehuetenango) .
Dry, open, brushy or forested, often rocky mountain slopes, fre-
quently on limestone, 800-1,950 meters; Baja Verapaz; Jalapa; El
Quiche" ; Huehuetenango. Southern Mexico.
Stout, erect shrubby or suffruticose plants, usually ca. 50 cm. tall but some-
times attaining 120 cm., branched, the branches densely tomentose or hirsute with
long, white hairs; leaves subsessile or on petioles to 1 cm. long, the blades elliptic
to oblong-lanceolate or ovate-lanceolate, 1-5 cm. long, 0.5-2 cm. wide, usually
acute, rarely obtuse, obtuse at the base and often abruptly contracted into the peti-
ole, green or gray, densely white-hirsute or strigose above, the veins impressed,
densely white-tomentose or strigose beneath; inflorescences terminal, pedunculate,
ebracteate, often geminate bostryces 3-7 cm. long; pedicels to 1 mm. long; calyx
white-strigose, the lobes narrowly or broadly lanceolate, acute or acuminate, 2-
3 mm. long, surpassing the fruits; corolla white, 4-6.5 mm. long, the tube surpass-
ing the sepals, the lobes triangular to lanceolate, acute, strigose outside, glabrous
within, the limb 3-4 mm. broad; stamens included, subsessile, inserted on tube at a
point about midway from its base; the anthers 1.6 mm. long, coherent at their
apices; style ca. 0.1 mm. long, stigma annular, the appendage elongated 1-1.3 mm.;
fruits subglobose, whitish-strigose, 1.5 mm. long, 2.5 mm. in diameter, separating
into 4 nutlets.
This has been reported from Guatemala as H. coriaceum Lehm.
Heliotropium filiforme Lehmann, Goett. Gel. Anzeig. 1817:
1515. 1817; HBK. Nova Gen. & Sp. 3: 86 and 451, t. 204. 1818; DC.
Prodr. 9: 545. 1845; I. M. Johnston, Contr. Gray Herb. 81: 61. 1928.
GIBSON: FLORA OF GUATEMALA 143
Open places, savannas, near sea level; Pet£n. Southern Mexico;
British Honduras; Nicaragua; West Indies (fide Johnston); tropical
South America.
Slender, erect, branching annuals, 10-40 cm. tall, the stems and branches
white-strigose; leaves on petioles 1-3 mm. long, the blades oblanceolate or linear-
oblanceolate, obtuse or subacute, acute or attenuate to the base, 8-20 mm. long,
1.5-3.5 mm. wide, strigose; inflorescences of solitary, slender bostryces, elongating
in age, usually 2-15 cm. long, bearing minute, lanceolate to linear-lanceolate, often
subulate bracts and numerous flowers on pedicels 1 mm. long or less; calyx 0.8-
1.5 mm. long, the sepals lanceolate or linear-lanceolate, strigose, usually very un-
equal, surpassing the fuits; corolla white, 1-2 mm. long, the limb 1.5-2 mm. in
diameter, the tube not surpassing the sepals; stamens sessile on tube, included,
not at all coherent; stigma subsessile, ca. 0.3 mm. long; fruits subglobose, obscurely
4-lobate, strigose, separating into 4 nutlets ca. 1 mm. long.
A Nicaraguan specimen, F. E. Schramm (1926), has considerably
longer, nearly shaggy hairs on leaves and stems, appearing almost
velutinous rather than strigose.
Heliotropium foliosissimum Macbride, Proc. Am. Acad. 51:
541. 1916.
Open, grassy hillsides, about 2,000 meters; Huehuetenango.
Southern Mexico.
Plants perennial, much branched from the base, the stems slender, hirsute,
mostly 5-15 cm. long, prostrate, densely leafy; leaves short-petiolate, the blades
oblong-lanceolate or narrowly elliptic-oblong, 4-10 mm. long, 1.5-3.5 mm. wide,
obtuse or acute, acute at the base, entire, sometimes revolute, strigose and with
some longer, spreading hairs on margins and costae; inflorescences terminal, usu-
ally short and dense, the bracts large and foliaceous, the flowers pedicellate; calyx
3-3.5 mm. long, hirsute to pilose, the sepals oblanceolate to spatulate, acute or
obtuse, nearly twice as long as the fruits; corolla white, the tube about equalling
the calyx, the short lobes rounded, often somewhat crenate, the limb 3.5-5 mm.
broad; stamens included, subsessile, the anthers coherent at their apices; style
evident, 0.5-1 mm. long; fruits depressed globose, somewhat strigillose, separating
into 4 nutlets.
Heliotropium fruticosum L. Syst. ed. 10, 2: 913. 1759 and
Amoen. Acad. 5: 394. 1759; I. M. Johnston, Contr. Gray Herb. 81:
66. 1928. H. campechianum HBK. Nova Gen. & Sp. 3: 86. 1818.
H. phyllostachyum Torr. Bot. U. S. & Mex. Bound. 137. 1859.
H. phyllostachyum var. erectum Macbride, Proc. Am. Acad. 51: 542.
1916. H. assurgens I. M. Johnston, Journ. Arn. Arb. 20: 375. 1939.
Damp or dry, open or brushy, often rocky plains and hillsides,
200-675 meters; Zacapa. Southern Mexico; British Honduras; Hon-
FIG. 26. Heliotropium fruticosum. A, habit, natural size; B, flower, X 5;
C, accrescent calyx with fruit, X 5; D, style and stigma, X 15; E, pubescence,
much enlarged.
144
GIBSON: FLORA OF GUATEMALA 145
duras; El Salvador; Nicaragua; Costa Rica; West Indies; northern
South America.
Slender, erect or ascending annuals, 8-30 cm. tall, the branches strigose or
appressed-pilose, usually leafy; leaves on petioles 1-4 mm. long, the blades small,
0.5-2.5 cm. long, usually 0.4-0.6 cm. wide, strigose on both surfaces, usually more
densely so beneath, linear-oblong to oblong-lanceolate, lanceolate, or oblong-ellip-
tic, apex usually acute, rarely obtuse, acute or attenuate to the base, the margins
entire, often somewhat revolute; inflorescences of slender, elongating bostryces bear-
ing scattered, large leaf-like bracts, the flowers sessile or on pedicels ca. 1 mm. long,
crowded at first but becoming remote (from 2 or 3 to as much as 10 mm. apart);
sepals lance-ovate to lanceolate, acute or subacute, strigose, unequal, 1.5-3 mm.
long, surpassing the fruits; corolla white, 2.5-3.5 mm. long, the limb 3-4 mm.
across, the lobes broadly lanceolate, the tube equalling or exceeding the sepals at
an thesis; stamens included, subsessile, inserted on lower half of tube, the anthers
coherent at apices; style evident, at an thesis as long as or longer than the ovary,
often persistent on the fruit, with the stigmatic disc and appendage still attached;
fruits 1.5-2 mm. in diameter, subglobose, strigose, separating into 4 nutlets about
1 mm. long.
Heliotropium indicum L. Sp. PL 130. 1753. Cola de alacrdn
(Retalhuleu, Sacatepe'quez) ; cola de zope (Santa Rosa); giieguecho
(Jutiapa) .
Usually in damp or wet thickets or fields, sandy river beds, some-
times on dry wasteland and along railways and roadsides, sea level to
1,200 meters; Alta Verapaz; Escuintla; Guatemala; Huehuetenango;
Izabal; Jutiapa; Pete"n; El Progreso; Retalhuleu; Sacatepe'quez ; San
Marcos; Santa Rosa; Zacapa. Florida; Mexico; British Honduras to
Panama; West Indies; South America; naturalized in the Old World
tropics.
Coarse, erect annuals to 1.5 m. tall, the stems stout, branched, usually hispid
with spreading whitish hairs; leaves on petioles 1-6 cm. long, the blades usually
broadly ovate to deltoid-ovate, sometimes lanceolate, 4-15 cm. long, acute or ob-
tuse, mostly rounded or subcordate at the base and narrowly decurrent, repand or
undulate, hispid and sericeous or variously pubescent; inflorescences of long, ebrac-
teate, mostly solitary, simple bostryces to 28 cm. long, the flowers sessile, crowded
or remote; sepals linear-lanceolate, acute, 1.5-3 mm. long, sparsely hispid with
long hairs; corolla pale blue or lavender, the tube 3-4 mm. long, more or less stri-
gose outside, the lobes rounded, less than 1 mm. long; style short but evident,
ca. 0.5 mm. long, stigma depressed-conic, ca. 0.5 mm. long, ovary 0.8 mm. long,
stamens to 1 mm. long, subsessile on tube at a point below the middle of tube,
fruits deeply bilobate, mitriform, 2-3 mm. long, glabrous, finally breaking into 4
angulate, beaked nutlets.
Heliotropium lagoense (Warm.) Giirke in Engler & Prantl,
Pflanzenf. IV, 3a: 97. 1893. Schleidenia lagoensis Warm. Kjoeb.
146 FIELDIANA: BOTANY, VOLUME 24
Vidensk. Meddel 1867: 15. 1868. H. antillanum Urban, Symb. Ant.
4: 528. 1910. H. trinitense Urban, Symb. Ant. 7: 350. 1912.
Damp fields or savannas, 200 meters or less; Pete"n. Panama:
Cuba and Puerto Rico; Trinidad; South America.
Plants annual or perennial, much branched from the base, the stems prostrate
and often forming mats, sparsely appressed-hirsute; leaves small and numerous,
mostly 3-8 mm. long, 1-2 mm. wide, lanceolate, linear-lanceolate, or lance-oblong,
petiolate, obtuse or acute, acute at the base, entire, sparsely pilose or almost gla-
brous; flowers solitary in or above the leaf axils, on pedicels 2-4 (-6) mm. long;
sepals lanceolate or linear-lanceolate, unequal in width and in length, acute,
sparsely hirsute to pilose, sometimes minutely papillose, ca. 1.5 mm. long at an-
thesis, 2-2.5 mm. long in fruit; corolla white, ca. 3 mm. long, the lobes 1 mm. long,
rounded ; stamens included, subsessile on tube, the anthers coherent at their apices;
stigma subsessile; fruits glabrous or sparsely strigillose, separating into 4 beaked
nutlets, almost equalling the sepals in length.
Heliotropium macrostachyum (DC.) Hemsl. Biol. Centr.
Amer. Bot. 2: 375. 1882. Heliophytum macrostachyum DC. Prodr.
9: 556. 1845. Cola de mico (Pet&i).
Brushy, rocky slopes and bordering lakes, 400-1,465 meters;
Chiquimula; Guatemala; Pete'n; Zacapa. Mexico; Honduras; Nica-
ragua.
Erect herbs about 1 m. tall, the stout branches densely pilose with soft, yellow-
ish, spreading to somewhat reflex ed, more or less viscid hairs; leaves on petioles
1-5 cm. long, the blades usually ovate to rounded-ovate, rarely lance-ovate, mostly
6-12 cm. long, rarely to 20 cm. long, 4-10 cm. wide, acute to acuminate, usually
cordate to obtuse at the base, sometimes decurrent, undulate, sparsely or densely
pubescent on both surfaces with spreading, more or less viscid hairs; inflorescences
pedunculate, forked, the 2 branches 12-33 cm. long, helicoid, many-flowered, the
flowers crowded or remote, sessile; sepals linear- lanceolate, acuminate, hispidulous,
2-3 mm. long; corolla white, 9-10 mm. long, pilosulous outside, the lobes rounded,
1-1.5 mm. long; stamens included, the anthers subsessile at a point just above the
middle of the corolla tube; style ca. 1 mm. long, stigma depressed-discoid; ovary
4-celled, but 2 cells empty; fruit depressed-globose, 3 mm. long, 4 mm. broad,
pilosulous, separating into 2 nutlets.
This species is much like Tournefortia in general appearance and
has been confused with that genus in herbaria, especially when the
specimens are not fruiting or bear immature fruits. It has also been
confused with H. indicum L., although the inflorescences of H. indi-
cum are never forked, its corollas are much shorter, and its fruits
separate into four nutlets.
Heliotropium pringlei Robins., Proc. Am. Acad. 26: 170. 1891.
Dry mountain slopes, 800-1,600 meters; Huehuetenango. Mexico.
GIBSON: FLORA OF GUATEMALA 147
Slender annuals, prostrate or spreading, much branched, the stems 3-12 cm.
long, rather densely hirsute with long, spreading white hairs; leaves short-petiolate,
the blades elliptic to elliptic-lanceolate or oblong lanceolate, sometimes nearly
ovate, 7-15 mm. long, 3-7.5 mm. wide, acute or obtuse, acute at the base, thinly
or densely white-hirsute; flowers short-pedicellate, borne in or above the leaf axils;
sepals lanceolate, acute to acuminate, 2-2.5 mm. long at an thesis, in fruit 4-5 mm.
long, ca. 2 mm. longer than the fruits; corolla white, 4-5 mm. long, the tube short,
the lobes very short, triangular, the limb 3.5-5 mm. broad; stamens included, sub-
sessile below throat of tube, anthers coherent at their apices in early anthesis but
separating later; style evident, at anthesis nearly as long as the ovary; fruits sub-
globose, minutely reticulate-veined, glabrate, separating into 4 nutlets.
Heliotropium procumbens Mill. Gard. Diet. ed. 8, 10. 1768;
Johnston, Contr. Gray Herb. 81: 52. 1928. H. americanum Mill.
Gard. Diet. ed. 8, 11. 1768; Johnston, Contr. Gray Herb. 92: 89.
1930. H. inundatum Swartz, Prodr. Veg. Ind. Occ. 40. 1788; DC.
Prodr. 9: 539. 1845. H. inundatum var. cubense DC. Prodr. 9: 540.
1845. H. rigidulum DC. I.e. H. eggersii Urban, Symb. Ant. 5: 481.
1908. Madre de alacrdn (Guatemala); rabillo (Pete"n).
Wet to dry thickets, fields, mud flats, waste land, or sandbars
along streams, sea level to 1,480 meters; Alta Verapaz; Guatemala;
Izabal; Jutiapa; Pete"n; El Quiche"; Retalhuleu; Santa Rosa. South-
ern United States; Mexico; El Salvador to Panama; West Indies;
South America.
Plants usually annual, usually much branched from the base, decumbent,
spreading, or ascending (rarely erect), strigose or appressed-pilose almost through-
out, the hairs white or grayish, the stems 10-50 cm. long, branched; leaves on peti-
oles 3-18 mm. long (commonly 4-10 mm.), sometimes with 2 or 3 smaller leaves
in the axil of an older leaf, the blades mostly 1-4 cm. long and 0.5-2 cm. wide,
usually thin and soft, appressed-pilose to strigose, elliptic to obovate or broadly
oblanceolate or (rarely) nearly linear, usually obtuse or rounded at the apex, rarely
acute, sometimes minutely apiculate, acute to broadly cuneate at the base, entire,
rarely revolute; inflorescences of mostly geminate or ternate, ebracteate, peduncu-
late bostryces, 2-10 cm. long, the flowers sessile or on pedicels ca. 1 mm. long; sepals
unequal, lanceolate or linear, 0.5-1 mm. long at anthesis, to 2.5 mm. long in fruit,
exceeding the fruits; corolla white, villous in throat, 1-1.5 mm. long, the limb 1-
1.4 mm. in diameter; stamens included, subsessile on lower half of tube, the anthers
not coherent at their apices; stigma sessile on the ovary; fruits subglobose, 4-lobate,
strigillose, separating into 4 nutlets ca. 1 mm. long.
A variable and weedy species often found where water has stood
during the rainy season.
Heliotropium rufipilum (Benth.) I. M. Johnston, Contr. Gray
Herb. 81: 44. 1928. Tournefortia rufipila Benth. Bot. Voy. Sulphur:
140. 1844. H. physocalycinum Donn.-Sm. Bot. Gaz. 49: 457. 1910.
148 FIELDIANA: BOTANY, VOLUME 24
H. jalicense Macbr. Proc. Am. Acad. 51: 542. 1916. Cola de alacran
(Guatemala and Retalhuleu) ; cola de mico (Pete"n) ; hierba de alacran
(Chiquimula and Jalapa) ; pie de shara (Chimaltenango, fide Mo-
rales) ; pie de gallo.
Damp or wet thickets, fields and slopes, along roadsides and in
coffee plantations, 600-2,400 meters; Alta Verapaz; Chimaltenango;
Chiquimula; Escuintla; Guatemala; Jalapa; Jutiapa; Pete"n; Retal-
huleu; Sacatepe"quez; Santa Rosa. Southern Mexico; El Salvador;
Honduras; Nicaragua; Ecuador; Peru; Bolivia.
Erect herbaceous or suffruticose plants to 2 m. tall, slender, branched, densely
hispid and hirsute throughout with spreading brownish, often viscid or glandular
hairs; leaves on petioles 0.5-2 cm. long, the blades lanceolate or oblong-lanceolate,
acute or acuminate, attenuate to the base, entire, 4-14 cm. long; inflorescences of
2-3 times dichotomous cymes, the peduncles slender, the bostryces usually densely
flowered, 5-15 cm. long, ebracteate, the flowers sessile; calyx at anthesis 2.5-3 mm.
long, the sepals lanceolate to lance-ovate, often subulate, the mid-vein sometimes
prominent, the calyx in fruit accrescent, assuming a pentagonal, subglobose form
enveloping the fruit, the segments then broadly ovate, cuspidate-acuminate, ca.
4 mm. long; corolla white, glandular-puberulent outside, the tube 2-3 mm. long,
the lobes semiorbicular, 1 mm. long or less; stamens included, the anthers sub-
sessile on the tube, about midway between its base and throat; style evident but
less than 0.5 mm. long; fruits glandular-puberulent, subglobose, separating into 4
nutlets ca. 1.5 mm. long.
Heliotropium ternatum Vahl, Symb. Bot. 3: 21. 1794; I. M.
Johnston, Journ. Arn. Arb. 30: 135. 1949. H. oaxacanum DC. Prodr.
9: 543. 1845. H. mexicanum Greenm., Proc. Am. Acad. 33: 484.
1898. Hierba de loro (Jutiapa) ; lorito bianco and pie de loro (Gua-
temala) .
Usually dry, often rocky, brushy or open slopes, fields, hillsides,
sometimes in pine-oak forests, rarely on sand bars, along streams or
swampy meadows, 85-1,900 meters; Baja Verapaz; Chiquimula;
Guatemala; Huehuetenango; Izabal; Jalapa; Jutiapa; El Quiche";
Santa Rosa; Zacapa. Mexico to Costa Rica; Colombia; Venezuela.
A stout, erect shrub to 6 cm. tall, or sometimes merely suffrutescent, branched,
the branches densely strigose with white or grayish hairs; leaves alternate or sub-
verticillate, sessile or on very short petioles, the blades lanceolate, linear-lanceolate
or linear, mostly 1.5-3 cm. long, 0.2-0.8 cm. wide, usually acute to acuminate,
rarely obtuse, acute at the base, subcoriaceous, revolute-margined, sparsely or
densely white-strigose on both surfaces, sometimes pustulate or rarely only pitted or
papillose on the upper surface; inflorescences of many-flowered bostryces, 2-15 cm.
long, bearing small, linear-lanceolate, sometimes subulate bracts, the flowers dense
or remote, sessile or nearly so; calyx 3-4 mm. long, white-strigose, the sepals lanceo-
late or ovate-lanceolate, acute, longer than the fruits; corolla white, strigose, the
GIBSON: FLORA OF GUATEMALA 149
tube equalling or exceeding the calyx, the limb 3-4 mm. broad; stamens included,
the anthers coherent at their apices; style evident; fruits subglobose, separating
into 4 strigose nutlets.
This plant has been known most commonly as H. fruticosum L.,
a name which has been found to apply properly to a quite different
plant, treated on a preceding page.
Heliotropium transalpinum Veil. Fl. Flum. 68. 1825 and
Icones 2, t. 40. 1827. H. tiaridioides Cham. var. schizocarpum I. M.
Johnston, Contr. Gray Herb. 81: 7. 1928.
Brushy slopes, thickets and along roadsides, 75-800 meters; Chi-
quimula; Izabal. Honduras; Costa Rica; Argentina; Bolivia; Brazil;
Ecuador; Paraguay.
Erect, shrubby or suffruticose plants, 1-2 m. tall, branching, more or less
pubescent throughout with largely malpighiaceous hairs; leaves often subopposite,
on petioles 0.5-1 cm. long, the blades entire, broadly lanceolate to oblong-elliptic,
with 7-10 pairs of primary lateral veins, acute or acuminate, acute or attenuate to
the base, 4-15 cm. long, upper surface usually glabrescent but may be somewhat
rough to the touch, lower surface with scattered pubescence, heaviest along the
costae; inflorescence of elongated, simple, pedunculate, ebracteate, usually solitary
bostryces to 20 cm. long, the flowers sessile, crowded or remote; sepals linear or
linear-lanceolate, acuminate, 3-4 mm. long, usually equalling or surpassing the
corolla tube, more or less strigillose; corolla white, 3-4 mm. long, strigillose out-
side; stamens subsessile on tube, inserted at a point below the middle of the tube;
anthers ca. 1 mm. long; stigma sessile, conic; ovary ca. 1 mm. long; fruits puberu-
lent, depressed-globose, bilobate, the carpels sulcate, breaking first into two parts
and finally into 4 nutlets.
Superficially much like H. angiospermum Murray, but readily
separated by the malpighiaceous hairs, longer calyx lobes and the
smooth, puberulent fruits.
LITHOSPERMUM L.
References: I. M. Johnston, Lithospermum, Contr. Gray Herb. 70:
18-31. 1924; A. Brand, Die amerikanischen Arten der Gattung
Lithospermum in Fedde, Rep. Sp. Nov. 28: 10-17. 1930; I. M. John-
ston, A survey of the genus Lithospermum, Journ. Arn. Arb. 33: 299-
362. 1952.
Mostly perennial herbs with rootstocks, usually with rough pubescence through-
out; leaves alternate, entire; inflorescences of leafy or bracteate cincinni, these
often elongating in age so that the flowers appear solitary in axils of leaves or
bracts; calyx usually 5-parted, the sepals linear; corolla usually yellow or white
in ours, small or large, in some species quite variable in size, the later flowers often
much smaller and cleistogamous, funnelform or salverform, the tube straight, the
150 FIELDIANA: BOTANY, VOLUME 24
throat often ampliate, sometimes gibbous or plicate, with or without intruding
appendages, the 5 lobes imbricate, obtuse, spreading at an thesis; stamens 5, in-
cluded, the short filaments inserted on the tube, anthers oblong, obtuse or apicu-
late; ovary 4-lobed, seated on a flat gynobase; style filiform or stout, the 2 stigmas
usually distinct; ovules erect; nutlets 4 or fewer by abortion, erect, ovoid, often
very hard and bone-like, smooth or rugose; seeds straight, the cotyledons flat.
About 40 species, chiefly in temperate regions of the northern
hemisphere, in the tropics confined to the mountains.
Only the following four species are known from Central America.
Corollas white, appendaged within throat.
Lateral veins of leaves conspicuous; calyx in fruit 6-9 mm. long, 2-3 times longer
than the nutlets L. mediale.
Lateral veins of leaves essentially obsolete; calyx in fruit 3-4.5 mm. long, little
longer than the nutlets L. distichum.
Corollas yellow, not appendaged within throat.
Corolla tubes usually 5-16 mm. long (in cleistogamous flowers only 1-3 mm.),
the lobes conspicuously erose-dentate or crisped; nutlets smooth, white.
L. calycosum.
Corolla tubes 18-25 mm. long, the lobes essentially entire; nutlets sulcate, gray-
ish-white or tawny L. guatemalense.
Lithospermum calycosum (Macbride) I. M. Johnston, Contr.
Gray Herb. 70: 30. 1924. L. strictum var. calycosum Macbride, Contr.
Gray Herb. 48: 56. 1916. L. obtusifolium I. M. Johnston, Contr. Gray
Herb. 70: 27. 1924, new synonym. L. galeottii Brand in Fedde, Rep.
Sp. Nov. 28: 17. 1930.
Open, grassy slopes, 1,350-2,800 meters; Huehuetenango. Mexico.
Coarse, erect perennials from thick, woody roots, the stems usually several,
stout, simple or branched, strigose and hispid, sparsely or densely leafy; basal
leaves attenuate to the base, oblanceolate, to 6 cm. long, the cauline leaves sessile,
linear-oblong to narrowly oblong, spatulate or oblong-lanceolate, usually obtuse,
sometimes acute, usually obtuse at the base, strigose, often hispid, the lower ones
often pustulate, the midvein usually prominent; inflorescences of simple or furcate,
leafy cincinni, short at anthesis but elongating to as much as 20 cm. in fruit, the
pedicels 1-5 mm. long; calyx 6-8 mm. long at anthesis, accrescent to as much as
15 mm. in fruit, the sepals linear; corolla lemon-yellow, variable in size, the tube
of early flowers often 10-16 mm. long, but the later ones sometimes only 1-3 mm.
long and cleistogamous, pubescent outside, not appendaged within, the lobes 1.5-
2 mm. long, the margins erose-dentate to erose-lobulate or crisped; style usually
about 7 mm. long; anthers subsessile just below throat of tube; nutlets 4 or fewer,
ovoid or broadly ovoid, 3-3.5 mm. long, white, lustrous, smooth.
Lithospermum distichum Ortega, Hort. Matr. Dec. 1:8. 1797;
Johnston, Journ. Arn. Arb. 33: 361. 1952. L. spathulatum Mart. &
Gal. Bull. Acad. Brux. 11: 337. 1844. Spakul (Huehuetenango).
GIBSON: FLORA OF GUATEMALA 151
Usually on grassy slopes, open hillsides and alpine meadows,
sometimes with Pinus rudis, 2,500-3,750 meters; Huehuetenango;
Quezaltenango; Sacatepe"quez. Mexico.
Perennials from woody, vertical roots, simple or sparsely branched, the stems
erect or decumbent, 10-30 cm. tall, densely strigose; leaves numerous, the basal
ones oblong-oblanceolate, attenuate to the base, to 8 cm. long, the cauline ones
usually oblong or oblanceolate, sometimes lanceolate, usually obtuse at the apex,
acute or attenuate to the base or broad and somewhat clasping, minutely strigose
on both surfaces, the lateral veins obsolete or nearly so; inflorescences of several
terminal cincinni, the flowers few or many, pedicellate, the branches in age elon-
gating and the flowers then remote; calyx at anthesis 2-3 mm. long, in fruit 3-
4.5 mm. long; corolla white, the length varible, usually 5-7 mm. long, rarely to
10 mm. long, puberulent outside, glabrous within, the faucal appendages well de-
veloped; anthers subsessile on the tube, inserted at a point 1-1.5 mm. below the
throat; style 2-3 mm. long; nutlets 2-2.5 mm. long, ovoid or broadly ovoid, white,
lustrous, smooth.
Lithospermum guatemalense Donn.-Sm., Bot. Gaz. 27: 436.
1899.
At 1,400-1,800 meters; Huehuetenango (type from Trinidad,
Seler & Seler 3082} ; also collected between Chacula and Canquintic.
Mexico (Chiapas).
Stout, erect perennials to 45 cm. tall, from thick, woody roots, the stems sev-
eral, simple below the inflorescence, densely leafy, densely hirsute or hispid with
white, more or less spreading hairs; basal leaves not seen, cauline leaves sessile,
linear-lanceolate to lance-oblong, strongly ascending or suberect, acute, obtuse or
acute at the base, hispid, the veins conspicuous; cincinni few-flowered, densely
leafy, the flowers short-pedicellate; calyx at anthesis 7-9 mm. long, accrescent,
10-12 mm. long in fruit, the sepals linear, hispidulous; corolla yellow, the tubes
18-25 mm. long, pubescent outside, not appendaged within, the broad lobes 2.5-
3 mm. long; anthers subsessile, inserted on tube at a point 2-3 mm. below the
throat; style filiform, exserted 1-2 mm.; nutlets ovoid, 3.5-4 mm. long, grayish-
white or tawny, lustrous, conspicuously sulcate and sparsely pitted.
Lithospermum mediale I. M. Johnston, Contr. Gray Herb.
70: 28. 1924. L. discolor var. subviride Kuntze, Rev. Gen. 2: 439.
1891. L. colombianum Brand in Fedde, Rep. Sp. Nov. 28: 16. 1930.
Chichavac (Chimaltenango).
Open, grassy or rocky hillsides, thickets, brushy or forested slopes,
often in oak and pine-oak forest, rarely on roadside banks, 1,400-
2,800 meters; Chimaltenango; Huehuetenango; Jalapa; Quezalte-
nango (type: Seler & Seler 2893); El Quiche"; Sacatepe"quez ; Santa
Rosa; Solola. Colombia and Venezuela.
FIG. 27. Lithospermum guatemalense. A, habit, 1A natural size; B, flower,
X 11A; C, pistil, X \Vz\ D, dissection of apex of corolla showing stamens, X 2;
E, nutlet, X 5.
152
GIBSON: FLORA OF GUATEMALA 153
Stout perennials, usually erect, rarely decumbent, 30-90 cm. tall, from thick,
woody roots, usually branched above, the stems strigose, often somewhat hispid;
leaves numerous, mostly ascending, the basal leaves oblong-lanceolate, attenuate
to the base, to 10 cm. long, the cauline leaves sessile or subsessile, the blades oblong
to lanceolate or oblanceolate, the lower ones to 7.5 cm. long, often to 2 cm. wide,
the upper ones 1-3 cm. long, 0.4-1 cm. wide, strigose, often pustulate, the veins
usually conspicuous; inflorescences terminal, the branches solitary, geminate, or
ternate, the flowers on short, thick pedicels, usually exceeded by the subtending
leaves; calyx 2-3 mm. long at an thesis, 6-9 mm. long in fruit, the sepals linear;
corolla white, the tube 5-6 mm. long, pubescent outside, glabrous within, the ap-
pendages within the throat well developed, the lobes rounded, minutely erose,
1-2 mm. long; the anthers subsessile on tube, inserted at a point ca. 2 mm. below
the throat; style 3-4 mm. long; ovary ca. 1 mm. long; nutlets 2-3, ovoid, 2.5-
3.5 mm. long, white, lustrous, smooth.
This has been reported from Guatemala as L. discolor Mart. &
Gal.
MACROMERIA D. Don
References: D. Don, Edinb. Phil. Journ. 239. 1832; A. DeCandolle,
in DC. Prodr. 10:68. 1846.
Erect, perennial, herbaceous or suffruticose plants, usually tall and coarse,
pubescent and often hispid; leaves alternate, the lateral veins oblique; inflores-
cences terminal on the branches, the flowers solitary and extra axillary from near
the bases of leaves or leaf-like bracts, the cincinni short at first, in age usually
more or less elongated, the flowers sessile or pedicellate (ours), conspicuous; calyx
5-parted, the segments linear; corolla yellow, whitish, or greenish, the tube elon-
gated, usually dilated above, naked within, the 5 lobes imbricate in bud, spreading
at an thesis; stamens 5, exserted, the filaments inserted on the tube, filiform; an-
thers oblong or linear, muticous, versatile; ovary 4-lobate, the lobes subglobose,
inserted upon a flat gynobase; style filiform, exserted, the stigma small; ovules
suberect; nutlets by abortion mostly 1-2, ovoid, erect; seeds straight, the cotyle-
dons flat.
About a dozen species, ranging from southwestern United States
to Peru. Only one is found in Central America.
Macromeria guatemalensis I. M. Johnston, Journ. Arn. Arb.
29: 232. 1948. Itamo real (San Marcos); te de monte (Huehuete-
nango) .
Dry, forested upper slopes, 2,400-3,900 meters; Huehuetenango;
San Marcos.
Erect, suffruticose perennials to 2 m. tall, much branched, the stems and
branches scabrous; leaves subsessile, the blades lanceolate to linear-lanceolate,
2.5-5 cm. long, 0.5-1.5 cm. wide, attenuate, rounded to acute at the base, the
margins narrowly revolute, veins and costae impressed above, evident below, both
FIG. 28. Macromeria guatemalensis. A, habit of plant, l/i natural size;
B, flower, X 2; C, dissection showing gynobasic style, two mature fruits which
are joined, two aborted ovules, and accrescent calyx lobes, X 3; D, pubescence
with enlarged bases, much enlarged.
154
GIBSON: FLORA OF GUATEMALA 155
surfaces scabrous, the indument of abundant, rigid, tuberculate, appressed hairs;
inflorescences terminal on the branches, the flowers solitary and extra axillary from
near the bases of leaves or leaf-like bracts; flowers on pedicels 3-10 mm. long; calyx
at an thesis 7-10 mm. long, in fruit 11-15 mm. long, the sepals linear, unequal in
length, prominently ribbed, pilose to strigose; corolla pale yellow, ca. 3.5 cm. long,
hirsute outside, glabrous within, basal part of tube ca. 10 mm. long and 1-1.2 mm.
thick, abruptly expanded upward, the upper part cylindrical, ca. 17 mm. long and
5 mm. wide, the lobes triangular, 6-8 mm. long, rounded at apex; stamens exserted,
the filaments 4-7 mm. long, affixed to tube at a point ca. 3 mm. below throat, an-
thers 2.5-3 mm. long; style filiform, exserted; nutlets ovoid, ca. 3 mm. long, white,
lustrous, smooth.
Fruits sometimes used for beads.
ROCHEFORTIA Swartz
References: M. Gurke, in Engler & Prantl, Pflanzenf. 4(3a): 89.
1893; I. Urban, Symb. Ant. 5: 479. 1908.
/
Dioecious shrubs, small trees, or woody vines, usually much branched, some-
times armed with spines; leaves alternate or fasciculate, entire, petiolate, the blades
variously pubescent; inflorescences cymose or the flowers solitary; flowers small,
sessile or pedicellate; calyx lobes 4-5, imbricate; corolla rotate, the tube very short;
stamens 4-5, exserted, the filaments of staminate flowers about equalling the an-
thers in length, those of pistillate flowers shorter, the anthers ovoid, introrse; disk
thick; ovary bilocular or 4-locular, the ovules 4; styles 2, terminal, those of the
pistillate flowers filiform, those of the staminate flowers subulate, the stigmas hir-
sute, dilated or flattened, fruits drupaceous, carnose, more or less globose, con-
taining 4 pyrenes.
Species five or six, all except the following in the West Indies and
northern South America.
Rochefortia hmdellii Camp in Lundell, Contr. Univ. Mich.
Herb. 7: 47. 1942.
Known only from the type, British Honduras, El Cayo District,
Arenal-Valentin Road, in high forest, C. L. Lundell 6167.
Woody vine, the branches slender, sparsely armed with rather blunt spines
2-6 mm. long, the bark gray; leaves fasciculate on short spurs, the petioles slender,
to 12 mm. long, glabrous in age, the blades coriaceous, pale, glabrous, cuneate
obovate to ovate, rounded or obtuse at apex, 3-7 cm. long, 1.5-3 cm. wide, attenu-
ate to the base, entire, with usually 5-7 (rarely 8-9) pairs of veins; inflorescences
appressed-pubescent to glabrate, the cymes to 2 cm. long, 2-6 flowered; calyx
sparsely appressed-pubescent, the 5 lobes imbricate, ovate or suborbicular, obtuse,
1.5-2.5 mm. long, ciliate, slightly accrescent in fruit; corolla glabrous, the tube
1 mm. long, the lobes reflexed, oblong-elliptic to broadly ovate, 2-3 mm. long,
rounded at the apex; stamens 5, alternate with the corolla lobes, the filaments
inserted at the apex of the corolla tube, the anthers oblong, sagittate; ovary gla-
brous, 4-celled; styles 2; fruits globose, lustrous, 5-7 mm. in diameter.
156
FIELDIANA: BOTANY, VOLUME 24
C
FIG. 29. Rochefortialundellii. A, habit of plant, natural size; B, calyx, X 10;
C, corolla opened to show stamens, X 10; D, staminate pistil, X 10.
A staminate plant from Pete"n, Tikal National Park, Lundell
1 7593, has been identified by other workers as R. lundellii. However,
the type specimen is described as a woody vine and according to
label data, the Pete"n collection is a "slender tree, 2" diam., 20 ft.
high ..." Further, it does not appear very different from some An-
tillean collections of the variable R. spinosa (Jacq.) Urban, although
GIBSON: FLORA OF GUATEMALA 157
the petioles of our plant are considerably longer. If additional mate-
rial becomes available it may be possible to place it correctly.
TOURNEFORTIA L.
References: Linnaeus, Gen. 68. 1754; A. de Candolle, in DC. Prodr.
9: 512-531. 1845; Ivan M. Johnston, Observations on the species of
Cordia and Tournefortia known from Brazil, Paraguay, Uruguay, and
Argentina, Contr. Gray Herb. 92: 3-89. 1930; The Boraginaceae of
northeastern South America, Journ. Arn. Arb. 16: 1-64. 1935; Boragi-
naceae of the Southern West Indies, Journ. Arn. Arb. 30: 111-138.
1949.
Shrubs to weak trees or woody vines, glabrous or pubescent; leaves alternate,
entire; inflorescences consisting of dichotomous helicoid cymes or the repeatedly di-
chotomous bostryces borne in panicles; calyx persistent, usually 5-lobate, one sepal
often longer than the others; corolla white, yellow or greenish, small, usually 5-
lobate, the tube cylindric, the limb spreading; stamens usually 5, borne on the
corolla tube, included, the filaments short; ovary 4- celled, sometimes 4-lobate, the
style solitary, terminal, usually bifid at apex, the stigma annular or conic; fruit
drupaceous, entire or lobate at maturity, when dry separating into 2-4 bony nut-
lets, these 1-2-seeded, or with 1-2 empty cavities; endosperm thin; cotyledons flat.
A genus of wide range and much diversification, presenting many
problems in delimitation. There are perhaps 100 species in the trop-
ics of both hemispheres, most numerous in America, with about 25
species in Mexico and Central America. Thirteen occur in Guate-
mala. The altitudinal range of the genus is very wide, often extend-
ing from sea level to 3,600 meters.
Sepals 6-9 mm. long.
Inflorescence dense, compact; corolla lobes 0.5-2 mm. long, obtuse; style short,
not more than one-third the length of corolla tube T. densiflora.
Inflorescence open, lax; corolla lobes 3-4.5 mm. long, acute; style elongated,
almost equalling corolla tube T. elongata.
Sepals less than 6 mm. long.
Corolla tubes 10-12 mm. long T. angustiflora.
Corolla tubes less than 10 mm. long.
Style at anthesis shorter than sepals.
Corolla lobes 3-4.5 mm. long.
Leaves linear or linear-spatulate T. gnaphalodes.
Leaves lance-oblong or lance-elliptic T. longiloba.
Corolla lobes 1-2 mm. long.
Sepals equalling or two-thirds as long as corolla tube T. belizensis.
Sepals less than two-thirds as long as corolla tube.
Leaves and stems glabrous or nearly so; if hairs present, these usually
appressed; leaf blades essentially smooth to the touch; fruits usu-
ally glabrous T. tricolor.
158 FIELDIANA: BOTANY, VOLUME 24
Leaves and stems abundantly hirsute or hirtellous with short, curved
hairs, or if leaf blades glabrate or only sparsely strigose, then rough
to the touch; fruits usually hispid T. hirsutissima.
Style at anthesis equalling or longer than sepals.
Woody vines; fruits conspicuously lobate; anthers usually adherent at their
apices.
Corolla tubes 3-4 mm. long; mature leaves usually essentially glabrous
beneath; sepals usually ovate-lanceolate T. maculata.
Corolla tubes 1.5-2.5 mm. long; mature leaves usually densely sericeous
or tomentose beneath; sepals linear-lanceolate T. volubilis.
Shrubs or small trees; fruits not lobate; anthers free from each other.
Corolla tubes 4-6 mm. long; sepals 2-4 mm. long.
Corolla lobes ovate or lanceolate, acute, usually less than 2 mm. long.
T. petiolaris.
Corolla lobes narrowly triangular, acuminate, 2-3 mm. long.
T. acuti flora.
Corolla tubes 3-4 mm. long; sepals less than 2 mm. long T. glabra.
Tournefortia acutiflora Mart. & Gal. Bull. Acad. Brux. 11(2):
333. 1844.
Usually in damp or wet thickets or forests, rarely on dry, forested
slopes, 1,500-3,400 meters; Chimaltenango; Huehuetenango; Jalapa;
San Marcos; Zacapa. Southern Mexico; Honduras.
Shrubs or small trees, to 8 m. tall, the branches usually stout, densely brown-
tomentose; leaves on petioles 1-8 cm. long, the blades usually ovate-oblong to
elliptic- oblong, rarely broadly obovate, acute or acuminate, rounded to acute at
the base, or abruptly contracted and narrowly decurrent, mostly 8-25 cm. long, 4-
12cm. wide, usually densely and softly velutinous-pilose on both surfaces or strigose
above and velutinous beneath, but may be glabrous or glabrate on both surfaces,
inflorescences usually lax, bostryces few, the flowers sessile or subsessile, usually 1-
3mm. apart; sepals linear to linear-lanceolate, often subulate, strigillose, 2. 5-4 mm.
long; corolla yellow-green or greenish, the tube strigillose outside, ca. 5 mm. long,
the lobes linear- lanceolate, acuminate, recurved, 2.5-3 mm. long; stamens inserted
in throat of tube, the anthers 1.5-2 mm. long; style elongated, surpassing calyx
segments at anthesis and reaching a point just below the anthers; fruits ovoid to
subglobose, glabrous, white, ca. 1 cm. in diameter.
Tournefortia angustiflora Ruiz & Pavon, Fl. Peruv. 2: 25,
t. 151. 1799.
Damp thickets and forest, often on banks along streams, 150-
600 meters; Escuintla; Santa Rosa. Honduras to Panama; South
America.
Weak, often subscandent shrubs, to 4 m. tall, the young branches slender,
sparsely short-hispidulous or almost glabrous; leaves short-petiolate, the blades
ovate-lanceolate to elliptic or oblanceolate, acute or acuminate, acute or attenuate
to the base, rarely obtuse and abruptly decurrent, essentially glabrous, sometimes
sparsely hispidulous beneath along the costa, densely and finely papillose, mostly
GIBSON: FLORA OF GUATEMALA 159
7-18 cm. long, with 5-7 pairs of primary lateral veins; inflorescences of dichotomous
cymes, the nearly straight or only slightly helicoid cymes usually not more than
10 cm. long, often densely flowered, the flowers subsessile to short pedicellate, erect
or nearly so; calyx short, the sepals broadly ovate, 1-1.5 mm. long, usually densely
papillose, minutely strigillose or almost glabrous; corolla greenish white, the tube
slender, narrowed upward, sparsely and minutely strigillose outside, 10-12 mm.
long, the lobes broadly ovate, short, rounded or obtuse and apiculate; stamens in-
serted below mid-tube, the anthers ca. 2 mm. long; style very short, the stigma
almost sessile on the ovary; fruits globose or globose-ovoid, white, glabrous.
Tournefortia belizensis Lundell, Bull. Torr. Bot. Club 69: 399.
1942.
Thickets and forest, 150-1,800 meters; Chiquimula; Pete"n. Brit-
ish Honduras.
Shrubs to 3 m. tall, the young branches hirtellous; leaves subsessile or short-
petiolate, the blades lance-oblong or elliptic-oblong, acute or acuminate, cuneate or
attenuate to the base, hirtellous on both surfaces, mostly 5-12 cm. long, 2-4 cm.
broad; inflorescences pedunculate, small or large, the bostryces usually 6-12 cm.
long, the flowers sessile, ca. 2 mm. apart; sepals linear to linear-lanceolate, acute,
hirtellous, 2.5-4 mm. long; corolla white or greenish, the tube 3-4.5 mm. long,
strigillose outside, the lobes linear or triangular and acuminate, spreading, 1-2 mm.
long; stamens sessile, inserted just above mid-tube, anthers 1 mm. long or less;
style at an thesis not exceeding calyx; fruits ovoid to depressed-globose, glabrous,
white, ca. 5 mm. in diameter.
This may prove to be a form or variety of T. umbellata HBK. of
the West Indies and the Yucatan Peninsula, Mexico, as it differs
only in vesture. The branches, stems, leaves, and calyces of T. um-
bellata are glabrous or nearly so, and the leaf blades are sometimes
scaberulous.
Tournefortia bicolor Swartz, Prodr. Veg. Ind. Occ. 40. 1788;
Johnston, Contr. Gray Herb. 92: 69. 1930 and Journ. Am. Arb. 16:
49. 1935. T. laevigata Lam. Encycl. 1: 416. 1791. T. nitida HBK.
Nova Gen. & Sp. 3: 84. 1819. T. laevigata var. latifolia DC. Prodr.
9: 519. 1845.
Damp or wet thickets or forest, sometimes in mangrove swamps,
sea level to 1,500 meters; Alta Verapaz; Huehuetenango; Izabal;
Quezaltenango; Sacatepe"quez; San Marcos. Southern Mexico to
Panama; West Indies; tropical South America.
Large shrubs to 5 m. tall, often subscandent, the branches almost glabrous or
with scattered, weak appressed hairs; leaves on petioles 1-2 cm. long, the blades
ovate to elliptic, lance-ovate or lance-oblong, 5-15 cm. long, 3-9 cm. broad, usually
rather thick and firm, acute to acuminate, usually obtuse or rounded (rarely acute)
at the base, nearly glabrous or with scattered, appressed hairs on one or both sur-
160 FIELDIANA: BOTANY, VOLUME 24
faces; inflorescences usually dense and much branched, small or large, the bostryces
short, the flowers sessile; sepals ovate or ovate-lanceolate, acute, 1-2 mm. long
(rarely to 3.5 mm.), sparsely and minutely strigose; corolla white, the tube strigil-
lose outside, usually 4-6 mm. long (rarely to 8 mm.), the lobes obtuse, usually
apiculate; stamens inserted about mid-tube, the anthers 1.5-2 mm. long; stigma
subsessile on ovary; fruits white, subglobose, glabrous, ca. 6 mm. long when dry.
Tournefortia densiflora Mart. & Gal. Bull. Acad. Brux. 11(2):
333. 1844. T. scholastica Standl. & L. Wms., Ceiba 3: 56. 1952.
Chompipito (Guatemala) .
Damp thickets to dry mountain slopes, 120-1,400 meters; Guate-
mala; Huehuetenango; Jalapa; Jutiapa; El Progreso; El Quiche;
Zacapa. Mexico; Honduras.
Erect shrubs to 2 m. tall, the branches terete, usually short-hirsute; leaves on
petioles 1-4 cm. long or the upper leaves sometimes subsessile, the blades lanceolate
to ovate or elliptic, 4-20 cm. long, acute or acuminate, often attenuate to the base,
sometimes acute, rarely rounded and somewhat abruptly decurrent (usually the
older leaves on lower stems), scabrous on both surfaces or scabrous above and softly
pubescent beneath; inflorescences long-pedunculate, the densely-flowered bostryces
few or rather numerous, usually less than 8 cm. long; calyx 6-9 mm. long, the
sepals linear, erect, densely hispidulous; corolla white, the tube densely hispidulous
outside, 7-9 mm. long, the lobes ovate, acute or obtuse, often somewhat crenate;
stamens inserted about mid-tube, the anthers 2 mm. long; style short, rising to
about one-third the length of tube; fruits globose-ovoid, sparsely hispidulous or
glabrous, ca. 6 mm. long when dry.
\
Apparently closely related to T. mutabilis Vent, of Mexico, which
includes T. hartwegiana Steud. and probably T. trichocalycina DC.
and T. capitata Mart. & Gal. All of these are variable in quantity of
indument, and all have sepals shorter than those of T. densiflora,
usually about half (rarely two-thirds) as long as the mature corolla
tube.
T. calycina Benth., also of Mexico, may be separated from all of
these by its broader, oblanceolate to spatulate sepals; in addition,
the few specimens I have seen appear to be uniformly scabrous.
Tournefortia elongata D. Gibson, Fieldiana: Botany, 32: 65,
/. 1. 1969. Yerba coche (Jalapa).
Upper volcanic slopes, wet, mixed broadleaf and cypress forest,
cloud forest, damp second-growth thickets, rarely in hedgerows on
upper slopes, 1,800-3,600 meters; Chimaltenango; Escuintla; Guate-
mala; Jalapa; El Progreso; Quezaltenango; El Quiche"; San Marcos;
Solola: Totonicapan. Mexico (Chiapas).
Shrubs or small trees to 8 m. tall, the branchlets densely brown hirsute or
hirsute-tomentose; leaves on stout, tomentose petioles mostly 0.5-4 cm. long, the
FIG. 30. Tournefortia elongata. A, habit, }/£ natural size; B, calyx opened to
show gynoecium, X 3; C, corolla opened to show stamens, X 3.
161
162 FIELDIANA: BOTANY, VOLUME 24
blades elliptic-oblong to oblong-ovate, usually 6-25 cm. long, 2-9 cm. wide, long
acuminate, long-attenuate to the base, densely hirsute or short-hispid on both
surfaces; inflorescences open, lax, of dichotomous helicoid cymes, long pedunculate,
the flowers secund; sepals linear to linear-lanceolate, acuminate, 6-9 mm. long,
erect, usually densely hispidulous; corolla green or greenish-yellow, the tube his-
pidulous outside, 8-9 mm. long, the lobes 2.5-3.5 mm. long, triangular, acute,
spreading or somewhat recurved; stamens subsessile in corolla throat, anthers ca.
2 mm. long; style elongated, 6-7 mm. long, the entire gynoecium to 9 mm. long,
always equalling or exceeding the calyx; fruits white, globose-ovoid, glabrous, ca.
1 cm. in diameter.
Tournefortia elongata closely resembles T. johnstonii Standley of
Costa Rica. However, the gynoecium of T. johnstonii at anthesis is
only 5-6 mm. long, the style always shorter than the calyx, the sepals
narrowly linear, and the corolla lobes 4-5 mm. long, subulate and
acuminate.
The single Mexican specimen seen, Breedlove 9708, is atypical in
that the sepals are mostly 3-5 mm. long. However, those of a few
flowers are 6 mm. long and occasionally one sepal of a flower will
attain 6 mm. while the others remain only 3-4 mm. long.
Tournefortia glabra L. Sp. PI. 141. 1753. T. cymosa L. Sp.
PI. ed. 2, 202. 1762.
Wet thickets or mixed forest, sometimes along streams, ofteh in
second growth, 50-2,600 meters; Alta Verapaz; Escuintla; Guate-
mala; Huehuetenango; Izabal; Pete"n; Quezaltenango; Retalhuleu;
Sacatepe"quez ; San Marcos; Solola. Southern Mexico to Panama;
South America.
Slender shrubs or small trees to 7 m. tall, the branches glabrous or nearly so;
leaves on petioles 1-5 cm. long, the blades lanceolate, oblanceolate, elliptic or ovate-
elliptic, apex acute or acuminate, acute to long attenuate at base, mostly 7-20 cm.
long, usually glabrous or nearly so but when young may be minutely appressed-
pilosulous, primary lateral veins usually 8-12 pairs; inflorescence usually broad
and lax, the dichotomous branches slender, often numerous, usually many-flowered,
the flowers sessile; calyx usually 1.3-2 mm. long (rarely 3 mm.), the sepals lan-
ceolate or oblong-lanceolate, acute, sparsely and minutely strigillose or almost
glabrous; corolla white, greenish white, or yellowish green, the tube minutely pu-
berulent on the outside, dilated near the top, 3-4 mm. long, the lobes 1-2 mm. long,
narrowly triangular, acute to acuminate; stamens inserted a little below sinuses,
the anthers 0.5-1 mm. long; style elongated, surpassing the sepals at anthesis, and
usually reaching a point just below the stamens; fruit subglobose, appearing some-
what turbinate when dry, glabrous, 3-5 mm. in diameter.
Tournefortia gnaphalodes (L.) R. Br. ex Roem. & Schult. Syst.
4: 538. 1819. Heliotropium gnaphalodes L. Syst. ed. 10: 913. 1759.
GIBSON: FLORA OF GUATEMALA 163
Mallotonia gnaphalodes Britt. Ann. Mo. Bot. Card. 2: 47. 1915.
Messerschmidia gnaphalodes I. M. Johnston, Journ. Arn. Arb. 6: 165.
1935.
On open, sandy Atlantic beaches; British Honduras. Southern
Florida; Yucatan Peninsula, Mexico; Nicaragua; West Indies.
Stout shrubs to 3 m. tall, often forming dense clumps, the branches white-
sericeous and densely leafy; leaves linear or linear-spatulate, 4-10 cm. long, obtuse
or rounded at apex, attenuate to the base, somewhat succulent, densely white-
sericeous; inflorescences equalling or shorter than the leaves, with 2-4 short,
densely-flowered bostryces; calyx 3-5 mm. long, the lobes broadly lanceolate, acute,
but appearing obtuse due to the densely sericeous indument outside; corolla white,
the tube tomentose outside, 4-5 mm. long, the lobes lanceolate, acute, 2-3 mm.
long; stamens inserted below the throat of the tube, the anthers ca. 1 mm. long;
style shorter than sepals; fruit ovoid, brown, glabrous, 5-6 mm. long, the lower half
composed of corky exocarp.
Tournefortia hirsutissima L. Sp. PL 140. 1753. Messer-
schmidtia hirsutissima Roem. & Schult. Syst. 4: 541. 1819. T. ellip-
tica Mart. & Gal. Bull. Acad. Brux. 9(2) : 332. 1844. T. schomburgkii
DC. Prodr. 9: 517. 1845. T. alba Splitberger ex DeVriese, Nederl.
Kruidkund. Arch. 1: 347. 1848; Schomburgk, Fauna u. Fl. Brit.
Guiana 1151. 1848. T. billbergiana Beurl. Vet. Akad. Handl. Stockh.
139. 1856. T. bicolor var. calycosa Donn.-Sm. Bot. Gaz. 14: 27. 1889.
Nigua (Pete"n).
Wet or dry thickets or mixed forest, frequently in second growth,
sea level to 1,800 meters; Alta Verapaz; Chimaltenango; Chiqui-
mula; Escuintla; Guatemala; Huehuetenango; Izabal; Pete"n; Re-
talhuleu; Sacatepe"quez ; San Marcos. Southern Florida; Mexico;
British Honduras to Panama; WTest Indies; South America.
Suberect shrubs to 3 m. high, or elongating and becoming more or less scandent,
the branches hispid or hirsute to nearly glabrous or glabrate; leaves on petioles
0.5-2 cm. long, the blades lance-oblong, oblong-ovate, ovate, or elliptic, mostly
7-20 cm. long, 3-7 cm. broad, usually acuminate, sometimes acute, rarely obtuse,
acute or obtuse at the base, usually scabrous-hirsute above or if glabrate, usually
rough to the touch, rarely nearly glabrous and smooth; inflorescences often large
and diffuse, of numerous dichotomous helicoid cymes, the bostryces usually less
than 4 cm. long; sepals ovate-lanceolate, lanceolate, oblong, oblanceolate, elliptic,
linear, or linear-spatulate, acute or acuminate, hispidulous or nearly glabrous,
sometimes papillose, usually 2-4 mm. long, rarely 5 mm.; corolla white or greenish,
the tube densely or sparsely strigose outside, 4-6 mm. long, the lobes short, obtuse,
apiculate; stamens sessile, inserted at or a little below mid-tube, the anthers 1-
2 mm. long; style subsessile on ovary; mature fruit globose, white, often hispid.
This is a common, weedy plant in many parts of the Central
American tierra caliente. Because of the great range of variation in
164 FIELDIANA: BOTANY, VOLUME 24
quantity of indument on the leaves and stems and the variation in
length and form of the sepals, it is difficult to delimit good taxonomic
units. The more glabrous forms of T. hirsutissima have been con-
fused with the closely related T. bicolor Sw., and Johnston, in his
discussion of T. bicolor var. calycosa Donn.-Sm. (Journ. Arn. Arb. 16:
52. 1935) concluded that the differences separating T. hirsutissima
and T. bicolor "are almost exclusively those of pubescence. ..." Be-
cause he felt this difference to be decisive, he retained the var. caly-
cosa as a variety of T. bicolor; however, the sepals of the true and
uniformly glabrous T. bicolor are consistently short and ovate to
ovate-lanceolate, while those of the var. calycosa are much like many
of the various forms of T. hirsutissima. Although Johnston held that
the fruits of T. bicolor and the var. calycosa are always glabrous,
while those of T. hirsutissima are usually hairy, a collection from
Sacatepe"quez in 1935, Skutch 2069, with elongated sepals, glabrous
outside and sericeous within, and nearly glabrous leaves, also has
many hairy young fruits.
Tournefortia billbergiana Beurl. is also placed here as the leaves
are definitely though somewhat sparsely strigose, the fruits are hairy,
and the calyx is like those of T. hirsutissima.
In view of the numerous variations and forms which suggest in-
tergradation, until monographic study can be done, it seems best to
treat this complex collectively as T. hirsutissima L., sens. lat.
Tournefortia longiloba D. Gibson, Fieldiana: Bot. 32: 68,
/. 2. 1969.
On steep, damp slopes, cloud forests, sometimes with Quercus and
Pinus, 2,400-3,400 meters; Huehuetenango; El Quiche"; El Progreso.
Mexico (Chiapas); Costa Rica.
Shrubs to 5 m. tall; leaves on stout, tomentose petioles 1-3.5 cm. long, the
blades lance-oblong or lance-elliptic, mostly 8-18 cm. long, 3-5 cm. wide, acute to
acuminate, usually attenuate to the base, rarely rounded to obtuse and abruptly
decurrent, usually puberulous above, tomentose to velutinous below (rarely gla
brate), with 8-14 pairs of lateral nerves; inflorescences paniculate, of few to many
dichotomous cymes, the bostryces many-flowered, the flowers secund, often placed
at intervals of 5-6 mm., sessile or on pedicels 1-6 mm. long; sepals ovate-lanceolate
to lanceolate, acute, hispidulous to hirsutulous, 3-5 mm. long; corolla white, the
tube hirsutulous on the outside, ca. 5 mm. long, the lobes narrowly triangular,
acuminate, spreading to recurved, 3-4.5 mm. long; stamens included, sessile or
subsessile, inserted in tube at a point just above the middle and ca. 3 mm. above
the base, anthers 1.5-2 mm. long; ovary subglobose, the style short, not exceeding
the sepals at an thesis; mature fruits globose-ovoid, white, 7-9 mm. long, 5-6 mm.
in diameter when dry.
GIBSON: FLORA OF GUATEMALA 165
Sometimes confused with T. acutiflora Mart. & Gal. which it
superficially resembles. However, the style of T. acutiflora is elon-
gated, considerably surpassing the sepals; the corolla lobes are only
2.5-3 mm. long, and the corolla tube is 6-7 mm. long. T. longiloba
is probably closely related to T. subspicata Donn.-Sm. of Costa Rica,
which also has a short style and long corolla lobes, but has a nearly
glabrous calyx, a longer corolla tube (7 mm. long) with appressed
hairs, and leaves that are glabrous beneath or very sparsely pubescent.
Tournefortia maculata Jacq. Enum. PL Carib. 14. 1760. T. sy-
ringaefolia Vahl, Symb. Bot. 3: 23. 1794. T. peruviana Poir. Encycl.
Suppl. 4: 425. 1816. T. sagraena DC. Prodr. 9: 522. 1845. T. guade-
lupensis Urban in Fedde, Rep. Sp. Nov. 17: 169. 1921. Messer-
schmidtia syringaefolia (Vahl) Roem. & Schult. Syst. 4: 543. 1819.
Damp thickets or mixed forest, 800-1,500 meters; Alta Verapaz;
Pete"n; Quezaltenango; San Marcos; Solola. Honduras to Panama;
West Indies; tropical South America.
Woody vines, the branches slender, sparsely or densely short- hirtellous or al-
most glabrous; leaves on petioles 0.5-1.5 cm. long, the blades ovate to lance-elliptic
or broadly lanceolate, mostly 3-10 cm. long, 1-5 cm. wide, acute to acuminate,
rounded to acute at the base, usually almost glabrous; inflorescences lax, of slender
dichotomous cymes with bostryces to 5 cm. long; flowers sessile or very short-pedi-
cellate, 2-8 mm. apart; calyx 0.5-1.5(2) mm. long, the sepals usually ovate-lanceo-
late, somewhat strigillose; corolla pale green to greenish white, the tube narrowly
cylindric, strigillose outside, 3-4(5) mm. long, the lobes 1.5-2 mm. long, acuminate,
spreading; stamens inserted in throat of tube, subsessile, the anthers usually con-
nate at their apices; style elongated, filiform, the gynoecium as long as the corolla
tube; fruit yellow or orange, glabrous, conspicuously 4-lobate, containing 4 one-
seeded nutlets.
A single specimen from Alta Verapaz, von Turckheim 1790, is
atypical; monographic study may prove it to be a form, or it may
represent a species new to Central America. The stems and petioles
are densely invested with erect to shaggy hairs, the sepals are acumi-
nate, ca. 2 mm. long and pilose; the costae and margins of young
leaves are pilose; the mature leaf blades are thick and glabrate. The
plant is much like the South American T. elegans Cham., which John-
ston considered synonymous with T. paniculata Cham. For a dis-
cussion of T. paniculata and its relation to T. maculata (as T.
syringaefolia Vahl and as T. peruviana Poir.) see Johnston, Contr.
Gray Herb. 92: 78. 1930 and Journ. Am. Arb. 16: 49. 1935.
There is also a plant in Honduras, A. Molina 7269, which closely
resembles those from the West Indies, described as T. laurifolia Vent.
166 FIELDIANA: BOTANY, VOLUME 24
It has longer sepals and corolla lobes and the corolla tube is glabrous
save for some puberulence below the lobes. (See Johnston, Journ.
Am. Arb. 30: 130-131. 1949.)
The vesture of plants from southern Central America is often
much heavier than that of ours.
Tournefortia petiolaris DC. Prodr. 9: 520. 1845. T. nelsonii
Donn.-Sm., Bot. Gaz. 23: 10. 1897.
Damp or dry slopes, ravines, wet cloud forest, 1,400-3,000 me-
ters; Chimaltenango; Chiquimula; Jalapa; El Quiche"; Sacatepe'quez ;
San Marcos; Solola; Zacapa. Southern Mexico; El Salvador.
Shrubs or weak trees to 8 m. tall, the stems and branches tomentose or strigil-
lose or nearly glabrous; leaves on petioles usually 2-3 (rarely to 10) cm. long, the
blades elliptic, ovate elliptic, oblong-ovate, oblanceolate, or lance-ovate, mostly
8-15 cm. long (rarely to 25 cm.), 3-12 cm. wide, acute or acuminate, often atten-
uate to the base but may be obtuse or almost rounded or abruptly contracted and
decurrent, somewhat scabrid to velutinous or nearly glabrous above, often gla-
brous or with only short puberulence beneath but may be velutinous; inflorescences
usually large, long pedunculate, repeatedly dichotomous, the bostryces slender, usu-
ally lax, many flowered, the flowers often 2-4 mm. apart, mostly sessile or subsessile
but occasionally with some flowers on pedicels 1-2 mm. long; calyx segments lan-
ceolate or oblong-lanceolate, acute to acuminate, 2-3.5 mm. long, usually hispidu-
lous; corolla greenish white or yellowish green, the tube 5-6 mm. long, hispidulous
outside, the lobes ovate to lanceolate, acute, 1-2 mm. long; stamens with short
filaments inserted below the throat, anthers 1-1.5 mm. long; style at anthesis sur-
passing the calyx and elongating in maturity; fruit greenish white, globose obovoid
when dry, glabrous.
Apparently closely related to T. acutiflora Mart. & Gal., which
has longer, acuminate corolla lobes.
The very large leaf form, in which the blade is abruptly con-
tracted and decurrent has also been observed on specimens of T.
acutiflora and T. longiloba D. Gibson. The plants of T. petiolaris
with large leaves usually have longer petioles. Donnell-Smith, a
year after proposing his new species, T. nelsonii, based on this large
and velutinous leaf form, rejected it, stating that "it does not appear
to differ from T. petiolaris DC."
Tournefortia volubilis L. Sp. PI. 140. 1753. T. sericea Vahl,
Ecol. 1: 17. 1796. T. floribunda HBK. Nova Gen. & Sp. 3: 79. 1818.
T. velutina HBK. I.e. t. 201. 1819. T. potosina Standl. Contr. U. S.
Nat. Herb. 23: 1230. 1924. Miona (Zacapa).
Damp or dry thickets, brushy, rocky slopes, sometimes in hedges
or clearings, 120-1,000 meters; Baja Verapaz; Guatemala; Jalapa;
GIBSON: FLORA OF GUATEMALA 167
Jutiapa; El Progreso; El Quiche"; Zacapa. Southern Florida and
Texas; Mexico; El Salvador; Honduras; Nicaragua; West Indies;
northern South America.
Woody vines, often diffusely branching, the branches slender, usually densely
white tomentose; leaves on petioles 2-10 mm. long, ovate to oblong, lance-oblong
or (rarely) linear-lanceolate, the blades 2-8 cm. long, 0.6-3.5 cm. wide, acute or
acuminate, usually rounded or obtuse at the base, usually densely tomentose be-
neath with soft grey or white hairs, often equally tomentose above or sometimes
glabrate; inflorescences slender-pedunculate, usually small, of few-many slender,
forking cymes to 4 cm. long with flowers 2-6 mm. apart; sepals 1-2 mm. long,
linear-lanceolate, acute or acuminate; corolla greenish-white, the tube 1.5-2.5 mm.
long, densely sericeous outside, the lobes linear-subulate, a little shorter than the
tube; style at anthesis about equalling the calyx ; stamens inserted in throat of tube,
the small anthers often connate at their apices; fruits depressed, white with black
dots, conspicuously 4-lobate, glabrous, 2-3 mm. long.
The fruits are often modified into insect galls and are then much
larger than normal and densely white-pilose.
VERBENACEAE. Verbena family
DOROTHY N. GIBSON
Reference: H. N. Moldenke, Verbenaceae (of the Maya area),
Carnegie Inst. Wash. Publ. 522: 148-223. 1940.
Mostly trees or shrubs, sometimes herbs or woody vines; branches often tetrag-
onous; leaves mostly opposite, without stipules, simple or palmately compound,
the blades entire or variously dentate, lobate, cleft, or incised; inflorescences axil-
lary or terminal, cymose, paniculate, racemose, or spicate, sometimes capitate, the
flowers perfect or imperfect, small or large; calyx gamopetalous, persistent and
often accrescent, usually with 4-5 lobes or teeth; corolla gamopetalous, regular or
irregular, funnelform or salverform, the lobes 4 or 5, the limb often somewhat
bilabiate; stamens usually 4 and didynamous, rarely 2, the filaments inserted on
the corolla tube, staminodes sometimes present; ovary of 2, rarely 4-5 carpels,
often with 4 lobes, usually with 2-5 locules, rarely unilocular, the locules with 1
or 2 ovules; ovules anatropous and basal or sometimes lateral on a central or axile
placenta; fruit a dry schizocarp or drupaceous, usually with 2-4 locules (rarely 1),
indehiscent or separating into 2 or 4 cocci or pyrenes.
About 75 genera, in both hemispheres, chiefly in tropical regions.
Only the following are native in Central America. One other, a
native of southern Asia, Holmskioldia sanguined Retz., is sometimes
cultivated in Guatemala for ornament. It is a pubescent or glabrate
shrub with ovate, entire or dentate leaves conspicuously punctate
beneath; the flowers are in dense terminal racemes; the calyx is
rotate, bright red or orange, orbicular, 2.5 cm. broad, entire; corolla
168 FIELDIANA: BOTANY, VOLUME 24
is brick-red or orange, 2-2.5 cm. long. In Costa Rica this plant is
called "sombrero chino."
Leaves palmately compound Vitex,
Leaves simple.
Inflorescence cymose or paniculate.
Fertile stamens 2; staminodes 2 Cornutia.
Fertile stamens 4 or 5.
Indument of stellate or dendroid hairs; stigma capitate or peltate.
Callicarpa.
Indument of simple hairs; stigma bifid.
Stigma deeply bifid, the lobes subulate Aegiphila.
Stigma shallowly bifid, the lobes acute Clerodendrum.
Inflorescence spicate or racemose.
Fertile stamens 2; staminodes 2; calyx closely appressed to rachis and often
more or less immersed in it Stachytarpheta.
Fertile stamens 4; staminodes 1 or none; calyx not appressed to rachis.
Fruits capsular; ovary unilocular Avicennia.
Fruits not capsular; ovary bilocular or with 4 perfect or imperfect locules.
Calyx deeply lobate, the lobes showy, elongating and much enlarged in
fruit, usually blue or purple, rarely white, reticulate veined . . Petrea.
Calyx not as above.
Plants herbaceous or sometimes woody at base.
Fruiting calyx globose, bearing minute, uncinate hairs Priva.
Fruiting calyx not globose, without uncinate hairs.
Corolla with 4 lobes; calyx minute and inconspicuous. . . .Lippia.
Corolla with 5 lobes; calyx evident, not minute.
Fruiting calyx broadly campanulate; fruit with slightly fleshy
exocarp Tamonea.
Fruiting calyx tubular; fruit dry.
Style filiform; fruit composed of 2 rostrate cocci. . . .Bouchea.
Style short; fruit composed of 4 cocci, not rostrate. . Verbena.
Plants trees or shrubs.
Fruit dry.
Corolla with 5 lobes; calyx early deciduous; ovary with 4 locules;
fruit 8-20 mm. long, naked Rehdera.
Corolla with 4 lobes; calyx persistent, enclosing fruit; ovary with
2 locules; fruit less than 4 mm. long Lippia.
Fruit drupaceous.
Fruiting calyx contracted at the apex and completely enclosing
the fruit Duranta.
Fruiting calyx not contracted nor completely enclosing the fruit.
Inflorescence headlike, the flowers in dense, usually short spikes ;
calyx small and inconspicuous; ovary bilocular. .Lantana.
Inflorescence not headlike, the flowers in elongated, often lax
racemes; calyx evident; ovary with 4 perfect or imperfect
locules Citharexylum.
AEGIPHILA Jacquin
Reference: Harold N. Moldenke, A monograph of the genus Aegi-
phila, Brittonia 1: 245-477. 1934.
GIBSON: FLORA OF GUATEMALA 169
Shrubs or small trees, sometimes scandent, the branches tetragonal or sub-
terete; leaves simple, usually opposite and decussate, rarely verticillate, deciduous,
estipulate; inflorescences cymose, the cymes often appearing paniculate or simu-
lating corymbs or heads, axillary or terminal; calyx more or less campanulate or
tubular, the margin truncate and entire or subtruncate and scarious or with 4-5
teeth or lobes, accrescent and thickened in fruit; corolla funnelform or salverform,
the tube cylindric, the limb 4-5-parted, the lobes imbricate; stamens 4-5, equal or
subequal, inserted below the mouth of the tube, included or exserted; style ter-
minal, the stigma bifid, the branches subulate; ovary perfectly or imperfectly
4-locular, each locule uniovulate, the ovules attached laterally; fruits drupaceous,
usually 4-seeded, or the seeds by abortion fewer, without endosperm.
Heterostyly is present in most (perhaps all) species; floral char-
acters within the genus are usually not especially significant; there
is considerable clinal variation in leaf form and size, and sometimes
in size of inflorescences. It is therefore, regrettably, often difficult to
separate species morphologically, especially those with large, pani-
culiform inflorescences, without relying heavily on differences in the
fruiting calyces.
Perhaps 150 species in tropical America. Only ten are known
from Guatemala, but 12 are treated here. Aegipkila deppeana Steud.
and A. valerii Standley occur in Chiapas, Mexico as well as in south-
ern Central America and are therefore included here. One plant
previously reported from Guatemala has been omitted, as A. hoff-
mannioides Standley & Steyermark has an inferior ovary and is ac-
tually Hoffmannia lenticellata Hemsl. (Rubiaceae).
Inflorescences appearing paniculate or corymbiform, usually appearing terminal.
Margins of fruiting calyces always entire, truncate or subtruncate.
A. laxicupulis.
Margins of fruiting calyces lobate or irregularly torn, often somewhat scarious.
Fruiting calyces shallowly cupuliform to patelliform, the margins always dis-
tinctly lobate A. martinicensis.
Fruiting calyces first urceolate, later deeply cupuliform, the margins irreg-
ularly torn.
Internodes usually 1-2 cm. long; calyces at anthesis truncate or subtrun-
cate (rarely irregularly lobate) ; fruiting calyces ca. 3 mm. long.
A. skutchii.
Internodes usually 3-8 cm. long; calyces at anthesis lobate; fruiting calyces
4-7 mm. long.
Pedicels and calyces minutely appressed-puberulent; leaves essentially
glabrous beneath with 7-12 pairs of lateral veins A. elata.
Pedicels and calyces shortly and densely tomentose; leaves densely or
sparsely tomentose beneath with 5-8 pairs of lateral veins.
A. deppeana.
Inflorescences cymose, axillary or lateral.
Inflorescences pedunculate.
Peduncles often half as long as the leaves, lax A. costaricensis.
Peduncles very short, only 4-15 mm. long, not lax.
170 FIELDIANA: BOTANY, VOLUME 24
Leaves very large, 25-45 cm. long, to 16 cm. wide, with 20-22 pairs of
lateral veins A. standleyi.
Leaves 3-20 cm. long, 2.5-7.5 cm. wide, with 7-16 pairs of lateral veins.
Corollas 5-parted; leaves with 10-16 pairs of lateral veins. . . .A. valerii.
Corollas 4-parted; leaves with 7-9 pairs of lateral veins.
Flowers pedicellate A. pauciflora.
Flowers sessile A. hastingsiana.
Inflorescences sessile and fasciculate.
Calyces 2-3.5 mm. long, minutely pubescent or almost glabrous.
A. monstrosa.
Calyces 6-8 mm. long, densely ochraceous-velutinous A. fasciculata.
Aegiphila costaricensis Moldenke in Fedde, Rep. Sp. Nov. 33:
119. 1933. Clerodendron matudae Standl. Field Mus. Bot. 17: 206.
1937.
Mixed lowland forest, 720 meters; Quezaltenango (near Colom-
bia, Skutch 2012). Mexico (Chiapas); Costa Rica; Panama.
Trees to 8 m. tall, the branchlets glabrous or sparsely pilose at the nodes, the
trunk as much as 20 cm. in diameter, the crown broad and flat, the bark pale
brown, slightly roughened; leaves opposite or ternate, often somewhat crowded
near the ends of the branches, on petioles 3-7 mm. long, the blades membrana-
ceous, lance-oblong to oblanceolate or obovate, 7-30 cm. long, 4-10 cm. wide,
acute or acuminate, cuneate-acute at the base, glabrous, entire, the leaves of a pair
often very unequal; cymes axillary on long, slender peduncles, often half as long
as the leaves, very lax, few-flowered, the pedicels 1-3 cm. long, filiform, glabrous;
calyx broadly campanulate, 2-5 mm. long, glabrate, subtruncate, usually 4-cuspi-
date; corolla white, the tube 4-8 mm. long, glabrous, the 5 lobes 6-8 mm. long;
stamens 5, usually long-exserted ; fruit ovoid, first yellow, at maturity black, 5 mm.
long or larger.
Readily distinguished from other local species by the 5-parted
rather than 4-parted corollas, and by the lax inflorescence with
greatly elongated pedicels and slender peduncles.
Aegiphila deppeana Steud. Norn. Bot. ed. 2, 1: 29. 1840. A.
brachiata Cham. & Schlecht. Linnaea 6: 371. 1831 (not A. brachiata
Veil. 1825). A. berteriana Schauer in DC. Prodr. 11: 654. 1847.
A. pacifica Greenm. Proc. Am. Acad. 33: 485. 1898.
Pete"n; at or near sea level. Southern Mexico; Nicaragua; Costa
Rica.
Shrubs or small trees, often subscandent, to 7 m. tall; branchlets obtusely
tetragonal or terete, shortly and densely tomentose, sometimes glabrate in age;
leaves on petioles 4-13 mm. long, these densely tomentose, the blades membranous
to subcoriaceous, ovate to oblong-ovate, 5-16 cm. long, 2-9 cm. wide, entire, acute
to short-acuminate, rounded or acute at base, indument of upper surface confined
GIBSON: FLORA OF GUATEMALA 171
to midrib and to the 4-8 pairs of lateral veins, usually densely tomentose beneath,
rarely glabrate in age; inflorescences cymose, appearing paniculate, the cymes in
the axils of the upper leaves, about half as long as the leaves, bracteate, many-
flowered, peduncles and pedicels tomentose, the pedicels ca. 3 mm. long; calyx
campanulate at anthesis, 3-4 mm. long, 4-lobate, pubescent to tomentose, in fruit
cupuliform, 6-9 mm. long, indurated, somewhat tomentose to glabrate, rugose,
irregularly and shallowly lobed; corolla white to pale yellow, glabrous, the tube
5-6 mm. long, the 4 lobes 3-4 mm. long; fruits obovoid, yellow, to 10 mm. long.
Aegiphila elata Swartz, Prodr. Veg. Ind. Occ. 31. 1788.
Damp thickets, brushy slopes, often in second growth, 75-350
meters; Alta Verapaz; Izabal; Pete"n (fide Moldenke). Southern
Mexico; British Honduras to Panama; West Indies; South America.
Erect shrubs or woody vines, the branchlets glabrous or minutely puberulous,
sometimes compressed and widened at the nodes; leaves on petioles 5-15 mm. long,
the blades ovate, oblong-ovate, or elliptic, 6-15 cm. long, 2-7 cm. wide, acute to
acuminate, obtuse or rounded at the base, entire, sometimes somewhat asymmetric,
glabrous above and often lustrous, glabrous beneath or appressed-pubescent only
on the costae and veins, minutely glandular-punctate beneath, lateral veins 7-12
pairs; inflorescences often large, the cymes many-flowered, dense or rather open,
the pedicels 4-8 mm. long, puberulent; calyx at anthesis more or less puberulent,
turbinate-funnelform, 3-5 mm. long, the margin usually with 4 distinct lobes but
may be subtruncate, in fruit first flask-shaped, later cupuliform, 4-7 mm. long, the
margin irregularly lobate, often becoming somewhat scarious; corolla cream-colored
to pale yellow, the tube 5-8 mm. long, ca. 1 mm. broad, glabrous, the 4 lobes 3-
6 mm. long; filaments of stamens inserted about midway of the tube; fruit sub-
globose to oblong-ovoid, yellow to orange, ca. 7 mm. long, 5-6 mm. broad.
Aegiphila fasciculata Donn.-Sm. Bot. Gaz. 57: 425. 1914.
Known only from the type, collected near Coban, Alta Verapaz,
1,350 meters, Tuerckheim 4013, and a single collection from Nica-
ragua, 1,400 meters, L. 0. Williams et al. 27757.
Trees to 6 m. tall, the branches stout, densely ochraceous-velutinous; leaves
on petioles 2-3.5 cm. long, the blades coriaceous, lance-oblong to ovate-elliptic,
12-22 cm. long, 5-8 cm. wide, acuminate, acute at the base, entire, sparsely or
densely spreading-pilose above, ochraceous-tomentose beneath, the lateral nerves
7-9 pairs; inflorescences axillary, sessile, the cymes fasciculate, subglomerate,
densely flowered, the pedicels 1-2 mm. long; calyx turbinate-campanulate, 6-8 mm.
long, 4-mucronulate, densely ochraceous-velutinous; corolla white, the tube 5-
6 mm. long, glabrous, the 4 lobes 4-5 mm. long; ovary glabrous; fruit unknown.
Aegiphila hastingsiana Moldenke, Phytologia 1: 207. 1937.
Known only from the type, said to have been collected in Guate-
mala in 1864 by R. Grosourdy.
172 FIELDIANA: BOTANY, VOLUME 24
Branchlets slender, obtusely tetragonal, densely short-pubescent; leaves char-
taceous, on petioles 3-10 mm. long, the blades elliptic to ovate-elliptic, 3-10 cm.
long, 2.5-4.5 cm. wide, short-acuminate, rather long- acuminate at the base, en-
tire, puberulent to short-pubescent above, densely velutinous-pubescent beneath,
densely punctate, the lateral nerves 7-9 pairs; inflorescences pedunculate, the pe-
duncles 5-10 mm. long, axillary, the cymes solitary, 1.5-2.5 cm. long, densely
flowered, the flowers sessile; calyx campanulate, 3 mm. long, densely short-pubes-
cent, the 4 lobes ovate, acute; corolla tube about 2.5 mm. long, glabrous, the 4
lobes 2 mm. long; ovary glabrous.
Aegiphila laxicupulis Moldenke in Fedde, Rep. Sp. Nov. 33:
130. 1933 (type, Heyde 191, perhaps from Santa Rosa). A. magnified
Moldenke, I.e. 132 (type from Nicaragua) .
Damp thickets and second growth woods, 330-1,050 meters;
Chiquimula; Escuintla; Quezaltenango ; Santa Rosa; Suchitepe"quez.
Honduras; Nicaragua; Costa Rica.
Large, often subscandent shrubs or small trees to 9 m. tall; branchlets slender,
obtusely tetragonal or subterete, often compressed and somewhat widened at the
the nodes, glabrous or puberulent; leaves on petioles 5-10 mm. long, the blades
membranaceous, lanceolate, ovate- oblong or oblong, 6-20 cm. long, acuminate,
rounded or acute at the base, very minutely puberulent to glabrate above and
below, minutely glandular-punctate beneath, the lateral veins 9-12 pairs; in-
florescences cymose, appearing paniculate, small or large, usually 9-20 cm. long,
the cymes many-flowered, the pedicels 2-3 mm. long at anthesis; calyx obpyrami-
dal at anthesis, 2-3 mm. long, truncate or subtruncate, usually minutely puberu-
lous, in fruit cupuliform, indurated, the margin truncate or subtruncate, 3-6 mm.
long, 7-10 mm. broad; corolla cream-colored; mature fruits subglobose to barrel-
shaped, orange, glabrous, to 1 cm. long.
Although the type specimen of A. magnified has larger leaves than
some specimens of A. laxicupulis, Moldenke's two species differ in
no other respect if one may judge from specimens determined by him.
Whether or not the calyx closely invests the fruit seems to depend
upon the stage of growth and development of the plant. A. laxi-
cupulis has also been confused by Moldenke and others with A. pa-
niculata Moldenke, a tree of the lowlands of southern Central
America and South America, which differs in its patelliform fruiting
calyces that are only 2-2.5 mm. long. As I am unable to separate
flowering material of any of these taxa from that of A. martinicensis
Jacq., perhaps they should all be placed there; however, the fruiting
calyces of A. martinicensis are lobate.
Aegiphila martinicensis Jacq. Obs. Bot. 2: 3, t. 27. 1767. A.
panamensis Moldenke, Trop. Woods 25: 14. 1931. A. glandulifera
Moldenke, Brittonia 1: 187. 1932. Chiploque (Retalhuleu).
FIG. 31. Aegiphila laxicupulis. A, habit, ^ natural size; B, long-styled
flower, X 3; C, calyx of long-styled flower opened to show pistil, X 3; D, stamens
of short-styled flower, X 5; E, short-styled flower, X 3; F, detail of stamens, X 10.
173
174 FIELDIANA: BOTANY, VOLUME 24
Damp or dry thickets or lowland forest, often in second growth,
30-1,060 meters; Escuintla; Quezaltenango; Retalhuleu; Suchitepe"-
quez; Santa Rosa. Southern Mexico; El Salvador to Panama; West
Indies; South America.
Shrubs or trees, sometimes 15 m. tall, sometimes more or less scandent, the
branchlets compressed and often widened at the nodes, glabrous or puberulent;
leaves on petioles 6-25 mm. long, the blades lanceolate, lance-ovate, lance-
oblong, or ovate-oblong, 6-20 cm. long, 3-8 cm. wide, long acuminate or abruptly
acuminate, rounded to acute at the base, entire, sometimes somewhat asym-
metric, glabrous or very minutely puberulent, minutely glandular-punctate be-
neath, the lateral nerves 8-16 pairs; inflorescences often large, cymose-panicu-
late, pedicels 2-5 mm. long, the flowers very numerous; calyx obpyramidal at an-
thesis, 2-3.5 mm. long, truncate or minutely apiculate or irregularly lobed, in fruit
often much enlarged, 2-5 mm. long, 6-10 mm. wide, shallowly cupuliform or patel-
liform, shallowly or deeply lobed, rarely merely repand; corolla white, cream-
colored, pale greenish-yellow or reddish-yellow, the tube 2-6 mm. long, 1-2 mm.
wide, minutely puberulous, the 4 lobes oblong lanceolate, 2-4 mm. long; fruit
7-10 mm. long, 6-8 mm. broad, glabrous, orange, with 4 seeds.
The two species described by Moldenke, although far separated
in his key, differ in no respect from A. martinicensis or from each
other if one may judge from specimens determined by him.
A. martinicensis f. falcata (Donn.-Sm.) D. Gibson, Fieldiana:
Bot. 32. (11): 176. 1970. A. falcata Donn.-Sm. Bot. Gaz. 18: 7. 1893.
Differs from A. martinicensis only by its larger leaves, 14-30 cm.
long, 6-14 cm. wide, and in its often more conspicuously thickened
and flattened nodes of the branchlets.
Aegiphila monstrosa Moldenke, Trop. Woods 25: 12. 1931.
Hulub (British Honduras).
Wet thickets or thin forest, often in second growth, 300 meters
or less; Alta Verapaz ; Izabal ; Pete"n. Mexico (Oaxaca) ; British Hon-
duras; Atlantic lowlands of Honduras.
Shrubs or trees, sometimes 9 m. tall but usually lower, the trunk often branched
from the base, the branches very soft and brittle, the younger ones very thick,
whitish or ochraceous, obtusely tetragonal, glabrous or minutely puberulent; leaves
on petioles 2-7.5 cm. long, the blades membranaceous, oblong to elliptic or elliptic-
obovate, 12-35 cm. long, 7-23 cm. wide, acute or subacuminate, acute or attenuate
to the base, minutely puberulent or in age glabrate, the lateral veins about 10
pairs; inflorescences sessile, cymose, the cymes numerous, usually at defoliated
nodes, very dense and glomerate, many- flowered, the flowers fragrant, on pedicels
1-2 mm. long; calyx campanulate, 2-3.5 mm. long and broad at anthesis, minutely
pubescent or almost glabrous, subtruncate, in fruit cupuliform, much enlarged and
indurated, verrucose; corolla white, the tube 8-10 mm. long, glabrous or minutely
GIBSON: FLORA OF GUATEMALA 175
puberulent, the 4 lobes 3-4 mm. long; fruit subglobose or ovoid, 7-10 mm. long and
broad, glabrous.
Aegiphila pauciflora Standley, Trop. Woods 16: 41. 1928.
Known only from the type, Indian guamil camp VI, Vaca, West-
ern Cayo District, British Honduras, Duncan Stevenson 5.
A tree, the branchlets obtusely tetragonal, puberulent or strigillose; leaves
chartaceous, on petioles 1.5-2 cm. long, the blades oblong-elliptic, 7-10 cm. long,
3-4 cm. wide, acuminate, cuneate at the base, entire, grayish-strigillose when
young, becoming glabrate, the lateral veins 7-8 pairs; cymes solitary, axillary,
lax, few-flowered, the peduncles 6-15 mm. long, the slender pedicels 1.5-3 mm.
long, strigillose; calyx broadly turbinate, 2.5-3 mm. long, grayish-strigillose, trun-
cate; corolla glabrous, the tube 3-4 mm. long, the lobes 2.5 mm. long; ovary
glabrous; fruits unknown.
Aegiphila skutchii Moldenke, Phytologia 1: 399. 1940.
Type collected at Finca Moca, Suchitepe"quez, in forest, 1,400
meters, A. F. Skutch 155. Mexico (Chiapas).
Trees to 15 m. tall, the trunk 50 cm. in diameter, the branchlets minutely
puberulent, the nodes 1-2 cm. apart; leaves on petioles 1-3 cm. long, the blades
membranaceous, elliptic-oblong, 8-15 cm. long, 1.5-5 cm. wide, acuminate, acute
at the base, entire, minutely puberulent, especially on the midrib and near the
base, or almost glabrous, the lateral veins 7-9 pairs; inflorescence 5-10 cm. long,
the cymes 4-9 cm. long, many-flowered, the peduncles 2-5 cm. long, the pedicels
1-2 mm. long, minutely puberulent; calyx at anthesis campanulate, truncate, 2-
3 mm. long, cinereous-puberulent, in fruit first urceolate, later cupuliform, irreg-
ularly torn, ca. 3 mm. long; corolla white, the tube 3 mm. long, the lobes 3 mm.
long; fruits apparently subglobose (only immature fruits seen).
Aegiphila standleyi Moldenke, in Fedde, Rep. Sp. Nov. 33:
140. 1933.
Damp forest, 1,000-1,250 meters; Sacatep£quez (Barranco Hondo,
Standley 65004). Costa Rica.
Large shrubs to 3 m. tall, the branchlets thick, glabrate; leaves on stout peti-
oles 1-2 cm. long, the blades obovate, thick, membranaceous, very large, as much
as 45 cm. long and 16 cm. wide, acute or short-acuminate, attenuate to the base,
finely and sparsely puberulent or glabrate, the margin somewhat repand-dentate
or entire, lateral veins about 20 pairs; inflorescences cymose, axillary, densely
flowered, the peduncles 6-10 mm. long, the pedicels ca. 3 mm. long; calyx at an-
thesis cupuliform, 3.5-4 mm. long, minutely puberulent, subtruncate, immature
fruiting calyx cupuliform to campanulate, 4 mm. long, verrucose; corolla not seen,
said to be glabrous, the tube said to be 1.5 mm. long and the 4 lobes 2 mm. long;
no mature fruits seen.
It should be noted that this record for Guatemala is based on a
sterile specimen annotated by Moldenke.
176 FIELDIANA: BOTANY, VOLUME 24
Aegiphila valerii Standl. Journ. Wash. Acad. Sci. 15: 481. 1925.
Not reported from Guatemala, but as it occurs in Chiapas, Mex-
ico (1,200 meters) as well as in Costa Rica (750-850 meters), it may
be expected in Guatemala.
Trees to 20 m. tall, the branchlets stout, covered with pale tomentum of short,
appressed hairs; leaves on petioles 1-2 cm. long, the blades obovate, mostly 7-
20 cm. long and 4-7.5 cm. wide, acute or short-acuminate, cuneate at the base,
entire, sparsely and minutely puberulent or glabrate, often more densely puberu-
lent on lower surface, lateral veins 10-16 pairs; inflorescences of short, axillary,
cymes, usually densely flowered, peduncles 4-10 mm. long, pedicels 1-3 mm. long,
both minutely appressed tomentose; calyx 3-5 mm. long, obconic, truncate, ap-
pressed tomentulose; corolla glabrous, the tube 3-5 mm. long, the 5 lobes ca. 3 mm.
long; fruits globose.
Often misidentified in herbaria as A. anomala Pittier, which has
flowers that are twice as large as those of A. valerii.
AVICENNIA L.
Trees or large shrubs of seashores, the branches nodose; leaves opposite, short-
petiolate, without stipules, the blades coriaceous, entire; inflorescences more or less
spicate, arising in the axils of the upper leaves, often forming pedunculate panicles;
flowers bracteate, small; calyx spreading, persistent, its 5 lobes imbricate, divided
nearly to the base; corolla broadly campanulate, the short tube cylindric, the limb
spreading, the 4 lobes subequal; stamens 4, the filaments inserted on the corolla
tube, the anthers introrse; ovary sessile, unilocular; ovules 4, on a central placenta;
style short, bilobate; fruit capsular, oblique, apiculate; seeds pendulous, without
endosperm, usually germinating in the capsule.
Three or four species, widely distributed in tropical regions in
coastal swamps. Although only one species is known from Guate-
mala, two are treated here, as A. bicolor Standley has been collected
in Mexico, El Salvador, Honduras, and Panama, and may be ex-
pected here.
This genus is placed by some authors in a separate family, Avi-
cenniaceae.
Leaves broadly elliptic to elliptic-ovate or ovate; flowers opposite, each pair 5-
8 mm. distant from the next pair on the rachis; corolla lobes glabrous inside;
style nearly obsolete, ca. 0.5 mm. long at anthesis A. bicolor.
Leaves oblong or oblong-lanceolate; flowers congested at ends of rachises; corolla
lobes sericeous inside; style 1-2 mm. long at anthesis A. germinans.
Avicennia bicolor Standl. Journ. Wash. Acad. Sci. 13: 354. 1923.
Swampy areas, sea level to 90 meters; Mexico (Chiapas); Hon-
duras; El Salvador; Costa Rica (fide Moldenke) ; Panama (type from
Aguadulce, Code", Panama, Pittier 4968).
GIBSON: FLORA OF GUATEMALA 177
Shrubs or trees to 13 m. tall; trunk to 30 cm. in diameter; young branchlets
glabrous; leaves on stout petioles 4-15 mm. long, the blades entire, broadly elliptic
to ovate, 5-13 cm. long, 3-7 cm. wide, rounded or obtuse at the apex, the base
usually cuneate or acute, often abruptly short-decurrent, glabrous above, densely
and finely pulverulent beneath; inflorescences spicate, tomentulose, the numerous
rachises forming pedunculate panicles 5-17 cm. long; the flowers opposite, sessile,
each pair 5-8 mm. distant from the next; bracts and bractlets ovate, usually
rounded and obtuse, tomentulose; corolla tube 2-3 mm. long, glabrous, the lobes
2-4 mm. long, obtuse to truncate, one or two often bifid, sericeous outside, gla-
brous inside; style 0.5 mm. long at anthesis, elongating to only 1 mm. by the time
the corolla falls; capsule obliquely oblong-ovate, 2-3 cm. long, pale green, very
finely pubescent.
This species closely resembles A. schaueriana Stapf & Leechman
of the West Indies and South America, which differs only in having
fewer, more congested, less complex inflorescences. If they should
prove to be synonymous, the name A. bicolor Standley takes pre-
cedence.
Called "mangle negro" in El Salvador.
Avicennia germinans (L.) L. Sp. PL ed. 3. 2: 891. 1764, pro
parte typica; P. Br. Civ. Nat. Hist. Jam. ed. 2, index I: 12, II: 7.
1789; Steam, Kew Bull. 1958: 34. 1958; Compere, Taxon 12: 150.
1963. Bontia germinans L. Syst. Nat. ed. 10: 1122. 1759; Sp. PL
ed. 2, 2: 891. 1763, pro parte typica. Avicennia nitida Jacq. Enum.
PL Carib. 25. 1760. A. africana P. Beauv. Fl. Oware 1 : 79-80, L 47.
1806. Black mangrove (British Honduras).
Common at or near sea level, in salt marshes, tidal flats, man-
grove swamps of both coasts; Escuintla; Izabal; Retalhuleu; San
Marcos. Florida; Mexico; British Honduras to Panama; along sea-
shores of South America and the Old World tropics.
Large shrubs or small trees, said to sometimes attain a height of 15 m., usually
smaller in Central America; the bark shallowly fissured, dark, orange-red within,
the young branchlets pale, densely and minutely puberulent; leaves on short, stout
petioles, the blades coriaceous, oblong or oblong-lanceolate, 3-12 cm. long, 1-4 cm.
wide, obtuse or acute, acute or attenuate to the base, grayish above and glabrous
or finely pubescent or puberulent, whitish or grayish beneath, and densely and
minutely pulverulent beneath, the veins usually conspicuous; inflorescences short,
forming panicles 2-5 cm. long, the flowers sessile, glomerate at the ends of the
rachises: bracts and bracteoles ovate-acute, imbricate; corolla greenish, cream, or
white, the tube 3-4 mm. long, glabrous, the lobes 3-5 mm. long, rounded, sericeous
inside and outside, one lobe often bifid; style usually 1-2 mm. long at anthesis,
sometimes attaining 3 mm. by the time the corolla falls; capsule obliquely oblong,
ovate, or oblong-ovate, laterally compressed, 2-3 (4) cm. long, pale green, finely
pubescent.
FIG. 32. Avicennia germinans. A, habit, l/i natural size; B, corolla dissected
to show stamens, X 4; C, calyx showing style and stigma, X 4; D, flower bud
showing subtending bracts and indicating sericeous pubescence, X 3.
178
GIBSON: FLORA OF GUATEMALA 179
Although the corollas have been described by some authors as
10 20 mm. long, I have seen none more than 8.5 mm.
This is one of the plants that characterize most tropical seashores.
In Guatemala it is one of the most abundant trees of coastal swamps.
The long, heavy roots grow in entangled arches, making almost im-
passable obstructions, and send to the surface of the mud large
masses of irregular asparagus-like aerial roots. The seeds usually
germinate on the tree and are growing and ready to take root when
they fall into the tidal mud. The flowers are said to be much visited
by bees and to supply a good grade of honey.
BOUCHEA Chamisso
References: Myrle Grenzebach, A revision of the genus Bouchea
(exclusive of Chascanum), Ann. Mo. Bot. Gard. 13: 71-90, t. 8-12.
1926; Harold N. Moldenke, A monograph of the genus Bouchea. II,
in Fedde, Rep. Sp. Nov. 49: 91-139. 1940.
Annual or perennial herbs, erect, glabrous or pubescent; leaves petiolate or
sessile, serrate, crenate dentate, or rarely incised, dissected, or entire; inflorescence
racemose or spicate, terminal or axillary, elongated, densely or laxly flowered,
bracteate; flowers short pedicellate, subsessile or sessile; bracts usually narrowly
linear-lanceolate; calyx persistent, tubular, 5-costate, the costae terminating in
more or less unequal teeth; corolla blue, lavender, rose, or purple, funnelform, the
tube straight or curved, the limb oblique, spreading, the 5 lobes unequal, the 2
posterior lobes shorter than the anterior ones; stamens 4, didynamous, included,
the filaments short, inserted on the corolla tube; anthers with 2 cells, ovate or sub-
cordate; ovary bilocular, each locule uniovulate; style filiform, the stigma bilobate;
fruit dry, linear, rostrate, included in the calyx or exserted, separating at maturity
into 2 cocci.
Ten species, one in Abyssinia, the others in tropical America.
Only the following species have been found in Central America.
Mature fruit 14-17 mm. long; beak 4-7 mm. long, densely puberulent; calyx con-
spicuously pubescent B. nelsonii.
Mature fruit 6-12 mm. long; beak less than 4 mm. long, glabrous or (rarely) only
sparsely and minutely puberulent; calyx puberulent or glabrate.
Fruit less than 10 mm. long.
Fruit 8-9.5 mm. long; beak ca. 1.5 mm. long B. prismatica.
Fruit 6-7 mm. long; beak 0.5 mm. long or less. .B. prismatica var. brevirostra.
Fruit 10-12 mm. long; beak 2-3 mm. long B. prismatica var. longirostra.
Bouchea nelsonii Grenzebach, Ann. Mo. Bot. Gard. 13: 83.
1926. Verbena.
Brushy, rocky slopes or plains, 150-800 meters; Huehuetenango;
Zacapa. Mexico (Oaxaca and Chiapas) ; Honduras.
180 FIELDIANA: BOTANY, VOLUME 24
Plants erect, stout, usually 16-60 cm. tall, rarely to 80 cm., simple or sparsely
branched, densely and finely pubescent throughout, the stems terete below, ob-
tusely tetragonal above; leaves on petioles 1-7 cm. long, the blades membrana-
ceous, broadly ovate, 2-6 cm. long, 1-4.5 cm. wide, acute or rounded at the apex,
broadly cuneate to subtruncate at the base, the margins coarsely serrate; racemes
terminal or axillary, up to 26 cm. long when mature, usually densely flowered, the
flowers short-pedicellate or subsessile; bracts linear, 5-6 mm. long; calyx very nar-
row, 7-9 mm. long at anthesis, 13-15 mm. long in fruit, pubescent, the teeth 2-
3 mm. long; corolla blue, rose, or purple; mature fruit 14-17 mm. long, linear, the
beak about one-third the total length, 4-7 mm. long, densely puberulent.
Bouchea prismatica (Jacq.) Kuntze, Rev. Gen. 2: 502. 1891.
Verbena prismatica Jacq. Coll. Bot. 2: 301. 1788. B. ehrenbergii
Cham. Linnaea 7: 253. 1832. Verbena.
Damp or wet fields, near sea level to 1,200 meters; Pete"n. South-
ern Arizona; Mexico; Honduras; West Indies; northern South
America.
Plants erect, to 60 cm. tall, simple or sparsely branched, the stems terete or
subtetragonal, pubescent or glabrate; leaves on petioles 1-6 cm. long, the blades
ovate to broadly ovate, 2-8 cm. long, 1-4.5 cm. wide, obtuse or acute, broadly
cuneate at the base, serrate to crenate-dentate, sparsely and finely pubescent on
both surfaces; racemes mostly terminal, 6-25 cm. long, densely or laxly flowered,
the flowers subsessile; bracts linear-lanceolate, 2-4 mm. long; calyx puberulent or
glabrate, 4-6 mm. long at anthesis, 5-9 mm. long in fruit, the teeth 1-2 mm. long;
corolla blue, rose, or purple; fruit linear, terete, 8-9.5 mm. long, the beak typically
1.5 mm. long and glabrous, rarely minutely puberulous.
Plants of this species usually wither when the rains end and are
seldom seen during the dry months.
B. prismatica var. brevirostra Grenzebach, Ann. Mo. Bot.
Gard. 13: 80. 1926 (type from Michoacan, Mexico, Arsene 2857}.
Shep-uon (Huehuetenango) .
Damp fields, grassy slopes, vacant lots, 800-1,900 meters; Hue-
huetenango; Santa Rosa. New Mexico; Mexico; El Salvador; West
Indies.
Differs from the typical variety in its shorter fruit, 6-7 mm. long,
shorter calyx, 4-6 mm. long, and especially in its very short, some-
times inconspicuous beak, 0.5 mm. long or less.
B. prismatica var. longirostra Grenzebach, Ann. Mo. Bot.
Gard. 13: 81. 1926 (type from Jamacia, Harris 11792}.
Damp fields, roadsides, 600 meters or less; British Honduras.
Southern Mexico; Honduras; West Indies; northern South America.
FIG. 33. Bouchea nelsonii. A, habit, H natural size; B, flower, X 3; C, calyx
enclosing fruit, X 3; D, detail of stigma, X 15; E, style and stigma, X 5.
181
182 FIELDIANA: BOTANY, VOLUME 24
Differs from the typical variety in its longer fruits, 10-12 mm.
long, fruiting calyx 8-11 mm. long, and much longer beak, 2-3 mm
long.
CALLICARPA L.
Reference: Moldenke, H. N. A monograph of the genus Calli-
carpa as it occurs in America and in cultivation, in Fedde, Rep. Sp.
Nov. 39: 288-317. 1936; 40: 38-131. 1936.
Trees or shrubs, the indument often of stellate or dendroid hairs; leaves oppo-
site, simple, without stipules; inflorescences cymose, axillary or supra-axillary;
flowers usually polygamous, small, actinomorphic; calyx at an thesis tubular or
campanulate, in fruit patelliform, with 4 lobes or teeth, or entire; corolla funnel-
form or salverform, the tube straight, ampliate above, the limb usually with 4 sub-
equal lobes, spreading; stamens 4 or rarely 5, the filaments filiform, glabrous,
inserted at or near the base of the corolla tube, usually exserted, anthers oblong or
elliptic, dorsifixed near the base, extrorse; style capillary, simple, glabrous, the
stigma capitate or peltate; ovary composed of 2 bilocular carpels, each carpel uni-
ovulate; fruit drupaceous, with a fleshy exocarp and hard endocarp, this separating
into 4 or by abortion fewer pyrenes; embryo straight; endosperm scant or none.
More than 100 species, widely dispersed in tropical regions of both
continents. Only one species is known in Central America.
Callicarpa acuminata HBK. Nova Gen. & Sp. 2: 252. 1817.
Ceniciento (Pete"n) ; fruta de chacha (Izabal).
On sand bars along streams, in wet to dry thickets or open forest,
sometimes in second growth, near sea level to 1,200 meters; Alta
Verapaz; Chiquimula; Guatemala; Izabal; Pete"n; Zacapa. Mexico;
British Honduras along the Atlantic lowlands to Panama; south-
ward to Bolivia.
Shrubs or small trees, usually not more than 5 m. tall, the branchlets densely
tomentose with stellate and dendroid hairs; leaves on petioles 0.5-3 cm. long, the
blades membranaceous or chartaceous, broadly ovate to lance-oblong or oblong-
elliptic, 8-25 cm. long, 2.5-11 cm. wide, long- acuminate, acute or acuminate at the
base, closely serrate or subentire, dark green above in age and sparsely stellate
tomentose, at least along the veins, pale and densely tomentose with stellate and
dendroid hairs beneath; inflorescences of numerous pedunculate cymes, appearing
corymbiform, borne in the upper leaf axils, usually less than half as long as the
leaves, densely flowered; flowers slightly fragrant, the pedicels 1-3 mm. long;
calyx campanulate, ca. 1 mm. long, subtruncate, minutely papillate and very
sparsely pubescent; corolla white, the tube 1.5-2.5 mm. long, the lobes scarcely
1 mm. long; stamens exserted; ovary glabrous; style glabrous, long-exserted ; fruit
dark purple, subglobose, juicy, 2-4 mm. long.
In Yucatan, called "zacpukim" or "xpucyim" (Maya).
FIG. 34. Callicarpa acuminata. A, section of flowering branch,
size; B, flower, X 5; C, calyx opened to show pistil, X 10.
natural
183
184 FIELDIANA: BOTANY, VOLUME 24
CITHAREXYLUM L.
Trees or shrubs, the branchlets usually tetragonous, rarely armed with spines;
leaf scars often large, corky, and elevated; leaves opposite or verticillate, estipulate,
entire or dentate, usually with 1 or 2 or more glands at the base of the blade; in-
florescences racemose or spicate, axillary and terminal, commonly elongated and
many-flowered, erect or nutant; flowers small, each subtended by an inconspicu-
ous bract; calyx at an thesis tubular or cupuliform, in fruit enlarged and indurated,
cupuliform or patelliform, the margin truncate or with 5 teeth or lobes; corolla
funnelform or salverform, white or yellowish, the limb spreading, the 5 lobes sub-
equal, usually pubescent in the throat; stamens 4, didynamous, inserted at or
above the middle of the corolla tube, included, a fifth stamen represented by a
rudimentary staminode, filaments very short, the anthers ovate or sagittate, in-
trorse; style often thickened toward the apex, the stigma shallowly bifid; ovary
with 4 perfect or imperfect locules, each locule with one lateral anatropous ovule;
fruit drupaceous, exceeding the calyx, with juicy exocarp and hard endocarp, con-
taining 2 bilocular pyrenes, each with 2 seeds.
More than 100 species, all in tropical and subtropical America.
Only ten occur in Guatemala but additional species are known from
other parts of Central America.
Indument of stellate hairs.
Bracts of inflorescence minute and inconspicuous, 1-2 (3) mm. long.
C. mocinnii.
Bracts of inflorescence conspicuous, 4-25 mm. long.
C. mocinniii var. longibracteolatum.
Indument of simple hairs.
Corollas lavender, 8-11 mm. long; angles of branches usually narrowly winged
or at least winglike C. pterocladum.
Corollas white, 4-7 mm. long; angles of branches not winged nor winglike.
Calyx at anthesis somewhat striate to conspicuously multistriate.
Lower leaf surface glabrous or obscurely puberulent; calyx 3-3.5 mm. long.
C. hexangulare.
Lower leaf surface pubescent, pilose on costa and veins; calyx 5-7 mm. long.
C. guatemalense.
Calyx at anthesis 5-costate.
Calyx conspicuously 5-lobate, even in fruit, the lobes ca. 1 mm. long; costa
and veins of lower leaf surface pilose with hairs ca. 1 mm. long.
C. cooperi.
Calyx sinuately 5-dentate, remotely denticulate, irregularly torn or trun-
cate; costa and veins of lower leaf surface not pilose.
Calyx at anthesis 4-7 mm. long (4 mm. in the short-styled flowers, 4.5-7
mm. in the long-styled flowers); fruiting calyx campanulate, closely
investing the mature fruit for half its length or more. .C. hirtellum.
Calyx at anthesis less than 4 mm. long; fruiting calyx cupuliform to pa-
telliform, not closely investing the mature fruit.
Lower leaf surface densely and finely pubescent; calyx more or less
pubescent C. crassifolium.
Lower leaf surface glabrous or only minutely puberulent; calyx gla-
brous or minutely puberulent.
Leaf blades mostly broadest below the middle, usually long-acumi-
nate at apex; calyx sinuately 5-dentate or denticulate.
C. donnell-smithii.
GIBSON: FLORA OF GUATEMALA 185
Leaf blades mostly broadest at or above the middle, obtuse, acute,
or short-acuminate at apex; calyx truncate.
Leaves typically 3 to 5 times as long as broad, apex usually obtuse,
sometimes acute; rachis of inflorescence glabrous.
C. caudatum.
Leaves typically about twice as long as broad, apex usually short-
acuminate: rachis of inflorescence puberulent.
C. steyermarkii.
Citharexylum caudatum L. Sp. PI. ed. 2. 872. 1763. C. erectum
Sw., Prodr. Veg. Ind. Occ. 91. 1788; Jacq., Ic. PI. Rar., ed. 2, 3. t. 501.
1793. C. macradenium Greenm. Field Mus. Bot. 2: 188. 1907.
Wet thickets, mangrove swamps, often in second growth, some-
times on pine ridges, at or near sea level; Izabal. Mexico (Chiapas
and Tabasco); British Honduras to Panama, along the Atlantic
coast; West Indies; northern South America.
Glabrous shrubs or trees, sometimes 10 m. tall (rarely to 20 m.) but often
flowering when only 1-2 m. tall, the trunk said to attain as much as 13 cm. in
diameter; branches and branchlets sub terete to tetragonous; leaves on petioles
0.5-2 cm. long, the blades more or less coriaceous, lustrous above, elliptic or nar-
rowly elliptic-oblong, usually broadest above the middle, 5-17 cm. long, 1.5-5.5
cm. wide, usually obtuse, but some leaves often acute, acute at the base, the basal
margins usually recurved and enclosing 2 dark, elongated glands; racemes axillary
or terminal, erect or pendant, solitary or forming lax panicles, 3-40 cm. long, usu-
ally densely flowered; rachis glabrous; pedicels 1-2.5 mm. long, the bracts minute,
triangular or subulate; calyx campanulate, 2-3.5 mm. long, glabrous, 5-costate,
subtruncate, remotely denticulate, in fruit broadly and shallowly cupuliform;
corolla white, the tube 3-4 mm. long, glabrous, the lobes 1-2 mm. long, glabrous
or minutely puberulent outside, puberulent inside; ovary glabrous, the style
minutely puberulent; fruit ovoid or subglobose, lustrous, yellow or orange to red at
first, purple black at maturity, to 12 mm. long.
Called "bird-seed" and "pigeon-feed" in British Honduras.
Citharexylum cooperi Standl. Trop Woods 10: 50. 1927.
In Guatemala, known only from Izabal, along Rio Tameja, alt.
50 meters. Panama.
Small trees, 7-8 m. tall, the branchlets quadrangular, pubescent or glabrate;
leaves on petioles 6-15 mm. long, the blades ovate to oblong-elliptic, 6-14 cm. long,
acuminate, acute and sometimes decurrent to the base, entire, subcoriaceous, essen-
tially glabrous above, softly pubescent below, pilose along the costae and veins
below, lateral veins 5-9 pairs; inflorescence paniculate, the racemes branching,
2-10 cm. long, densely flowered, the rachis short hirsute to hirtellous; bracts subu-
late, equalling or exceeding the pedicels; calyx narrowly campanulate, 3-3.5 mm.
long, sparsely hirtellous, 5-costate, the 5 lobes ovate; corolla tube ca. 2.5 mm. long,
glabrous outside, the lobes minutely puberulent on both surfaces; fruits subglo-
bose, 6-7 mm. long.
186 FIELDIANA: BOTANY, VOLUME 24
Called "wild lime" in Panama.
Citharexylum crassifolium Greenm. Field Mus. Bot. 2: 186.
1907.
Damp or dry mixed forest, 1,500-2,000 meters; Baja Verapaz
(type from Santa Rosa, Tuerckheim 1308}; Chimaltenango; Zacapa.
Mexico (Chiapas) ; British Honduras.
Shrubs or small trees, sometimes 9 m. tall, the branches subterete or tetrago-
nous, stout and stiff, the young ones usually densely pubescent; leaves on petioles
0.5-2 cm. long, the blades rigid-coriaceous, oblong, oblong-elliptic, or obovate,
5-15 cm. long, 2-8 cm. wide, entire or rarely (probably abnormally) coarsely ser-
rate near the apex, short acuminate to obtuse, acute at the base or rounded and
abruptly decurrent, essentially glabrous above, densely and finely pubescent be-
neath with spreading hairs, the veins elevated, the lateral ones 5-8 pairs; racemes
simple, 5-15 cm. long, usually few flowered, the rachis hirtellous or puberulent;
the bracts minute; pedicels 1-3 mm. long; calyx at anthesis narrowly campanulate,
2-3 mm. long, more or less pubescent, with 5 costae terminating in 5 very short
teeth, fruiting calyx broadly and shallowly cupuliform, truncate; corolla white,
the tube 3-4 mm. long, glabrous, the lobes 1.5-2 mm. long, pubescent on both sur-
faces; fruit obovoid-globose, 7-8 mm. long or more, first yellow, doubtless black
at maturity.
Citharexylum donnell-smithii Greenm. Field Mus. Bot. 2:
186. 1907. C. recurvatum Greenm. op. cit. 189 (type from Costa Rica,
Cooper 5889}. Coralillo (Guatemala and Sacatepe"quez) ; cuul, chuul
(Quezaltenango).
Damp, mixed forest, sometimes in oak and pine forest, often seen
along roadsides, sometimes planted, 1,000-2,700 meters; Escuintla;
Guatemala (type from Volcan de Pacaya, J. D. Smith 1879}; Que-
zaltenango; Sacatep£quez ; San Marcos; Suchitepe'quez; Zacapa.
Southern Mexico; Honduras; El Salvador; Costa Rica; Panama.
Trees, glabrous throughout, often 15 m. tall with a trunk to 70 cm. in diam-
eter, sometimes flowering when only 2 or 3 m. tall, the branches subterete; leaves
on petioles 1-3 cm. long, the blades coriaceous, lustrous, lanceolate or lance-oblong,
usually broadest below the middle, 8-20 cm. long, 2-5.5 cm. wide, long acuminate,
acute at the base, glabrous on both surfaces; racemes 5-35 cm. long, often forming
lax panicles; rachis usually glabrous, sometimes puberulent; pedicels 1-3 mm. long,
the bracts minute, subulate; flowers numerous; calyx tubular-campanulate, 2-3.5
mm. long, 5-costate, sinuately 5-dentate, ciliolate, in fruit cupuliform to patelli-
form; corolla greenish white to cream, the tube ca. 4 mm. long, glabrous outside,
the lobes 1-2 mm. long, glabrous outside, densely pubescent within; ovary gla-
brous, the style glabrous or minutely puberulent; fruit oblong-globose, 6-7 mm.
long, first yellow-orange to red, purple black at maturity.
FIG. 35. Citharexylum donnell-smithii. A, habit, ^ natural size; B, flower,
X 10; C, calyx opened to show pistil, X 10; D, corolla opened to show stamens
and staminode, X 10; E, fruit, X 2.
187
188 FIELDIANA: BOTANY, VOLUME 24
Citharexylum guatemalense (Moldenke) D. Gibson, Field
Mus. Bot. 32. (11) : 176. 1970. C. hirtellum var. guatemalense Mold-
enke, Phytologia 17: 113. 1968.
Known only from two collections, Steyermark 41818, the type,
and Steyermark 4-1816, both from Izabal, along Rio Tameja, alt. 50
meters.
Shrubs or small trees 4-5 m. tall, the branchlets glabrous, tetragonous; leaves
on stout petioles 1-2 cm. long, the blades oblong to oblong-ovate, 10-20 cm. long,
4-8 cm. wide, mostly acuminate, some acute, entire, glabrous above, long hirsute
to pilose below, sparsely so except along costae and veins, the hairs to 1 mm. long;
racemes simple, 10-18 cm. long, somewhat pubescent, densely or remotely flowered;
bracts minute; pedicels ca. 2 mm. long; calyx 5-7 mm. long, striate, sparsely pubes-
cent or glabrate, conspicuously 5-lobate, each lobe ca. 1 mm. long; corolla white,
essentially glabrous outside, the tube 4-5 mm. long, the lobes ca. 2 mm. long,
pubescent within; ovary and style glabrous, the style 2-3 mm. long; fruit obovoid-
globose, orange at first, doubtless black at maturity.
Although the flowers of C. guatemalense resemble those of the
long-styled form of C. hirtellum Standley in size of calyx and corolla,
the style of C. hirtellum is 4-5 mm. long, its calyx is shallowly sinuate
and denticulate or irregularly torn, and its leaves are smaller and
hirtellous below rather than pilose.
Citharexylum hexangulare Greenm. Field Mus. Bot. 2: 187.
1907.
Wet, mixed forest near sea level to 350 meters; Alta Verapaz
(type from Cubilguitz, Tuerckheim 7765) ; Pete"n. Southern Mexico;
British Honduras; Honduras; Nicaragua.
Trees or shrubs to 12 m. tall, the trunk to 15 cm. in diameter; branchlets
mostly acutely quadrangular, the larger ones mostly hexangular, glabrous or nearly
so; leaves opposite or sometimes ternate, on slender petioles 0.5-2 cm. long, the
blades chartaceous, lanceolate or lance-oblong, sometimes elliptic, 4-16 cm. long,
1-5.5 cm. wide, acute or acuminate at each end, entire, often with 2 elongated
glands at the base beneath, glabrous above, glabrous or minutely and obscurely
puberulent below, lateral veins 5-10 pairs; racemes axillary or terminal, simple or
paniculate, mostly erect, 5-30 cm. long, many-flowered, the rachis glabrous or
puberulent; pedicels ca. 1 mm. long, the bracts subulate, equalling or exceeding
the pedicels; calyx tubular campanulate at anthesis, in fruit first cupuliform, some-
times nearly patelliform in age, 3-4 mm. long, somewhat striate to conspicuously
multistriate, essentially glabrous, with 5 teeth or nearly truncate and merely den-
ticulate, ciliolate; corolla white, the tube 3-4 mm. long, glabrous outside, pubescent
in throat, the lobes 1-2 mm. long, puberulent outside and inside; style pubescent;
fruits ovoid, 6-8 mm. long, first yellow, black when mature.
GIBSON: FLORA OF GUATEMALA 189
The leaves are quite variable; those of some young shoots often
very narrow, resembling those of Salix species, while more mature
stems may have large, broadly elliptic leaves.
A plant of Costa Rica and Panama, C. viride Moldenke, is very
much like C. hexangulare and may prove to be synonymous with it.
The type of C. viride, Cooper & Slater 157, from Panama, is a fruiting
specimen, and the species was described without flowering material.
Later collections of flowering material identified by Moldenke as
C. viride differ from C. hexangulare only in their slightly broader
calyces. Although the leaves of C. viride are described as densely
puberulent beneath, leaves of the type specimen are essentially gla-
brous beneath or only very minutely pubescent along the lower part
of the costae and in the axils of some veins.
Citharexylum hirtellum Standl. Field Mus. Bot. 4: 257. 1929.
On ridges and along creeks, at or near sea level ; Izabal ; British
Honduras (type from Tower Hills, Orange Walk District, J. S.
Karling 9} .
Shrubs or small trees, the branches acutely quadrangular, hirtellous, or becom-
ing glabrate; leaves on petioles 4-15 mm. long, the blades first chartaceous but
usually becoming coriaceous, elliptic or elliptic ovate to elliptic-oblong, acute or
acuminate, rarely obtuse, acute at the base, entire, 4-12 cm. long, 3-6 cm. wide,
with 5-7 pairs of lateral veins, finely hirtellous above at first, becoming glabrate,
sparsely or densely hirtellous below; racemes simple, pedunculate, erect or pendant,
4-14 cm. long, the rachis densely hirtellous, many flowered, the flowers of 2 forms;
pedicels ca. 1 mm. long at anthesis, 2 mm. long in fruit; bracts linear subulate,
1-2 mm. long, or the lower ones sometimes elongated and 3-5 mm. long; calyx
tubular campanulate, sparsely hirtellous, 5-costate at anthesis, remotely denticu-
late or irregularly torn, ca. 4 mm. long in the short-styled flowers, 4.5-7 mm. long
in the long-styled flowers, fruiting calyx campanulate, closely investing the fruit,
4-7 mm. long; corolla white, ca. 6 mm. long in the short-styled flowers, 7-8 mm.
long in the long-styled flowers, the tube glabrous outside, pubescent within,
the lobes minutely puberulent outside, puberulent within; ovary glabrous, style
minutely pubescent, those of the small flowers ca. 3 mm. long, those of the larger
flowers 4-5 mm. long; fruit ovoid, 8-10 mm. long, first red, doubtless black at
maturity.
This has sometimes been confused with C. cooperi Standley, per-
haps partly because C. hirtellum was not understood to be hetero-
stylous. The calyx of C. cooperi, which is 3-4 mm. long, is always
conspicuously and evenly dentate. In addition, the racemes of
C. cooperi branch to form short panicles, 2-8 cm. long, and the leaves
usually have 8 -10 pairs of lateral veins.
190 FIELDIANA: BOTANY, VOLUME 24
Citharexylum niocinnii D. Don, Edinb. New Phil. Journ. 11:
238. 1831 (type from Mexico). C. rugendasii Cham., Linnaea 7: 120.
1832. C. rugendasii var. endlichii Loes. in Fedde, Rep. Sp. Nov. 9:
367. 1911. C. lankesteri Moldenke in Fedde, Rep. Sp. Nov. 37: 229.
1934.
Damp mountain forest, 1,500-2,300 meters; Guatemala; Que-
zaltenango; El Quiche"; Sacatepe"quez. Southern Mexico; Honduras;
El Salvador; Nicaragua; Costa Rica.
Trees to 18 m. tall, to 60 cm. in diameter, the branches usually stout, tetrago-
nous, stellate-tomentose or in age glabrate; leaves on stout petioles 1-6 cm. long,
the blades thick membranaceous to subcoriaceous, broadly ovate, oblong-ovate,
oblong-elliptic, lanceolate-ovate, or lanceolate, acute to acuminate (rarely obtuse),
the base rounded, acute, or cuneate, margins normally entire but rarely coarsely
dentate near the apex, 6-30 cm. long, 3-15 cm. wide, glabrate above except along
the costae and veins, pale or yellowish beneath and usually densely stellate tomen-
tose, the lateral veins prominent beneath; racemes solitary or forming panicles,
erect or pendent, many flowered, 6-30 cm. long, densely stellate-tomentose, the
pedicels short and thick, the bracts subulate, to 3 mm. long; flowers fragrant; calyx
tubular-campanulate at anthesis, 2.5-4 mm. long, pubescent or glabrate, the 5
costae-terminating in 5 remote, very small teeth, fruiting calyx ultimately patelli-
form, 6-7 mm. broad; corolla white, to 7 mm. long, the tube exceeding the calyx,
glabrous outside or the lobes somewhat pubescent, densely pubescent within; ovary
glabrous, the style minutely puberulent; fruit first subglobose, becoming ovoid, lus-
trous, first orange or red, black in maturity.
This species is extremely variable in leaf form and size.
Citharexylum mocinnii var. longibracteolatum Moldenke,
Phytologia 1: 415. 1940. Cana (Quezaltenango).
Damp forest, 1,500-2,800 meters; Quezaltenango; Sacatepe"quez ;
San Marcos. Southern Mexico; Honduras.
Differs from C. mocinnii only in its persistent, accrescent, con-
spicuous bracts which may be either subulate or spatulate and are
4-25 mm. long.
Citharexylum pterocladum Donn.-Sm., Bot. Gaz. 33: 255.
1902.
Dense, wet or dry mixed forest, sometimes in Cupressus forest,
sometimes in coffee plantations, 200-1,000 meters; Alta Verapaz
(type from Cubilgiiitz, Tuerckheim 7922)', Chimaltenango; Pete'n;
Quezaltenango. Southern Mexico; British Honduras.
Glabrous trees, often to 30 m. tall and to 40 cm. in diameter, the crown loosely
branched, open, the bark rough, brown, shed in small scales; young branches very
thick, acutely quadrangular, the angles narrowly winged or at least winglike, the
GIBSON: FLORA OF GUATEMALA 191
ultimate branchlets often compressed; leaves on slender petioles 1-8 cm. long,
the blades membranaceous, oblong-elliptic to ovate-oblong or lance-oblong, 6-29
cm. long, 3-14 cm. wide, acute to long-acuminate, usually cuneate-attenuate to
the base but sometimes nearly rounded; racemes usually branching and paniculate
but may be solitary, mostly 15-40 cm. long, many-flowered, very slender and lax,
apparently somewhat pendent; pedicels ca. 2 mm. long, bracts subulate- triangular,
very short; calyx tubular-campanulate, 5-costate, nearly truncate, remotely re-
pand-denticulate, 3-4 mm. long, in fruit indurated, 5-6 mm. broad; corolla pale
lavender with darker purple stripes, the tube 5-6 mm. long, glabrous outside, the
lobes 4-5 mm. long, glabrous except within at the base; ovary and style glabrous;
fruit yellow, probably black at maturity, ovoid-globose, about 8 mm. long.
Closely related to the Mexican C. affine D. Don. Moldenke sep-
arates the two by such tenuous characters as whether the branches
are "always alate" or "sometimes alate" and the leaf blades "glandu-
liferous at the base" or "usually not glanduliferous at the base."
The only constant difference is the smaller size of the flowers of
C. affine, in which the corolla tube is only ca. 4 mm. long with lobes
2-3 mm. long. Because they are so much alike in every other respect,
the flowers of both were examined for heterostyly but in both, the
stigma at anthesis is just a little below the anthers.
Citharexylum steyermarkii Moldenke, Phytologia 2: 14. 1941.
Dense, wet cloud forest, 2,000-2,600 meters, Chiquimula (type
from Volcan de Quezaltepeque, Steyermark 314.33); Zacapa.
Shrubs or small trees to 8 m. tall, the branches quadrangular or subterete, gla-
brous, lustrous, the younger branchlets obscurely puberulent; leaves on stout
petioles 0.5-2 cm. long, the blades coriaceous or thick-chartaceous, oblong-elliptic,
6-18 cm. long, 3-8 cm. wide, short-acuminate or acute, acute at the base, entire,
essentially glabrous above, glabrous or minutely puberulent below, the veins prom-
inent beneath, the lateral veins 7-8 pairs; racemes simple, 6-18 cm. long, many-
flowered, the rachis minutely puberulent; bracts minute; pedicels 1-3 mm. long;
calyx campanulate, ca. 3 mm. long, truncate, glabrous or puberulent and ciliolate,
in fruit shallowly cupuliform, lustrous; corolla white, the tube 3-4 mm. long, gla-
brous outside, the lobes 2 mm. long, puberulent outside, densely pubescent inside;
fruit 8 mm. long or more, subglobose, greenish-orange, doubtless black at maturity.
It should be noted that the specimen from Zacapa, Steyermark
428^5, is atypical. Although Moldenke previously identified it as
C. crassifolium Greenm. the lower leaf surface is not densely pubes-
cent, but is minutely and obscurely puberulent. Some of the leaves
are three times as long as broad, as in C. caudatum L., but they are
short-acuminate at the apex and the rachis of the inflorescence is
puberulent as in C. steyermarkii. It may be that C. steyermarkii is
only a glabrate form of C. crassifolium, or both may represent only
a broad-leaved form of C. caudatum L.
192 FIELDIANA: BOTANY, VOLUME 24
CLERODENDRUM L.
Plants herbaceous or woody, often trees or shrubs, sometimes scandent, gla-
brous or pubescent; leaves opposite or verticillate, the margins entire or dentate;
inflorescences cymose, the cymes borne in the upper leaf axils or (in C. fragrant
Vent., an exotic species) diposed in terminal, often headlike panicles, often bracte-
ate; flowers more or less asymmetric, often showy, white to blue, violet, or red;
calyx usually campanulate, rarely tubular, subtruncate or with 5 teeth or lobes,
often accrescent and subtending or enclosing the fruit; corolla salverform, the tube
straight or curved, often much elongated, the limb 5-parted, the lobes subequal or
the 2 posterior ones shorter; stamens 4, didynamous, inserted in the corolla tube,
long-exserted ; anthers ovate or oblong, the cells parallel, opening by longitudinal
slits; ovary bicarpellate, imperfectly 4-locular, each locule uniovulate, the ovules
inserted laterally near the apex of the locule; the style terminal, stigma bifid, the
lobes short; fruit drupaceous, globose or obovoid, often somewhat 4-lobate, the
exocarp fleshy, the endocarp bony or crustaceous, smooth or rugose, separating at
maturity into 4 pyrenes, or these sometimes coherent in pairs; seeds oblong, with-
out endosperm.
About 350 species in tropical regions of both hemispheres, most
abundant in Asia and Africa, with only a few species in America.
Two species are native in Guatemala, but three are treated here, as
C. fragrans Vent., a native of Asia, has become established in many
places. Four plants previously reported from Guatemala as Cleroden-
drum are omitted, as C. standleyi Moldenke has been referred to the
Acanthaceae, and C. pithecobium Standley & Steyermark, C. mimi-
cum Standley & Steyermark, and C. moldenkeanum Standley, all with
numerous ovules in bilocular ovaries, have been referred to the Scro-
phulariaceae.
Leaves almost as broad as long, truncate to deeply cordate or (rarely) rounded at
the base, the margins coarsely dentate; corollas double, the limb 2.5-3 cm.
across C. fragrans.
Leaves much longer than broad, acute to cuneate-acute at the base, the margins
entire or nearly so; corollas single, the limb less than 2 cm. across.
Calyx nearly truncate and dentate; leaves 1-4 cm. long, obtuse or rounded at
the apex, lateral veins 3-5 pairs, often obscure C. pittieri.
Calyx deeply 5-lobate, the lobes lanceolate and acuminate; leaves 2-10 cm. long,
acute to subacuminate at apex, lateral veins 5-9 pairs, conspicuous.
C. ligustrinum.
Clerodendrum fragrans Vent. Jard. Malm. 2, t. 70. 1804. C.
fragrans var. pleniflorum Schauer ex A. DC. in DC. Prodr. 11: 666.
1847. Boca de Amelia (Escuintla) ; camelia (Zacapa) ; jasmin (Hue-
huetenango); jasmin de Amelia (Alta Verapaz); Spanish jasmine
(British Honduras).
Native of Asia, widely cultivated in Guatemala and often natural-
ized, forming dense colonies in damp thickets, in waste ground, or
GIBSON: FLORA OF GUATEMALA 193
near dwellings, 200-1,800 meters, Alta Verapaz; Escuintla; Guate-
mala; Huehuetenango; Izabal; Sacatepe"quez ; Santa Rosa; Zacapa.
Also escaped in Mexico, British Honduras, El Salvador, Costa Rica
and West Indies.
Stout shrubs to 1.5 m. tall or almost wholly herbaceous, simple or sparsely
branched, the branches stout, obtusely angulate, pubescent; leaves on long, slender
petioles, the blades membranaceous, usually broadly ovate, 6-25 cm. long, 5-25
cm. wide, acute or acuminate, the base usually cordate or truncate, sometimes
rounded, short-hirtellous, especially beneath, the margins irregularly and coarsely
dentate; inflorescences terminal, cymose, often headlike, densely flowered, sessile
or short-pedunculate; bracts numerous, oblong or elliptic, foliaceous; calyx cam-
panulate, 1-1.5 cm. long, red or purple, strigillose-puberulent or glabrate, the 5
lobes lanceolate, acuminate; corolla white, often tinged with pink or purple, in
Central American plants always double, the limb 2.5-3 cm. across.
One of the common ornamental plants of Central American gar-
dens, maintaining itself when neglected, and blooming through the
dry months.
All Guatemalan material is referable to var. pleniflorum.
Clerodendrum ligustrinum (Jacq.) R. Br. in Ait. Hort. Kew.
ed. 2. 4: 64. 1812. Volkameria ligustrina Jacq. Coll. Bot. Suppl. 118,
t. 5, f. 1. 1796. Clerodendron mexicanum Brandegee, Univ. Calif.
Pub. Bot. 3:391. 1909.
Thickets along stream banks or in forest margins, 50-250 meters;
Pete"n. Mexico; British Honduras; Nicaragua; Panama.
More or less scandent, woody shrubs, the branchlets slender or stout, subterete
or obtusely angulate, puberulent or glabrate, the foliar scars large and elevated:
leaves on petioles 5-14 mm. long, the blades thick-membranaceous, elliptic to ellip-
tic-oblong or elliptic-lanceolate, 2-10 cm. long, 1-5 cm. wide, acute or subacumi-
nate at apex, acute at base, entire, glabrous, densely punctate beneath, the lateral
veins 5-9 pairs; inflorescences axillary, sometimes appearing terminal, the cymes
pedunculate, often lax, 3-7 cm. long, few-flowered, puberulent or glabrate, the
bracts few, foliaceous, caducous, the pedicels slender, 3-6 mm. long; calyx cam-
panulate, 6-9 mm. long, deeply 5-lobate, the lobes lanceolate, acuminate, 3-5 mm.
long; corolla white, the tube 10-12 mm. long, the 5 lobes subequal, 4-8 mm. long;
stamens long-exserted ; fruit subglobose, ca. 1 cm. long, splitting into two 2-seeded
halves at maturity.
It is reported that in Pete"n the leaves are used in cooking to
flavor fish.
Clerodendrum pittieri Moldenke ex Standl. Field Mus. Bot.
18: 1003. 1938 (without Latin diagnosis); Moldenke, Phytologia 1:
416. 1940.
FIG. 36. Clerodendrum ligustrinum. A, habit, ^ natural size; B, corolla dis-
sected to show stamens, X 3; C, calyx and style, X 3; D, fruit, X 1.
194
GIBSON: FLORA OF GUATEMALA 195
Coastal thickets or mangrove swamps; Escuintla; San Marcos.
Costa Rica; Panama.
Shrubs of 1-2 m. with stout branches, the young branchlets puberulent or
strigillose; leaf scars thickened and pyramidal, often somewhat spinelike; leaves on
petioles 1-8 mm. long, the blades subchartaceous, oblanceolate-oblong to narrowly
elliptic or obovate, 1-4 cm. long and 0.4-2 cm. wide, rounded or obtuse at the apex,
cuneate-acute at the base, entire, glabrous, densely punctate beneath, the lateral
veins 3-5 pairs, often obscure; inflorescences of solitary axillary cymes, 1-3-flow-
ered, the peduncles to 1 cm. long, the pedicels 5-8 mm. long; calyx 3-5 mm. long,
minutely puberulent or glabrate, truncate or nearly so and 5-dentate; corolla white,
the tube 2 cm. long, slender, the lobes 4-6 mm. long; fruit subglobose, ca. 5 mm.
long.
Closely related to C. aculeatum (L.) Schlecht., which is common
from eastern Mexico through the West Indies to northern South
America, and which has a smaller calyx tube with definite calyx
lobes. It should be noted that no flowering material of C. pittieri
from Guatemala was seen; the only two Guatemalan collections,
both identified by Moldenke, are poor fruiting specimens with only
fragmentary calyces and with leaves considerably larger than those
of the type of C. pittieri. Both might easily be placed in C. aculea-
tum. It is also interesting that a Field Museum photograph (Neg.
No. 24622), of a specimen collected in Peru by Ruiz and Pavon and
identified by Moldenke as C. buxifolium (Willd.) Spreng, has the
same calyx and corolla as the type of C. pittieri.
CORNUTIA Plum, ex L.
Reference: Harold N. Moldenke, A monograph of the genus Cor-
nutia, in Fedde, Rep. Sp. Nov. 40: 153-205. 1936.
Trees or shrubs with brittle branches and usually abundant pubescence, the
branches tetragonous; leaves opposite, petiolate; inflorescences terminal, usually
pyramidal-paniculate, the flowers small, zygomorphic; calyx cupuliform to cam-
panulate, or obconic, often becoming patelliform in fruit, entire or minutely 4-den-
tate; corolla salverform, irregular, blue or purple, the tube often inflated at the
base and ampliate above, straight or curved, the limb spreading, bilabiate, with
4 lobes; 2 fertile stamens inserted in the corolla tube at about the middle, 2 stami-
nodes usually inserted at a point above the middle, the fertile ones usually exserted;
filaments usually pubescent, the anthers dorsifixed, the cells widely divergent at
the base, dehiscent by longitudinal slits; style terminal, simple, glabrous or pubes-
cent, the stigma bifid, the lobes short; ovary pubescent, composed of 2 bilocular
carpels, each locule uniovulate; fruits small, drupaceous, subglobose, the exocarp
fleshy and juicy, the endocarp bony, the stone with 4 locules; seeds without en-
dosperm.
196 FIELDIANA: BOTANY, VOLUME 24
About ten species, in tropical America, usually very showy and
attracting attention because of their abundant blue or purple flowers.
Two other species are known from southern Central America.
Mature corolla tube abruptly and strongly incurved, 2.5-5 mm. wide, broadest
near base and often somewhat saccate, the lower lip usually equalling the
abaxial side of the tube; style glabrous or minutely pubescent; branchlets,
leaves, and inflorescence usually densely velutinous-tomentose.
C. grandifolia.
Mature corolla tube somewhat incurved or nearly straight, 1-3 mm. wide, usually
broadest near base but never saccate, the lower lip usually not more than half
as long as the abaxial side of the tube; style conspicuously short-pubescent;
branchlets, leaves, and inflorescence short-pubescent, puberulent, or tomentu-
lose, often somewhat viscid C. pyramidata.
Cornutia grandifolia (Schlecht. & Cham.) Schauer in DC.
Prodr. 11 : 682. 1847. Hosta grandifolia Schlecht. & Cham. Linnaea 5:
97. 1830. C. pyramidata var. dentata Kuntze, Rev. Gen. 2: 506. 1891.
C. grandifolia var. quadrangular is Moldenke in Fedde, Rep. Sp. Nov.
40: 168. 1936. C. grandifolia var. purpusi Moldenke, op. cit. 169.
C. grandifolia var. storkii Moldenke, op. cit. 169.
Damp thickets or dense forest, often in second growth, 150-1,300
meters; Alta Verapaz; Guatemala; Izabal; Sacatepe"quez. Mexico;
British Honduras to Panama.
Shrubs or weak trees to 5 m. tall, the branchlets densely velutinous-tomentose
with spreading hairs; leaves on stout petioles 1-5 cm. long, the blades ovate, ellip-
tic-ovate or oblong-ovate, 7-30 cm. long, 5-19 cm. wide, acuminate, attenuate to
the base and decurrent, usually densely velutinous-tomentose on both surfaces,
more densely so beneath (rarely only sparsely puberulent), the margins usually
dentate, rarely entire; inflorescences terminal, paniculate, 15-40 cm. long, the
flowers numerous, short-pedicellate, the cymes pedunculate, rachis, peduncles and
pedicels usually velutinous-tomentose; calyx at anthesis shallowly cupuliform
and truncate, sometimes obscurely toothed, 1-2.5 mm. long, densely short-pubes-
cent, in fruit usually patelliform, sometimes shallowly cupular; corolla light blue
to deep purple, puberulent, the tube abruptly and strongly incurved, 6-9 mm.
long, 2.5-5 mm. wide, broadest near the base and often more or less saccate, the
lower lip of the limb almost as long as the abaxial side of the tube; ovary densely
pubescent, style glabrous or minutely pubescent; fruits subglobose, 3-5 mm. long,
puberulent or pubescent.
Cornutia pyramidata L. Sp. PI. 628. 1753. Hosta pyramidata
A. Dietr. in Willd. Sp. PI. 1, 253. 1831. Hosta latifolia HBK. Nova
Gen. & Sp. 2: 248. 1817. Cornutia latifolia Moldenke in Fedde, Rep.
Sp. Nov. 40: 179. 1936. C. grandifolia var. intermedia Moldenke,
op. cit. 167. C. lilacina Moldenke, op. cit. 181 (type from Gualan,
Guatemala, Deam 6383). C. lilacina var. velutina Moldenke, op. cit.
183. C. pyramidata var. isthmica Moldenke, op. cit. 187. C. lati-
FIG. 37. Cornutia grandifolia. A, flowering branch, '2 natural size; B, flower,
X 3; C, calyx and pistil, X 3; D, corolla dissected to show stamens and stami-
nodes, X 3.
197
198 FIELDIANA: BOTANY, VOLUME 24
folia f. alba Moldenke, Phytologia 2: 131. 1946. Flor lila (Guate-
mala) ; hoja de zope (Izabal) ; lat-che (Maya, Pete"n) ; tzultesnuk and
matasano (British Honduras).
Damp forest or thickets, often in second growth, 100-1,500 me-
ters; Alta Verapaz; Chimaltenango; Chiquimula; El Progreso; Es-
cuintla; Guatemala; Izabal; Pete"n; Quezaltenango; Sacatepe"quez ;
San Marcos; Santa Rosa; Solola; Suchitepe"quez; Zacapa. Mexico;
British Honduras; El Salvador and Honduras to Nicaragua; West
Indies.
Large shrubs or weak trees to 12 m. tall, the trunk reported as sometimes
15 cm. in diameter, the branchlets densely short-pubescent, puberulent, or tomen-
tulose, often somewhat viscid; leaves on petioles 0.5-3 cm. long, the blades elliptic,
elliptic-ovate, or broadly ovate, 4-20 cm. long, 4-14 cm. wide, acute or acuminate,
usually attenuate (rarely acute) to the base and decurrent, the margins repand-
denticulate or entire, both surfaces densely short-pubescent or puberulent or the
lower surface tomentulose; inflorescences terminal or sub terminal, paniculate, 10-
40 cm. long, 5-10 cm. wide, the pedicels and peduncles of the cymes densely short-
pubescent or puberulent, often somewhat viscid; calyx cupuliform at anthesis,
1-3 mm. long, densely short- pubescent, truncate, sometimes obscurely toothed,
in fruit patelliform to shallowly cupular; corolla blue to purple, densely short
pubescent or puberulent, the tube 7-11 mm. long, 1-2.2 mm. wide, somewhat in-
curved or nearly straight, the lower lip of the limb usually half as long as the ab-
axial side of the tube, or shorter; ovary pubescent, style short-pubescent; fruits
subglobose, puberulent or pubescent, 3-6 mm. long.
The indument of West Indian plants is albidous, while that of
Central American species is usually fulvous, roseate, or purplish, but
they differ in no other respect.
Although specimens determined by Moldenke to be C. grandi-
folia var. intermedia Moldenke tend to have larger leaves and smaller
calyces, as in C. grandifolia, I have placed them in C. pyramidata
because of their corollas. Although straight and curved corolla tubes
can often be found on the same plant, they are usually straighter,
narrower and much less dilated than those of C. grandifolia and the
lower lip of the corolla is usually not more than half as long as the
tube. Their indument is also shorter, as in C. pyramidata.
Because there is such a wide range of variation in both C. grandi-
folia and C. pyramidata, some forms may appear quite different.
However, it is not always easy to separate even these two taxa. The
fruits are identical. Considerable variation in calyx length and co-
rolla tube width is sometimes seen on a single plant. Attempts to
describe separately all the intermediate representatives of such vari-
able plants have resulted in the creation of species and varieties
separated only by inconstant and ill-defined characters.
GIBSON: FLORA OF GUATEMALA 199
DURANTA L.
Shrubs or small trees, glabrous or pubescent, sometimes armed with spines,
the branches often elongated and recurved or pendent; leaves opposite or verticil-
late, entire or dentate; inflorescences racemose, terminal or axillary, the flowers
bracteate; calyx tubular, truncate, 5-costate, each costa terminating in a small
tooth; corolla blue, lavender, or white, salverform, the tube exserted from the
calyx, the limb spreading, regular or oblique, usually pubescent within at the ori-
fice, the lobes 5; stamens 4, didynamous, included, inserted at or above the middle
of the corolla tube (occasionally a staminode present), the filaments very short;
anthers sagittate, dorsifixed, the cells parallel; ovary more or less completely
8-locular, composed of 4 bilocular carpels, each locule uniovulate; style terminal,
the stigma obliquely subcapitate; calyx accrescent in fruit, usually longer than and
often enclosing the fruit; fruit drupaceous, the exocarp fleshy, the endosperm hard,
nutlike; pyrenes 4, each bilocular, each containing 2 seeds.
About 35 species, in tropical America. There are two species in
Guatemala, and a third occurs in southern Central America.
Branches usually armed; branchlets, rachis, and pedicels copiously pilosulous or
tomentose; calyx 5-7.5 mm. long at anthesis D. guatemalensis.
Branches usually unarmed; branchlets, rachis, and pedicels appressed-pubescent,
appressed-puberulent, or glabrate; calyx 3.5-5 mm. long at anthesis.
D. repens.
Duranta guatemalensis Moldenke, Bull. Torr. Bot. Club 68:
501. 1941.
Dry, brushy or wooded slopes, 1,400-2,600 meters; El Quiche"
(type from Chiul, Heyde & Lux 2947); Huehuetenango (between
San Ildefonso Ixtahuacan and Cuilco). Mexico (Chiapas).
Shrubs or small trees to 4.5 m. tall, the branchlets densely tomentose or some-
times glabrate in age, armed with stiff spines 3-15 mm. long; leaves short-petiolate,
the blades oblanceolate, obovate, or elliptic, 1.5-6 cm. long, usually obtuse or
rounded at the apex, sometimes acute, cuneate and often decurrent at the base,
the margins entire or sparsely serrate near the apex, sparsely pilosulous above or in
age glabrate, paler beneath, puberulent-pilosulous; racemes few-many-flowered,
3-5 cm. long; bracts minute, 1-2 mm. long; pedicels 1-2 mm. long; calyx tubular,
accrescent, minutely 5-apiculate, 5-7.5 mm. long at anthesis, 2-3.5 mm. wide,
pubescent outside, densely so within; corolla puberulent outside, the tube 7-9 mm.
long, ca. 3 mm. wide, the limb 9-11 mm. across, the lobes densely pubescent within;
ovary and style glabrous; fruit yellow, subglobose, almost 1 cm. in diameter, com-
pletely enclosed in the yellow, glabrate calyx which is prolonged by the persistent
apiculum into a short, curved beak.
Duranta repens L. Sp. PI. 637. 1753. D. plumieri Jacq. Sel.
Stirp. Amer. 186. 1. 176, f. 76. 1763. D. repens var. alba L. H. Bailey,
Manual Cult. PI. 843. 1949. Coralillo rosado (Guatemala).
FIG. 38. Duranta repens. A, flowering branch, % natural size; B, flower, X 3;
C, calyx opened to show pistil, X 3; D, corolla dissected to show stamens and
staminode, X 3.
200
GIBSON: FLORA OF GUATEMALA 201
In thickets, secondary forest, hedges, along roadsides, 500-2,600
meters; Guatemala; Quezaltenango; Santa Rosa; Solola; Suchitepe"-
quez. West Indies; Mexico to northern South America.
Shrubs or small trees to 6 m. tall, usually unarmed, the branches tetragonous,
commonly long and pendent, the branchlets glabrous or appressed-pubescent or
puberulent; leaves short-petiolate, the blades elliptic to obovate, 2-8 cm. long,
obtuse or acute at the apex, cuneate and decurrent at the base, the margins entire
or serrate above the middle, glabrous or somewhat appressed-puberulent beneath,
especially on the costae and lateral veins; racemes 3-16 cm. long, glabrous or ap-
pressed-puberulent, laxly many-flowered, often recurved or pendent, the bracts
2-4 mm. long and linear, or the lower ones foliaceous and often exceeding the calyx;
pedicels 2-5 mm. long, usually appressed-puberulent; calyx tubular, accrescent,
3.5-5 mm. long at anthesis, 5-costate, the costae prolonged and forming apicula
0.5-1 mm. long, sparsely appressed-puberulent or glabrous outside, sericeous with-
in; corolla puberulent or essentially glabrous outside, the tube 7-8 mm. long, 1.5-
2 mm. wide, the limb 8-15 mm. across, the lobes usually puberulent within; ovary
and style glabrous; fruit yellow, globose, 7-12 mm. in diameter, completely en-
closed in the yellow calyx which is prolonged by the persistent apiculum into a
short, curved beak.
This species, which is frequent in cultivation in Guatemala, is
thought by some to have been introduced into Central America. If
this is true, it is certainly widely distributed and has become natural-
ized in a variety of places.
LANTANA L.
Erect, ascending, or scandent shrubs, the branches aculeolate or unarmed;
leaves opposite or verticillate; inflorescences spicate, the spikes often short and
headlike, densely flowered, the peduncles axillary, solitary or binary in the leaf
axils; flowers small, bracteate; calyx small, membranaceous, truncate or irregularly
sinuate- dentate; corolla white, pink, purple, yellow, or red, the tube cylindric,
slender, usually more or less minutely puberulent outside, often somewhat pubes-
cent within the throat, the limb with 4 or 5 lobes, the lobes obtuse or retuse; sta-
mens 4, didynamous, included, inserted at about the middle of the corolla tube;
anthers ovate, the cells parallel; ovary unicarpellate, bilocular, each locule uni-
ovulate; style terminal, stigma thick, oblique or sublateral; ovules basal and erect
or attached laterally near the base of the locule; fruit drupaceous, juicy, bilocular
or splitting into 2 unilocular pyrenes, the endocarp hard; seeds without endosperm.
Perhaps 50 species, mostly in tropical America, with five in Gua-
temala.
Many species are separated by poor and variable differences, only
a few of those found in North America being clearly differentiated.
Stems usually aculeolate; corolla yellow to orange, or red, the tube 7-10 mm. long.
L. camara.
202 FIELDIANA: BOTANY, VOLUME 24
Stems never aculeolate; corolla white, cream, pale pink, or purple, the tube 2-6 mm.
long.
Lowermost bracts of inflorescence lanceolate, acuminate to cuspidate.
Margins of leaf blades acutely serrate; leaves usually binate; peduncles often
twice as long as the leaves L. achy ranthi folia.
Margins of leaf blades crenate-serrate; leaves usually ternate; peduncles often
equalling or exceeding the leaves but not twice as long L. trifolia.
Lowermost bracts of inflorescence ovate, broadly ovate, or oblong-elliptic, acute
or obtuse.
Leaf blades subacute to acuminate at apex, margins crenate, indument com-
posed of erect or ascending hairs; lowermost bracts of inflorescence ovate
to broadly ovate L. hispida.
Leaf blades rounded at apex, margins closely crenulate, indument composed
of appressed hairs; lowermost bracts of inflorescence lanceolate to oblong-
elliptic L. involucrata.
Lantana achyranthifolia Desf. Cat. PL Paris ed. 3. 392. 1829.
L. purpurea Benth. & Hook. f. Gen. PI. 2: 1142. 1876. L. macropo-
dioides Greenm. Field Mus. Bot. 2: 339. 1912.
Damp or dry thickets, 500-800 meters; Huehuetenango. Mexico;
Honduras.
Erect, unarmed shrubs to 1.5 m. tall, the stems appressed-strigose; leaves lan-
ceolate to broadly ovate, short-petiolate, the blades 4-8 cm. long, acuminate to
long-acuminate, obtuse or acute at the base and often abruptly decurrent, acutely
serrate, strigose on both surfaces, pale beneath; inflorescence pedunculate, capitate,
the peduncles stiff, straight, much longer than the leaves and often twice as long
or more, the flower heads short at anthesis but elongating in age, often becoming
much longer than broad, in fruit sometimes 2.5 cm. long, 1.5 cm. in diameter;
bracts lance-ovate or lanceolate, acuminate, pale green, densely hispidulous or
strigose, the lowermost ones sometimes to 12 mm. long; corolla usually white but
may be purplish, the tube 3-4 mm. long, puberulent.
This is often confused with the Mexican L. macropoda Torrey,
which has smaller, less sharply serrate leaves, acute at the apex.
Lantana camara L. Sp. PL 627. 1753. L. aculeata L. loc. cit.
L. mista L. Syst. ed. XII, 2: 417. 1767. L. horrida HBK. Nova Gen.
& Sp. 2: 261. 1817. L. tiliaefolia Cham. & Schlecht. Linnaea 7: 122.
1832. L. hirsuta Mart. & Gal. Bull. Acad. Brux. 11: 326. 1844.
L. horrida var. grandiflora Schauer in DC. Prodr. 11: 598. 1847. L.
horrida var. parviflora Schauer in DC. loc. cit. L. camara var. crocea
Bailey, Cycl. Am. Hort. 884. 1900. L. camara var. mista (L.) Bailey,
loc. cit. L. glandulosissima Hayek in Fedde, Rep. Sp. Nov. 2: 161.
1906. L. crocea var. guatemalensis Loes. Verh. Bot. Ver. Brandenb.
53: 76. 1911 (type from Huehuetenango, Seler 2805). L. camara var.
macrantha Loes. loc. cit. L. camara var. aculeata (L.) Moldenke,
Torreya 34: 9. 1934. L. scorta Moldenke, Carnegie Inst. Wash. Pub.
FIG. 39. Lantana camara. A, flowering branch, V£ natural size; B, corolla,
X 3; C, corolla dissected to show stamens, X 3; D, calyx opened to show pistil,
X 10.
203
204 FIELDIANA: BOTANY, VOLUME 24
522. 161. 1940. L. scandens Moldenke, Phytologia 2: 18. 1941. L.
hispida var. ternata Moldenke, Phytologia 2: 225. 1947. L. camara
var. parvifolia Moldenke, Phytologia 2: 467. 1948. L. hirta f. ter-
nata Moldenke, Phytologia 8: 160. 1962. Chiligua nigrita (Jalapa);
cinco negritos (Guatemala, Retalhuleu, Sacatepe'quez) ; ek-cuaiyak
(Alta Verapaz) ; mora de muerto (Alta Verapaz) ; sincuria (Izabal) ;
vivarana (Guatemala) .
Common in wet to dry thickets or open forest, sometimes in pine
forest, abundant in second growth, frequently on limestone, a weed
of banana plantations, often invading cultivated ground, sea level to
2,200 meters; Alta Verapaz; Baja Verapaz; Chimaltenango; Es-
cuintla; Guatemala; Huehuetenango; Izabal; Jalapa; Pete"n; Que-
zaltenango; Retalhuleu; Sacatepe'quez ; San Marcos; Santa Rosa;
Solola; Zacapa. Southern Florida; Mexico; British Honduras to El
Salvador and Panama. West Indies and South America; naturalized
in the Old World tropics.
Shrubs, usually 1-3 m. tall, often subscandent, the stems then attaining greater
lengths, stems usually aculeolate, rarely unarmed, pilose or hirsute with gland-
tipped or eglandular hairs; leaves opposite, binate or rarely ternate, petiolate,
broadly ovate to oblong-ovate, 2-12 cm. long, the margins crenate-serrate, the
apex acute or short-acuminate, acute or rounded at the base and abruptly decur-
rent or cordate; often bullate or rugose, scabrous to scabrous -tomentose above,
variously pubescent beneath (densely viscid-tomentose to strigose or rarely gla-
brate); inflorescences densely flowered heads, the peduncles 2-14 cm. long, the
bracts linear, linear-lanceolate, lanceolate, or lanceolate-oblong (rarely spatulate),
usually shorter than the corolla tube; calyx ca. 3 mm. long; corolla usually yellow
to orange, fading to red, sometimes red from the first, the tube 7-10 mm. long, the
limb 2-6 mm. broad (sometimes larger in cultivated specimens) ; drupes few, blue
to black at maturity, lustrous, juicy.
This is a common, weedy shrub in much of tropical America. It
is often grown in gardens of temperate America where it is a popular
bedding plant. The heads of ripe black fruits often resemble black-
berries. Although of poor flavor, they are eaten by children and are
sought by birds.
The material here referred to L. camara is divided by Moldenke
and others into several species but the characters on which the sup-
posed "species" are based are variable and inconstant.
Called "wild sage" in British Honduras.
Lantana hispida HBK. Nova Gen. & Sp. 2: 260. 1817. L. hirta
R. Grah. Edinb. Phil. Journ. 186. 1826. L. velutina Mart. & Gal.
Bull. Acad. Brux. 11, pt. 2: 325. 1844. L. involucrata var. velutina
GIBSON: FLORA OF GUATEMALA 205
Standl., Field Mus. Bot. 11: 172. 1936. L.frutilla Moldenke, Phyto-
logia 1 : 419. 1940. L. velutina f. violacea Moldenke, I.e. 8: 161. 1962.
L. hispida f. alba Moldenke, I.e. 9: 99. 1963. L. frutilla var. obtusi-
folia Moldenke, I.e. 14: 217. 1967. Chiligua (Jutiapa) ; chinkuro (Ja-
lapa) ; corronchocho (Chimaltenango, Guatemala) ; morita negra and
oregano del monte (Guatemala).
Damp forest or thickets, sometimes in oak, pine-oak, or pine for-
est, occasionally on dry, rocky hillsides, 300-2,700 meters; Alta
Verapaz; Baja Verapaz; Chimaltenango; Chiquimula; Escuintla;
Guatemala; Huehuetenango; Izabal; Jalapa; Jutiapa; Quezaltenan-
go; El Quich^ ; Sacatepe"quez ; San Marcos; Solola; Zacapa. Mexico
to Panama.
Erect, unarmed shrubs, 1-2 m. tall, the branches puberulent to hispidulous;
leaves usually opposite, rarely verticillate, usually short-petiolate, rarely subsessile,
the blades rugose, broadly ovate to lance-ovate, oblong-lanceolate, or ovate-
oblong, 1-9 cm. long (mostly 2-5 cm.), subacute to acuminate, acute or cuneate
at the base or broad and abruptly contracted, decurrent, usually scabrous above,
rarely merely puberulent or pilosulous, glandular-pellucid beneath and thinly his-
pidulous along the nerves to densely tomentulose or velutinous-pilosulous; margins
crenate (usually 18-30 indentations on each side) ; inflorescence headlike, peduncu-
late; bracts lance-ovate to broadly ovate, acute, usually as long as the corolla tube
or longer, the lowermost ones broadest, sometimes foliaceous, often venose; corolla
white, cream-colored, pale pink, or purplish, often with a yellow throat, the tube
4-6 mm. long; fruits purple black, juicy.
The two most diverse forms of this complex species are those
plants with oblong-lanceolate, finely crenate, scabrous to hispidulous
leaves (1,300-2,700 meters) and those with broadly ovate, coarsely
crenate, always velutinous-tomentulose leaves (600-1,900 meters),
and the temptation to retain L. velutina as a repository for all plants
in the latter category is understandable. However, one would then
be confronted with the problem of where to place the numerous inter-
mediate forms with various combinations of leaf shape, margin, and
indument that occur throughout the range (300-2,700 meters). A
photograph of the type specimen of L. hispida (Field Museum Neg.
No. 39493) clearly shows the lower leaf surface to have been fairly
tomentose rather than merely hispidulous, and, in fact, is almost
identical with some specimens which have previously been identified
as L. velutina.
The indument of stems and lower leaf surfaces of a single Guate-
malan collection which was reported as L. microcephala A. Rich.,
Watson 4.58c, is not strigose-canescent but is composed of ascending
hairs. Only in the shape of the leaves is it like L. microcephala
(which may prove to be synonymous with L. canescens HBK.).
206 FIELDIANA: BOTANY, VOLUME 24
Lantana involucrata L. Cent. PI. 2: 22. 1756; Amoen. Acad.
4: 319. 1759. L. odorata L. Syst. ed. XII, 2: 418. 1767. L. odorato
var. berlandieri Torrey, U. S. & Mex. Bound. Survey (Bot.) 128.
1858. L. involucrata var. odorata (L.) Moldenke, Phytologia 2: 53.
1941.
Open places, margins of thickets, chiefly along sea beaches; fre-
quently invading coconut plantations; southern Florida; Mexico;
British Honduras; West Indies; northern South America.
Erect, unarmed shrubs to 1.5 m. tall, the branches strigose; leaves on petioles
2-10 mm. long, the blades ovate, oblong-ovate or oblong-elliptic to obpyriforme,
2.5-7 cm. long in ours, the apex obtuse or rounded, acute at base or contracted and
decurrent, the margins very finely and closely crenulate, rugose and scabrous
above, usually strigillose but sometimes glabrate beneath; inflorescences globose,
usually densely flowered, long-pedunculate, the bracts oblong to elliptic, strigillose,
the lowermost ones 8-11 mm. long; corolla usually pink, lilac, or purple, sometimes
white, the tube puberulous, 2-4 mm. long; drupes 2-3 mm. long, blue or purple.
These plants are very aromatic.
Lantana trifolia L. Sp. PI. 626. 1753. L. trifolia var. geminata
Loes. Verh. Bot. Ver. Brandenb. 53: 75. 1911. Comida de paloma
(Izabal).
Usually in damp thickets, sometimes in low pine forest, rarely in
cleared land, near sea level to 1,200 meters; Alta Verapaz; Guate-
mala; Izabal (type from Los Amates, Seler & Seler 3378} ; Pete"n; El
Quiche". Mexico; British Honduras to El Salvador and Panama.
West Indies; South America.
Erect, unarmed shrubs, to 3 m. tall, the stems rather harshly pilose; leaves
usually ternate, sometimes binate, short petiolate, the blades lanceolate, oblong
lanceolate, elliptic-lanceolate, or occasionally ovate, 4-14 cm. long, acute or acu-
minate, acute or attenuate to the base, crenate serrate, scabrous or strigose above,
puberulent or pilosulous beneath, resinous-punctate; inflorescences at first head-
like, in age elongated and spikelike, sometimes 5 cm. long, densely flowered, pe-
dunculate, the peduncles 1 or 2 in each leaf axil, shorter than, equalling, or longer
than the leaves; bracts green, lanceolate or ovate, cuspidate; corolla usually pink,
lilac, or purple, rarely white, the tube puberulent, 4-6 mm. long; drupes lilac or
purple, juicy.
A common, weedy plant in many parts of the Central American
lowlands, often seen in second growth.
LIPPIA Houst. ex L.
Herbs, shrubs, or small trees, often aromatic, glabrous or pubescent; leaves
opposite or ternate, entire or dentate; inflorescences usually pedunculate, the pe-
GIBSON: FLORA OF GUATEMALA 207
duncles solitary to several in the leaf axils, spicate, the spikes short and nearly
globose or elongated and cylindrical, dense and many-flowered, the bracts generally
conspicuous; calyx small, membranaceous, ovoid-campanulate or compressed and
bicarinate; corolla tube cylindrical, straight or incurved, the limb oblique, spread-
ing, somewhat bilabiate, the lobes 4; stamens 4, didynamous, inserted near the
middle of the corolla tube, included or only slightly exserted; anthers ovate, not
appendaged in ours, the cells parallel; ovary bilocular, 1 ovule in each locule; style
terminal, stigma obscurely bilobate, oblique or recurved; ovules basal and erect or
affixed laterally near the base; fruit small, dry, included in the persistent calyx, at
maturity separating into 2 pyrenes; pericarp hard and dry; seeds without endo-
sperm.
A few herbaceous species of this genus have been treated by Mol-
denke and some earlier workers as constituting a distinct genus,
Phyla, but I prefer to leave them, as most authorities have, in Lippia.
Nearly 200 species have been reported, the majority of them from
tropical America. Additional studies may result in the reduction of
a considerable number of them to synonymy. Only 13 are known
from Guatemala.
Plants herbaceous to suffruticose, often prostrate or procumbent; flower heads
elongating in fruit; bracts cuneate-obovate to rhombic.
Peduncles much shorter than the flower heads, usually 2 or more in each leaf
axil L. betulaefolia.
Peduncles much longer than the flower heads, solitary in each leaf axil.
Leaf blades ovate or rhombic-ovate.
Hairs of leaf surface minutely bulbous-based, never malpighiaceous; calyx
completely villous L. dulcis.
Hairs of leaf surface malpighiaceous; calyx with a fine line of hairs on each
keel L. reptans.
Leaf blades linear-oblong, cuneate- oblong, spatulate, or cuneate-obovate.
Leaf blades linear-oblong; coarsely dentate along the whole margin; 10-
18 pairs of lateral veins, these prominent L. stoechadifolia.
Leaf blades spatulate to cuneate-obovate, usually dentate only above the
middle; the 5-6 pairs of lateral veins more or less obsolete.
L. nodiflora.
Plants erect shrubs or small trees; flower heads usually not or scarcely elongating
in fruit (except L. alba) ; bracts ovate or lanceolate.
Peduncles normally solitary in each leaf axil L. alba.
Peduncles normally 2 or more in each leaf axil.
Bracts of the inflorescence conspicuously 4-ranked; leaf blades uniformly
small, mostly 2-4 cm. long L. graveolens.
Bracts of the inflorescence not evidently 4-ranked; leaf blades variable in size,
at least some of them more than 4 cm. long.
Leaves densely and softly tomentose or pilose beneath.
Flower heads at anthesis 5-9 mm. in diameter.
Corolla yellow, twice as long as the calyx L. chiapasensis.
Corolla white, scarcely exceeding the calyx L. salamensis.
Flower heads at anthesis 10-16 mm. in diameter L. substrigosa.
Leaves glabrate beneath, strigose, thinly hispidulous, or scabrous.
208 FIELDIANA: BOTANY, VOLUME 24
Leaves smooth to the touch, the margins entire or obscurely crenate or
serrulate L. myriocephala.
Leaves rough to the touch, the margins conspicuously crenate, dentate,
or serrate.
Flower heads small (4-8 mm. in diameter at anthesis, in fruit 7-10 mm.
in diameter) ; lowermost bracts 3-8 mm. long L. cardiostegia.
Flower heads large (8-15 mm. in diameter at anthesis, in fruit to 25
mm.); lowermost bracts 10-15 mm. long L. controversa.
Lippia alba (Mill.) N. E. Browne ex Britton & Wilson. Fl.
Porto Rico 6: 141. 1925. Lantana alba Mill. Gard. Diet. ed. 8, no. 8.
1768. Lippia geminata HBK. Nova Gen. & Sp. 2: 266. 1818. Oro-
zuz; salvia santa; salvia siga (Alta Verapaz and Guatemala).
Usually on brushy hillsides and along roadsides (where it may
have escaped from cultivation), sometimes on riverbanks and sand-
bars, sea level to 1,800 meters; Alta Verapaz; Chiquimula; Escuintla;
Guatemala; Huehuetenango; Sacatepe"quez ; Solola. Texas; Mexico;
British Honduras to Panama; West Indies; South America.
Shrubs to 2 m. tall, usually sparsely branched, rather densely puberulent to
strigose; leaves opposite or sometimes ternate, the petioles 2-10 (14) mm. long,
the blades 2-7 cm. long, oblong, lance-oblong, or ovate-oblong, acute or obtuse,
cuneate or attenuate to the base and decurrent on the petiole, strigose-hirtellous
or puberulent, sometimes canescent, the margins finely serrate; peduncles usually
solitary (rarely geminate) in the leaf axils; flower spikes first subglobose, ca. 6 mm.
long, usually elongating to 8-12 mm. long; bracts puberulous, ovate, abruptly
acuminate, the lower ones sometimes mucronate, 3-5 mm. long; calyx villous,
1.5-2 mm. long; corolla pale lilac, purple, or white with purple, 5-6 mm. long.
Grown frequently in gardens throughout Central America as a
medicinal plant, a tea made from the leaves being a favorite domestic
remedy for both intestinal and respiratory disturbances.
Lippia betulaefolia HBK. Nova Gen. & Sp. 2: 264. 1818.
Phyla betulaefolia Greene, Pittonia 4: 48. 1899.
Open, wet fields or swamps, sea level to 225 meters; Escuintla:
Izabal. British Honduras to Panama; West Indies; South America.
Plants annual or perennial, erect or decumbent, often rooting at the nodes, the
stems strigillose; leaves on petioles 0.5-2.5 cm. long, the blades ovate or rhombic-
ovate, 1.5-6.5 cm. long, acute or obtuse, cuneate and decurrent at the base,
coarsely dentate, densely or sparsely strigose with largely malpighiaceous hairs;
peduncles short, 1-4 mm. long, 2-6 or more at each node; flower spikes oblong,
0.5-2 cm. long; bracts of the inflorescence cuneate, cuspidate; calyx minute; co-
rollas greenish or pinkish, to 1 mm. long.
Lippia cardiostegia Benth. Voy. Sulphur 153. 1844. L. lucens
Standl. Trop Woods 37: 30. 1934. L. brenesii Standl. Field Mus.
GIBSON: FLORA OF GUATEMALA 209
Bot. 18: 1009. 1938. L. hypoleia var. ovatifolia Moldenke, Phytologia
14: 217. 1967. Varafina (Guatemala) ; chiliqua (Jutiapa) ; coronchoch
(Huehuetenango) .
Damp or dry, often rocky, brushy plains and hillsides, sometimes
in open, mixed forest, 150-1,500 meters; Alta Verapaz; Chimaltenan-
go; Chiquimula; Escuintla; Guatemala; Huehuetenango; Jutiapa;
El Quich^ ; Sacatepe"quez ; Santa Rosa. Southern Mexico; El Sal-
vador; Honduras; Nicaragua; Costa Rica.
Shrubs to 3 m. tall, the branches strigose or scabrous; leaves petiolate, the
blades lanceolate, oblong-ovate to ovate or elliptic-oblong, usually 2-7 cm. long,
1-3 cm. wide (rarely to 12 cm. long, 4-5 cm. wide), 5-7 pairs of lateral veins, acute
or subobtuse at apex, cuneate-attenuate to the base, the margins crenate to serrate,
upper surface bullate-rugose, usually hirtellous-scabrous, harsh to the touch, often
lustrous, usually glabrate beneath but somewhat hirtellous-scabrous on the veins;
peduncles 2-several in leaf axils, equalling or shorter than the leaves, usually 1.5-
3 cm. long (rarely 4-5 cm. long); flowering spikes nearly globose, variable in size,
most often 8-12 mm. long; bracts ovate to rounded-ovate, sometimes cordate,
acute, acuminate, or apiculate, puberulent, scabrous, or nearly glabrous, minutely
ciliate, the outer ones 3-8 mm. long; calyx 1-2 mm. long, usually densely hirtellous;
corolla 2-3 mm. long, pale greenish yellow or greenish white, puberulent, often
glandular below lobes.
This species is extremely variable, especially in leaf size and length
of peduncles, but there seem to be no definitive characters by which
the various forms can be separated. One specimen, Steyermark 31596,
is atypical because of its relatively narrow, oblanceolate to elliptic-
oblong leaves.
Several collections from Honduras of L. oxyphyllaria (Donn.-
Sm.) Standley (once thought to be endemic to Costa Rica and Pan-
ama) have been misidentified and distributed in herbaria as L. lucens.
Those which I have seen, Williams and Molina R. 10802, Molina
R. 1456, 1837 and 3213, and Standley 26473 were also incorrectly
cited by Moldenke. Although the leaves of L. oxyphyllaria are bul-
late-rugose and sometimes glabrate and lustrous, they are usually
more or less pilose; the flower heads are much larger than those of
L. cardiostegia, often 2.5 cm. wide, and in fruit 3-4 cm. long, with
large bracts, sometimes as long as 1.8 cm.
Lippia chiapasensis Loes. Verh. Bot. Ver. Brandenb. 53: 78.
1911. L. curtisiana Moldenke, Phytologia 1: 425. 1940.
Wet or dry, often rocky thickets or forests, frequently in pine-oak
forest, sometimes in meadows, 1,500-3,000 meters; Baja Verapaz;
Huehuetenango; San Marcos; Solola; Totonicapan. Mexico.
210 FIELDIANA: BOTANY, VOLUME 24
Shrubs or weak trees to 4 m. tall, the branches densely pubescent, leaves on
petioles 4-14 mm. long, the blades ovate or ovate-elliptic, 2-6(9) cm. long, 1.5-
4.5 cm. wide, usually acute, sometimes obtuse, usually cuneate (rarely rounded)
at the base, bullate and scabrous above, usually densely and softly pubescent be-
neath, venation prominent, margins crenate to serrate; peduncles 2-4 in each axil,
densely pubescent, 1.5-2.5 cm. long, the flower spikes 8-9 mm. wide, in fruit to
13 mm. broad, dense and many-flowered; bracts ovate at an thesis, 4-5 mm. long,
3-6 mm. wide, reniform in age, 6-9 mm. wide, venose, puberulent to strigillose,
ciliate; calyx 2-3 mm. long, hirsute; corolla yellow, twice as long as the calyx,
usually puberulous at apex and within throat.
Although this plant has smaller flowering heads and generally
smaller leaves than L. substrigosa Turcz., they appear to be closely
related.
Lippia controversa Moldenke, Phytologia 1 : 423. 1940. L. pine-
torum Moldenke, Phytologia 2: 20. 1941. L. controversa var. brevi-
pedunculata Moldenke, Phytologia 4: 56. 1952.
Damp or dry, brushy fields or hillsides, 500-1,400 meters; Es-
cuintla; Jutiapa; Sacatepe"quez ; Santa Rosa. Mexico; Honduras.
Shrubs to 2 m. tall, the branches hispidulous or glabrate; leaves short-petio-
late, the blades broadly ovate or ovate-elliptic, mostly 4-9 cm. long, acute, often
abruptly so, usually rounded or obtuse at the base and abruptly long-decurrent,
very rough to the touch, hispidulous above with bulbous- based hairs, sparsely his-
pidulous beneath, especially on the veins, rugose, the margins crenate or dentate;
peduncles 1-2 in each leaf axil, 2-4 cm. long, slender, glandular-pubescent; bracts
usually ovate, sometimes nearly lanceolate, acute, the lowest 1-1.5 cm. long, ve-
nose, strigillose to puberulent; the spikes in fruit sometimes 2.5 cm. broad; calyx
ca. 2 mm. long at anthesis, to 3 mm. in fruit, villous; corolla 4-6 mm. long, puberu-
lent, cream-colored, yellow, or greenish-yellow.
Sometimes confused with L. oxyphyllaria (Donn.-Sm.) Standley,
with which it is probably closely related.
Lippia dulcis Trev. Nov. Act. Acad. Caes. 13, pt. 1: 187. 1826.
Zapania scaberrima Juss. ex Pers. Syn. PI. 2: 140. 1806, not Lippia
scaberrima Sond. 1850. Phyla dulcis Moldenke, Torreya 34: 9. 1934.
Phyla scaberrima Moldenke in Fedde, Rep. Sp. Nov. 41: 64. 1936.
Orozuz (Alta Verapaz).
Damp thickets and waste ground, wooded river banks, borders of
ponds, or in open clearings and pastures, sea level to 1,800 meters;
Alta Verapaz; Chiquimula; Guatemala; Pete"n; Retalhuleu; Sacate-
pe"quez; Santa Rosa; Solola. Southern Mexico; British Honduras to
Panama; West Indies.
GIBSON: FLORA OF GUATEMALA 211
Perennial plants, erect or decumbent, rarely to 40 cm. tall, usually more or less
fruticose near the base, strong-scented, the stems often rooting at the lower nodes,
strigillose or glabrate; leaves on petioles 0.5-1.5 cm. long, the blades rhombic to
ovate, 1-6 cm. long, acute to somewhat acuminate, broadly cuneate at the base,
the margins crenate-serrate, upper surface strigose, rough to the touch, sparsely
to densely strigillose beneath and obscurely glandular ; peduncles solitary in the leaf
axils, 1-5 cm. long; flower spikes ovoid- globose at first, later cylindric, ca. 6 mm.
thick, the heads occasionally elongating to as much as 3 cm. long in age but usually
shorter; bracts cuneate-ob ovate, obtuse and abruptly acuminate; calyx minute,
villous; corolla white, 1-1.5 mm. long.
Lippia dulcis is one of the group of Lippias which some workers
hold to belong to the separate genus Phyla, but does not have mal-
pighiaceous hairs.
This plant is said to be much used in domestic medicine. The
leaves are aromatic; the root when chewed has the flavor of licorice,
hence the name "Orozuz," common in Central America.
Lippia graveolens HBK. Nova Gen. & Sp. 2: 266. 1818. Lan-
tana origanoides Mart. & Gal. Bull. Acad. Brux. 11(2): 327. 1844.
Lippia berlandieri Schauer in DC. Prodr. 11: 575. 1847. Gonio-
stachyum graveolens Small, Fl. Southeast. U. S. 1012. 1903. Oregano.
Rocky slopes or damp thickets on plains, 350 meters or less;
Pete"n; Zacapa. Southern Texas; Mexico; Nicaragua.
Slender shrubs to 2 m. tall, the branches short-pilose; leaves on petioles usually
5-10 mm. long, the blades oblong to elliptic or ovate to ovate-oblong, 2-4 cm. long,
usually obtuse or rounded at the apex, sometimes acute, rounded or subcordate at
the base, densely soft-pilosulous above, soft to the touch, glandular and densely
tomentose or pilosulous beneath, the margins finely crenate; peduncles 2-6 in leaf
axils, 4-12 mm. long; flower spikes subglobose to oblong, 4-12 mm. long; bracts
4-ranked, ovate to lanceolate, acute, glandular and densely pilosulous; calyx 1-
2 mm. long, glandular and villous; corolla white, the tube strigollose, 3-6 mm. long.
The aromatic leaves, either fresh or dried, are used in Central
America for flavoring food, and the dry leaves are often sold in the
markets.
Lippia myriocephala Schlecht. & Cham. Linnaea 5: 98. 1830.
L. hypoleia Briq. Ann. Cons. Jard. Bot. Geneve 4: 236. 1900. L. my-
riocephala var. integrifolia Loes. Verh. Bot. Ver. Brandenb. 53: 77.
1911 (type from Coban, Alta Verapaz, Seler 2^85). L. hypoleia var.
ovatifolia Moldenke, Phytologia 14: 217. 1967. Cutujume (Izabal);
sacatzum (Alta Verapaz) .
Wet or dry, often rocky thickets, open hillsides, outer slopes of
cloud forest, often in pine-oak forest, 300-2,700 meters; Alta Vera-
212 FIELDIANA: BOTANY, VOLUME 24
paz; Baja Verapaz; Chimaltenango; Chiquimula; Guatemala; Iza-
bal; Pete"n; El Progreso; Quezaltenango; Sacatepe"quez; San Marcos;
Santa Rosa. Mexico; British Honduras; Honduras to Costa Rica.
Shrubs or trees to 12 m. tall, younger branches pilose with appressed or spread-
ing hairs; leaves on petioles 3-14 mm. long, the blades lance- oblong to narrowly
elliptic- oblong, oblong-ovate or (rarely) ovate, mostly 5-15 cm. long, 1-5 cm. wide,
usually acuminate to long-acuminate, cuneate-attenuate to the base, thinly ap-
pressed-pilosulous above or glabrate, not rough to the touch, sparsely pilosulous
beneath with spreading or appressed hairs to almost glabrous, the margins ob-
scurely crenulate or serrulate or entire; peduncles 3-10 in each axil, 1-4 cm. long,
the densely flowered spikes small, only 5-6(7) mm. long and broad, in fruit 8-
9 mm. long; bracts broadly ovate to subreniform, abruptly acute or short acumi-
nate, puberulent and ciliate, the lower ones 2-3 mm. long; calyx villous, ca. 1 mm.
long; corolla white or cream, sometimes tinged with lavender, the tube 0.5-1 mm.
long.
The wood is said to be used for rafters and poles in the poorer
dwellings in warmer regions.
Lippia nodiflora (L.) Michx. Fl. Bor. Amer. 2: 15. 1803. Ver-
bena nodiflora L. Sp. PL 20. 1753. Phyla nodiflora Greene, Pittonia
4: 46. 1899. P. incisa Small, Fl. Southeast. U. S. 1012. 1903. P.
nodiflora var. longifolia Moldenke, Phytologia 2: 22. 1941.
In damp soil, usually in open places, often on sandbars, salt flats,
beaches, wet meadows, and irrigation ditches, sea level to 1,400 me-
ters; Chiquimula; Esquintla; Guatemala; Huehuetenango; Izabal;
Jutiapa; Pete"n; Retalhuleu; Zacapa. United States and Mexico;
British Honduras to El Salvador and Panama; West Indies; South
America; Old World.
Plants herbaceous, creeping and often forming large mats or colonies, the stems
strigillose or glabrate; leaves short-petiolate, often succulent, the blades spatulate,
oblanceolate, or cuneate-obovate, mostly 1-4 cm. long, rounded or obtuse at the
apex, cuneate at the base, dentate above the middle, strigillose with malpighiaceous
hairs or almost glabrous, peduncles solitary in the axils, usually longer than the
leaves; flower spikes at first globose, elongating and becoming cylindric in age,
1-2 cm. long, 6-9 mm. thick in fruit; bracts purplish, closely imbricate, cuneate-
obovate to rhombic, abruptly acuminate, the lower ones often cuspidate; calyx
bicarinate, with a line of very fine hairs along each keel; corolla white or purplish,
3-4 mm. long.
Lippia reptans (Spreng.) HBK. Nova Gen. & Sp. 2: 263. 1818.
Zapania reptans Spreng. PI. Pugill. 2: 70. 1813. Lippia repens
Spreng. Syst. 2: 752. 1825. L. strigulosa Mart. & Gal. Bull. Acad.
Brux. 11(2) : 319. 1844. L. nodiflora var. reptans 0. Ktze. Rev. Gen.
2: 508. 1891. Phyla reptans Greene, Pittonia 4: 47. 1899. P. nodi-
GIBSON: FLORA OF GUATEMALA 213
flora var. reptans Moldenke, Torreya 34: 9. 1934. P. yucatana Mol-
denke, Phytologia 2: 140. 1946. P. yucatana var. parviflora Moldenke,
Phytologia 2: 141. 1946. P. strigulosa Moldenke, I.e. 233. 1947.
P. strigulosa var. parviflora Moldenke, I.e.
Usually in wet, open places, often bordering lakes or streams, near
sea level to 1,500 meters; Alta Verapaz; Escuintla; Huehuetenango;
Izabal; Jutiapa; Pete"n; Sacatepe"quez. Mexico; British Honduras;
Honduras; El Salvador; Nicaragua; Costa Rica; northern South
America.
Plants herbaceous, usually prostrate and rooting at the nodes, strigillose
throughout with largely malpighiaceous hairs; leaves on petioles 2-12 mm. long,
the blades rhombic to broadly ovate, 1-5 cm. long, obtuse or acute, broadly cuneate
at the base, coarsely and sharply dentate, usually 6-9 pairs of lateral veins, these
conspicuous beneath; peduncles solitary in the leaf axils, longer than the leaves,
sometimes to 10 cm. long, the flower spikes subglobose to oblong, usually 1 cm.
long or less, rarely to 2 cm.; bracts often purplish, cuneate- obovate to rhombic,
abruptly acuminate, the lower ones sometimes cuspidate; calyx bicarinate with a
line of very fine hairs along each keel, 2-2.5 mm. long, corolla white or tinged with
purple, or with a purplish throat, ca. 3 mm. long.
Lippia salamensis Loes. Verh. Bot. Ver. Brandenb. 53:77. 1911.
Damp or dry, open hillsides or in oak forest, about 1,600 meters;
Baja Verapaz (type from Cuesta Choacuz, Seler 24.83 and 3404.) ; col-
lected also in the mountains near Morazan; Jalapa (Chahuite, north-
west of Jalapa) .
Slender shrubs about 2 m. tall, the branches densely pilose with short appressed
hairs; leaves thick, short-petiolate, the blades ovate to ovate-oblong or oblong,
(2)4-10 cm. long, rounded or acute at the apex, acute and decurrent to the base,
the margins crenate-serrulate, upper surface somewhat rugose and short-strigose
or sericeous-strigose, densely pubescent to velutinous-tomentose beneath with
short, more or less spreading, soft hairs; peduncles several in each axil, usually
about 5, commonly 1-2 cm. long, slender, appressed-pilosulous; flower spikes sub-
globose to ovoid, in anthesis 5-7 mm. long and 7 mm. broad, in fruit elongating
to 10 mm. long; bracts thin, reticulate-veined, pubescent or hirtellous, ciliolate,
broadly ovate to cordate-reniform; calyx 2-2.4 mm. long, hirtellous; corolla white,
scarcely longer than the calyx, puberulent.
This closely resembles a fragment of the type of L. jurgensenii
Briq. from Oaxaca, Mexico (Ann. Conserv. Jard. Bot. Geneve 4: 239.
1900). The description is very similar except for the shorter calyx
of the Mexican plant. If they should prove to be synonymous, as
seems likely, then L. jurgensenii would, of course, take precedence.
Because of inadequate material, it is probably best to maintain them
separately for the present.
214 FIELDIANA: BOTANY, VOLUME 24
Lippia stoechadifolia (L.) HBK. Nova Gen. & Sp. 2: 265. 1818.
Verbena stoechadifolia L. Sp. PI. 19. 1753. Phyla stoechadifolia (L.)
Small, Bull. Torrey Club 36: 162. 1909.
Wet thickets, damp, open fields, swampy land bordering lakes
and streams, rarely on open, rocky banks, sea level to 1,500 meters;
Alta Verapaz; Huehuetenango; Jutiapa; Pete"n. Southern Florida;
Mexico; British Honduras; Honduras; West Indies; northern South
America.
Erect shrubs to 1.5 m. tall, or suffruticose and spreading, often much branched,
the stems strigose, leaves nearly sessile to short-petiolate, the blades thick, oblong-
linear, 2-10 cm. long, 0.3-1 cm. (rarely 1.5 cm.) wide, acute, cuneate at the base,
the margins serrate-dentate, both surfaces strigose with both simple and malpighia-
ceous hairs, the veins very conspicuous beneath, the lateral ones numerous, usually
10-14 pairs; peduncles solitary in the leaf axils, usually longer than the leaves;
flower spikes at first globose, in age oblong, 2-3 cm. long, 0.5-0.7 cm. in diameter;
bracts cuneate- obovate; calyx bicarinate, with a line of fine hairs along each keel,
2-2.5 mm. long; corolla white or purplish, 2.5-3.5 mm. long.
Lippia substrigosa Turcz. Bull. Soc. Nat. Moscou, pt. 2: 202.
1863. L. kellermanii Greenm. Field Mus. Bot. 2: 341. 1912 (type
from Lago de Amatitlan, Guatemala, W. A. Kellerman 6372}. L. gua-
temalensis Gandoger, Bull. Soc. Bot. France 65: 63. 1918 (type from
Guatemala, Tuerckheim s.n.). Chichicaste de venado (Guatemala);
salvia santa (Quezaltenango) ; supup (Alta Verapaz) .
Wet or dry thickets, oak-pine forest, sometimes on open, rocky
slopes, 1,200-2,800 meters; Alta Verapaz; Baja Verapaz; Chimalte-
nango; Chiquimula; Guatemala; Huehuetenango; Jalapa; Quezal-
tenango; El Quiche* ; Sacatepe"quez ; San Marcos; Santa Rosa; Solola;
Zacapa. Southern Mexico; British Honduras; Honduras; El Sal-
vador; Nicaragua.
Shrubs or trees to 7 m. tall, the branches usually densely viscid-pubescent with
harsh, brownish, spreading hairs, sometimes pilose with pale, spreading hairs;
leaves on petioles 0.7-3 cm. long, the blades ovate to broadly ovate, ovate-elliptic
or lance-ovate, mostly 5-24 cm. long, 2.5-12 cm. wide, acute to acuminate, cordate
to abruptly contracted and obtuse at the base or cuneately long attenuate, the sur-
face rugose or finely bullate, scabrous-hispid, usually very rough to the touch,
densely hirsute-tomentose or pilose beneath or both surfaces rarely glabrate, the
margins crenate-serrate; peduncles usually 4-8 in each leaf axil, 2.5-6 cm. long,
hirsute and usually glandular-pubescent; flower spikes subglobose, at an thesis 10-
20 mm. long, 10-16 mm. wide, in fruit to 28 mm. long and 25 mm. wide; bracts
broadly ovate to subreniform, acute or rounded and abruptly acuminate to cuspi-
date at the apex, hispidulous or glandular-pubescent and ciliate, the lower ones
sometimes to 14 mm. long; calyx 2-2.5 mm. long, villous; corolla pale yellow, 4-6
mm. long.
FIG. 40. Lippia substrigosa. A, flowering branch, J/a natural size; B, flower,
X 5; C, corolla dissected to show pistil and stamens, X 5.
215
216 FIELDIANA: BOTANY, VOLUME 24
This plant has been reported from Guatemala as L. umbellata
Cav. and some of the material has been determined by Moldenke
and cited by him as L. callicarpaefolia HBK. Lippia umbellata,
which has been poorly understood and which probably includes L.
pringlei Briquet, was originally described (Cav. Icon. PL 2: 75, 1. 19^.
1793) as having "flores umbellati, umbellis pluribus axillaribus in
ramorum summitatibus" and the illustration clearly shows primary
peduncles with several "rays" forming definite dichasia. I have
seen no material of L. umbellata nor of the closely related L. pringlei
from either Guatemala or Chiapas. The Mexican L. callicarpaefolia,
which in foliage and large flowering heads does resemble L. substri-
gosa, differs markedly in its dichasial inflorescence and conspicu-
ously colored (rose to purple) bracts.
PETREA L.
Reference: Moldenke, H. N., A monograph of the genus Petrea,
in Fedde, Rep. Sp. Nov. 43: 1-48; 48: 161-221. 1938.
Shrubs, trees, or woody vines; leaves opposite, sub-opposite, or verticillate,
the venation prominent, both surfaces often very rough; inflorescences axillary or
terminal, racemose, usually elongated and many-flowered; bracts small, caducous;
flowers long-pedicellate and often remote on the rachis; calyx tube cylindric or
campanulate, costate, 5-lobate, the lobes subequal, elongated, cristate ventrally
at the base; corolla salverform, usually blue or violet, rarely white, slightly zygo-
morphic, the tube urceolate, cylindric, or funnelform, usually pubescent within near
the middle, the limb spreading, 5-lobate; stamens 4, didynamous, the filaments
very short, inserted near the middle of the corolla tube, included, or the anthers
scarcely exserted; anthers oblong or ovate, dorsifixed near the base, the cells open-
ing by an introrse slit; style terminal, the stigma capitate, oblique; overy bilocular,
one ovule in each locule; ovules lateral, ascending, sometimes pendent; fruiting
calyx accrescent, the lobes subcoriaceous, reticulate- veined ; fruit drupaceous, com-
pletely enclosed in the accrescent calyx, the exocarp coriaceous, the endocarp hard,
of 2 pyrenes.
Perhaps 30 species, in tropical America. One other species occurs
in southern Central America.
Petrea volubilis L. Sp. PL 626. 1753. P. arborea HBK. f. albi-
flora Standl. Field Mus. Bot. 11: 140. 1932 (type from Sittee River,
Stann Creek District, British Honduras, Schipp 727). P. volubilis
var. albiflora (Standl.) Moldenke, Revista Sudamer. Bot. 5: 2. 1937.
P. volubilis var. pubescens Moldenke in Fedde, Rep. Sp. Nov. 43:
45. 1938. P. volubilis f. albiflora Standl. Field Mus. Bot. 18: 1011.
1938. Colacidn (Guatemala); corona de la reina and Santa Rita
(Pete"n); cuerode sapo (El Progreso); yochopptzimin (Yucatan, Maya).
FIG. 41. Petrea volubilis. A, flowering branch, y% natural size; B, accrescent
calyx and style, X 2; C, fruiting calyx enclosing fruit, X 1; D, corolla dissected
to show stamens, X 3.
217
218 FIELDIANA: BOTANY, VOLUME 24
Dry thickets or forest, rocky slopes, near sea level to 1,800 me-
ters; often cultivated; Alta Verapaz; Baja Verapaz; Chimaltenango;
Escuintla; Guatemala; Izabal; Jutiapa; Pete"n; El Progreso; Quezal-
tenango; El Quiche; Retalhuleu; San Marcos; Santa Rosa; Suchite-
pe"quez; Zacapa. Mexico; British Honduras to El Salvador and
Panama; West Indies.
Woody vines or subscandent shrubs, the stems puberulent or hirtellous, some-
times as much as 9 cm. in diameter; leaves short-petiolate, the blades coriaceous,
rough to the touch, oblong, obovate, or elliptic-oblong, mostly 6-15 cm. long, acute
or obtuse, acute to rounded at the base, entire, scabrous or pilosulous above, sca-
brous beneath or rather densely pilosulous; racemes usually pendent, solitary in the
leaf axils, often clustered near the ends of the branches, mostly 8-20 cm. long, lax;
bracts linear or linear-lanceolate, 4-8 mm. long, caducous; calyx tube ca. 3 mm.
long at anthesis, the lobes oblong, in fruit as much as 2 cm. long, blue or purple
(rarely white) ; corolla tube 6-8 mm. long, usually somewhat puberulent, at least
near the top, the lobes minutely puberulent outside, essentially glabrous within;
ovary and style glabrous.
This handsome vine is very common in the dry lower Motagua
Valley, where it is covered with flowers in late March, and is frequent
in many places along the Pacific plains and foothills. It has been
introduced into cultivation in many warm regions and in Florida is
known as "purple- wreath." This species has been reported from
Guatemala as P. arborea HBK., a species which is usually arbores-
cent, with sessile or subsessile, often clasping leaves, and apparently
confined to the West Indies and South America. The other Central
American species is P. aspera L., with pedicels and calyx essentially
glabrous.
PRIVA Adanson
References: C. E. Kobuski, A revision of the genus Priva, Ann.
Mo. Bot. Gard. 13: 1-35, t.1-5. 1926; H. N. Moldenke, A monograph
of the genus Priva, in Fedde, Rep. Sp. Nov. 41: 1-76. 1936.
Herbaceous annuals or perennials with usually rough pubescence; leaves oppo-
site, sessile or petiolate, the blades membranaceous, dentate; inflorescences race-
mose, terminal or axillary, the flowers small, pedicellate, bracteate, the small bracts
lanceolate or ovate; calyx tubular or cylindric in anthesis, 5-costate, the costae
terminating in short teeth, the calyx persistent, accrescent and enclosing the fruit,
usually more or less contracted at the orifice; corolla tube cylindric, the limb more
or less bilabiate, the 5 lobes spreading, oblique; stamens 4, didynamous, inserted
at about the middle of the corolla tube, included or scarcely exserted; anther cells
parallel or slightly divergent; ovary bilocular, the ovules 2 or by abortion 1; style
terminal, stigma bilobate; fruit dry, included in the enlarged calyx, separating at
maturity into 2 bilocular or by abortion unilocular cocci, the pericarp hard, the
dorsal surface of the cocci echinate, ridged and rugose, or smooth, the inner surface
excavated, concave, or plane.
GIBSON: FLORA OF GUATEMALA 219
About 20 species, in tropical America, Asia, and Africa, inclined
to be weedy. Only three occur in Guatemala; no others are known
in Central America.
Fruiting calyx broadly ovoid, not closely investing the fruit; fruits distinctly
echinate P. lappulacea.
Fruiting calyx globose or nearly so, closely investing the fruit; fruits reticulate-
rugose to reticulate- verrucose but not echinate.
Pedicels reflexed in fruit; calyx 1.5-2.5 mm. long P. mexicana.
Pedicels erect in fruit; calyx 3-4 mm. long P. aspera.
Priva aspera HBK. Nova Gen. & Sp. 2: 278. 1817. P. orizabae
S. Wats. Proc. Am. Acad. 23: 282. 1888. Chile hueco (Guatemala).
Damp thickets, brushy slopes, sometimes in pine-oak forest,
1,000-2,100 meters; Alta Verapaz; Chimaltenango; Escuintla; Gua-
temala; Huehuetenango; Jalapa; El Quiche"; Sacatepe"quez ; Solola.
Mexico; Honduras; Nicaragua; Costa Rica.
Erect or ascending annuals, the stems often hollow, acutely tetragonous, pu-
berulent; leaves on petioles 0.3-5 cm. long, the blades ovate or lanceolate-ovate,
mostly 4-15 cm. long, rarely to 20 cm. long, acuminate to long-acuminate, cuneate
or obtuse at the base, the margins crenate to crenate- dentate, scabrous above,
puberulent beneath; racemes terminal or axillary, 6-40 cm. long; flowers often re-
mote, short- pedicellate, the pedicels erect in fruit, the bracts lanceolate, usually
exceeding the pedicels; calyx at anthesis tubular, 3-4 mm. long, in fruit globose,
closely investing the fruit, 3-4 mm. long, obscurely rostrate, usually densely puber-
ulent and bearing some longer, uncinate hairs; corolla blue to purple, 5-7 mm. long;
cocci of the fruit 2, the dorsal surface reticulate-rugose, not echinate, the inner sur-
face usually shallowly excavated.
The ripe fruit is black and lustrous.
Priva lappulacea (L.) Pers. Syn. PI. 2: 139. 1807. Verbena lap-
pulacea L. Sp. PI. 28. 1753. P. echinata Juss. Ann. Mus. Paris 7:
70. 1806. Pega-pega (Zacapa).
Wet or dry thickets, clearings, waste ground, common near dwell-
ings, sea level to 900 meters; Alta Verapaz; Chiquimula; Escuintla;
Guatemala; Izabal; Pete"n; Retalhuleu; Santa Rosa; Solola; Zacapa.
Southern Florida; Mexico; British Honduras to El Salvador and
Panama; West Indies; South America; introduced in the Old World
tropics.
Erect, ascending, or decumbent annuals, often much branched, to 75 cm. tall,
the stems tetragonous, pubescent; leaves on petioles 0.5-4 cm. long, the blades
ovate or lanceolate-ovate, 2-10 cm. long, acute or acuminate, rounded or subcor-
date at the base, the margins crenate-dentate, hispidulous or glabrate above,
puberulent beneath; racemes lax, 5-20 cm. long, the flowers short-pedicellate, the
FIG. 42. Priva lappulacea. A, habit, % natural size; B, calyx opened to show
pistil, X 10; C, corolla dissected to show stamens and staminode, X 3; D, fruiting
calyx enclosing fruit, X 5; E, mature fruit, X 8.
220
GIBSON: FLORA OF GUATEMALA 221
bracts narrowly lanceolate, usually shorter than the pedicels; calyx at an thesis
tubular, 2-3 mm. long, densely uncinate-hispidulous, in fruit broadly ovoid, ap-
pearing inflated, constricted at the apex and often somewhat rostrate, 4-7 mm.
long; corolla usually blue or purple, sometimes pink or white, 4-5 mm. long; fruit
of 2 bilocular cocci, these echinate dorsally, the inner surface plane or nearly so.
Throughout most of the tierra caliente of Central America this
is a common weed, often abundant about dwellings. The fruiting
calyces adhere to clothing and even to the feathers of birds by the
abundant small uncinate hairs; the common names "pega-pega,"
and in El Salvador and Honduras "mozote" and "mozotillo," indi-
cate this characteristic.
Priva mexicana (L.) Pers. Syn. PL 2: 139. 1806. Verbena mexi-
cana L. Syst. 66. 1784. P. hispida Juss. Ann. Mus. Paris 7: 70. 1806.
Damp thickets and hedges, wooded slopes, sometimes in pine or
scrub-oak forest, or a weed in cultivated ground, 50-1,800 meters;
Baja Verapaz; Chimaltenango; Escuintla; Guatemala; Jalapa; Pe-
ten; Sacatepe"quez. Mexico; Honduras.
Erect or decumbent annuals, to 1 m. tall, often much branched, the stems
tetragonous, pubescent or glabrate; leaves on petioles mostly 0.5-2.5 cm. long, the
blades ovate to lanceolate-ovate, acute or acuminate, rounded at the base or the
lowest ones subcordate, crenate, hispidulous-strigose above, paler beneath and
viscid- puberulent; racemes terminal or axillary, pedunculate, 5-30 cm. long, the
bracts lanceolate, equalling or more often exceeding the pedicels, these 0.5-1 mm.
long, reflexed in age; calyx densely uncinate-hispidulous, cylindric at anthesis,
1.5-2.5 mm. long, in fruit 2-3 mm. long and closely investing the fuit; corolla usu-
ally blue to purple, sometimes white, about twice as long as the calyx; fruit com-
posed of 2 unilocular cocci, these reticulate-verrucose dorsally, not echinate,
somewhat concave or excavated on the inner surface, separating easily.
REHDERA Moldenke
Reference: H. N. Moldenke, A monograph of the genus Rehdera,
in Fedde, Rep. Sp. Nov. 39: 47-55. 1935.
Trees; leaves subcoriaceous, triplinerved or with 5-7 pairs of lateral veins;
inflorescences short-racemose, the flowers small, bracteate; calyx tubular, with 5
costae terminating in 5 short teeth, the fruiting calyx usually splitting into 5 seg-
ments and early deciduous; corolla salverform, the tube cylindric, about equalling
the calyx, the 5 lobes long and narrow, tomentose or pilose within; perfect stamens
4, included, inserted near the apex of the corolla tube; anthers oblong, introrse,
dorsifixed near the base; style simple, terminal, the stigma subcapitate, obscurely
bilobate; ovary bicarpellate, each carpel bilocular, the ovules attached basally;
fruit oblong, dorsally compressed, lustrous, composed of 2 cocci, often more or less
marginally winged, rounded or subretuse at the apex; seed linear.
222 FIELDIANA: BOTANY, VOLUME 24
Leaf blades usually obtuse or rounded at the apex (rarely acute or abruptly short-
acuminate); fruiting calyces sinuately denticulate R. trinervis.
Leaf blades acuminate to long-acuminate; fruiting calyces subtruncate.
R. penninervia,
Rehdera penninervia Standl. & Moldenke in Fedde, Rep. Sp.
Nov. 39: 50. 1935. Palo bianco (fide Standley); raspa sombrero
(Pet&i).
Upland limestone forest or thickets, 300 meters or less; Pet£n
(type from Uaxactun, H. H. Bartlett 12317}. British Honduras.
Trees to 25 m. tall, the trunk 45-75 cm. in diameter, the branchlets glabrate;
leaves on slender, often minutely puberulent petioles 5-15 mm. long, the blades
subcoriaceous, lustrous, 5-10 cm. long, with 5-7 pairs of lateral veins, lance- oblong
or elliptic-oblong, long-acuminate, cuneate at the base, the margins entire, glabrous
on both surfaces or sometimes barbellate beneath in the axils of the costae and low-
ermost veins; racemes axillary and terminal, 1-3 cm. long, sometimes branched,
with few or many flowers; pedicels 1 mm. long or less; bracts soon deciduous, linear
or narrowly triangular, 1-3 mm. long; calyx tubular at an thesis, subtruncate, the 5
costae terminating in 5 minute teeth, glabrous outside, sericeous within, ca. 5 mm.
long, in fruit cupuliform, truncate, soon deciduous; corolla white, glabrous outside,
the tube 4-5 mm. long, the lobes 3-4 mm. long, tomentose to pilose inside; fruit
oblanceolate-oblong, compressed, about 17 mm. long and 8 mm. broad.
The crushed leaves are reported to have a distinctive odor.
Called "hinge hinge" and "roble del mico" in British Honduras.
Rehdera trinervis (Blake) Moldenke in Fedde, Rep. Sp. Nov.
39: 52. 1935. Citharexylum trinerve Blake, Proc. Biol. Soc. Wash.
34: 45. 1921. C. macrocarpum Standl. Journ. Wash. Acad. Sci. 14:
243. 1924. Rehdera mollicella Standl. & Moldenke in Fedde, Rep.
Sp. Nov. 39: 51. 1935 (type from Baja Verapaz, Kellerman 7992).
Dry, brushy, often rocky plains and hillsides, 200-800 meters;
Baja Verapaz; Chiquimula; Jutiapa. Mexico; Honduras; El Salva-
dor; Nicaragua; Costa Rica.
Trees to 20 m. tall, the branches puberulent or glabrate; leaves on puberulent
petioles 5-15 mm. long, the blades subcoriaceous, broadly or narrowly elliptic or
obovate, 2-9 cm. long, usually rounded or obtuse, rarely acute or abruptly short-
acuminate at the apex, cuneate or obtuse and abruptly contracted at the base, the
margins entire, lateral veins 3-7 pairs, plinerved or not, glabrate and often lus-
trous above, glabrous or more or less pubescent beneath, or pubescent only along
the costae; racemes axillary, usually 1-2 cm. long, glabrous or puberulent, with few
or many sessile flowers; bracts linear or narrowly triangular, puberulent, usually
1-3 mm. long; calyx tubular at an thesis, in fruit cupuliform, glabrous or minutely
puberulent outside, sericeous inside, 4-6 mm. long, 5-costate and denticulate; co-
rolla white, cream, or greenish, glabrous outside, the tube 4-6 mm. long, the lobes
D.iSuJotl t)
FIG. 43. Rehdera trinervis. A, fruiting branch, Y^ natural size; B, flower, X 4;
C, corolla opened to show stamens, X 4; D, calyx opened to show pistil, X 5.
223
224 FIELDIANA: BOTANY, VOLUME 24
pilose or tomentose inside, 4-5 mm. long; fruit oblong or obovate-oblong, 8-20 mm.
l°ng. glabrous and lustrous.
Since Standley and Moldenke described R. mollicella, numerous
intermediate specimens from various localities have been collected,
in which some of the smaller leaves appear triplinerved, but with 5-7
pairs of lateral veins in some larger leaves, these anastomosing near
the margin or not. Standley 28593 from Honduras, Standley 94.12
from Nicaragua, and Jimenez 312 from Costa Rica have leaves that
are essentially glabrous but with 5-7 conspicuous pairs of lateral
veins. Standley 28600 from Honduras has uniformly small, tripli-
nerved leaves, but they are densely pubescent beneath. Molina
14321 and 14350 from Honduras have most leaves triplinerved but
all have patches of indument along the costae and most have sparsely
scattered pubescence on the lower surfaces. Calyces of these inter-
mediate specimens are usually glabrous, but a few on Molina 14321
are somewhat puberulent.
STACHYTARPHETA Vahl
Annual or perennial herbs or low shrubs, glabrous or pubescent; leaves mostly
opposite, the blades dentate to serrate; inflorescences terminal, spicate, the spikes
short or long, densely or laxly flowered, the flowers numerous, sessile or partly im-
mersed in the rachis of the spike, each flower solitary in the axis of a bract; calyx
tubular, 5-dentate, the teeth usually unequal; corolla small, white to blue, purple,
pink, or red, the tube cylindric, the limb spreading, 5-lobate; stamens 2, inserted
above the middle of the corolla tube, included, staminodes 2, small and inconspicu-
ous; anthers not appendaged, the cells divergent, dehiscent in one continuous line;
ovary bilocular, each locule uniovulate; style elongated, the stigma terminal, sub-
capitate; ovules attached laterally near the base of the locule; fruit dry, oblong:
linear, included in the persistent calyx, splitting at maturity into 2 hard cocci;
endosperm none.
Perhaps 30 or 40 species, in tropical America. One other species,
S. mutabilis (Jacq.) Vahl, is known from Central America.
Leaves linear or oblong-linear, 6-10 times longer than broad S. angustifolia.
Leaves not as above.
Rachis of inflorescence at maturity thick and stout, 4-7 mm. in diameter.
S. jamaicensis.
Rachis of inflorescence at maturity slender, less than 4 mm. in diameter.
Mature rachis usually less than 2 mm. in diameter; calyx 4-6 mm. long.
S. cayennensis.
Mature rachis 2-3.5 mm. in diameter; calyx 7-12 mm. long.
Bracts conspicuously outcurved; leaves glabrate or the pubescence incon-
spicuous, usually confined to costae and veins of lower surface.
S. miniacea.
Bracts appressed or only slightly divergent; leaves usually pubescent on
both surfaces S. frantzii.
GIBSON: FLORA OF GUATEMALA 225
Stachytarpheta angustifolia (Mill.) Vahl, Enum. PL 1: 205.
1804. Verbena angustifolia Mill. Card. Diet. ed. 8, no. 15. 1768.
Open places, clearings, weedy fields, often near wet places or
along roadsides, sea level to 900 meters; Pete"n. Mexico; British
Honduras; Honduras; West Indies; northern South America.
Erect annuals, to 1 m. tall, essentially glabrous, the stems terete or subterete;
leaves linear or oblong linear, 4-14 cm. long, usually less than 1 cm. wide, rarely
to 1.5 cm., attenuate at each end, serrate-dentate, often somewhat hirsutulous
beneath on the veins; mature spikes of inflorescence usually stout, 3-5 mm. wide,
10-40 cm. long; bracts ovate or lanceolate, subulate-acuminate, 5-6 mm. long;
calyx 6-7 mm. long; corolla blue or purple.
Stachytarpheta cayennensis (L. Rich.) Vahl, Enum. PL 1:
208. 1804. Verbena cayennensis L. Rich., Act. Soc. Hist. Nat. Paris
1: 105. 1792. S. guatemalensis Moldenke, Carnegie Inst. Wash.
Publ. 522: 181. 1940 (type from Coban, Alta Verapaz, Tuerckheim II.
1823). S. tabascana Moldenke, Phytologia 1: 437. 1940. S. guate-
malensis L albiflora Moldenke, Phytologia 9: 99. 1963. Styetolok
(Alta Verapaz, Quecchi) ; San Diego vervena (Pete"n) .
Damp thickets, forest, or swamps, sometimes in pine forest, often
a weed in waste places, sea level to 1,500 meters; Alta Verapaz; Chi-
maltenango; Izabal; Pete"n. Mexico; British Honduras to Panama;
West Indies; South America; said to be naturalized in some parts of
the Old World.
Plants erect and stiff, branching, to 1 m. tall, sometimes suffrutescent below,
the stems subterete, pubescent or glabrate; leaves petiolate, the blades ovate or
elliptic, 3-10 cm. long, rounded to acute at the apex, cuneate or attenuate to the
base, the margins crenate-serrate, usually scaberulous above, usually glabrate be-
neath, often strigillose on the veins; inflorescence mostly 10-40 cm. long, glabrous
or nearly so, the rachis very slender, usually 2 mm. or less in diameter; bracts linear
or lanceolate, often subulate, 3.5-5 mm. long; calyx 4-6 mm. long; corolla usually
blue or purple, rarely pink or white, the limb to 5 mm. broad.
The characters Moldenke uses in separating S. guatemalensis and
S. tabascana from S. cayennensis do not hold in all specimens deter-
mined by him.
The Indians of Alta Verapaz employ the plant as a supposed
remedy for malaria and other fevers. In British Honduras it is used
to treat dysentery.
Called "mozote" in Honduras and "wild verbena" and "camac-
olal" (Maya) in British Honduras.
FlG. 44. Stachytarphetafrantzii. A, flowering and fruiting branches, ^natural
size; B, flower, X 5; C, corolla dissected to show stamens, X 5; D, calyx opened to
show pistil, X 5; E, fruit, X 5.
226
GIBSON: FLORA OF GUATEMALA 227
Stachytarpheta frantzii Polak. Linnaea 41: 593. 1877. S.
frantzii var. patentiflora Moldenke, Phytologia 1: 430. 1940 (type
from Guatemala, Tonduz 626). S. guatemalensis var. lundelliana
Moldenke, I.e. S. robinsoniana Moldenke, Carnegie Inst. Wash.
Publ. 522: 182. 1940. Cola de armado (Guatemala); pije de gato
( Jutiapa) .
Damp fields or thickets, 150-1,500 meters; Guatemala; Huehue-
tenango; Jutiapa; Santa Rosa. Southern Mexico to Costa Rica.
Annuals or perennials to 1 m. tall, herbaceous throughout or suffrutescent be-
low, usually much branched, the stems often tetragonous, pilose, tomentose, or
glabrate; leaves on winged petioles or the blades often cuneate to the base, ovate
or oblong-ovate, 3-10 cm. long, acute or obtuse, coarsely serrate, scabrous to pilose
or strigose-pubescent above, pubescent to tomentose beneath, often densely so;
inflorescence 8-30 cm. long, the rachis slender, usually 2-3 mm. in diameter,
densely or sparsely pilosulous or glabrate, many-flowered; bracts narrowly lanceo-
late, 6-8(9) mm. long, appressed or slightly divergent; calyx 7-11(12) mm. long;
corolla normally blue, violet, or purple, rarely white, red, or pink, the tube 10-
16 mm. long.
This has been reported from Guatemala as S. mutabilis (Jacq.)
Vahl, a more robust plant with the rachis of the inflorescence often
5-6 mm. in diameter.
Stachytarpheta jamaicensis (L.) Vahl, Enum. PL 1: 206. 1804.
Verbena jamaicensis L. Sp. PL 19. 1753. Stachytarpheta jamaicensis
f . albiflora Standley, Field Mus. Bot. 4 : 320. 1929. Chilillo (Jutiapa) .
Usually on sea beaches, sometimes along roadsides, sea level to
900 meters; Izabal; Jutiapa. Southern United States; Mexico; Brit-
ish Honduras to Panama; West Indies; northern South America.
Erect herbs to 1 m. tall, essentially glabrous throughout, the stems terete or
obscurely tetragonous; leaves petiolate, the blades usually broadly ovate to oblong-
ovate, sometimes oblong-elliptic, 3-10 cm. long, obtuse or acute, cuneate at the
base and usually long-decurrent, coarsely serrate; inflorescence glabrous, stiff,
stout, the mature rachis 4-7 mm. in diameter, 10-70 cm. long; bracts lanceolate
or oblong-lanceolate, 4-6 mm. long; calyx 5-7 mm. long; corolla usually blue or
purple, rarely white.
Called "verbena" in Honduras and, fide Standley, "mes" (Que-
cchi); "ibinxiu" and "talche" (Yucatan, Maya).
Stachytarpheta miniacea Moldenke, Phytologia 1: 170. 1935.
S. petenensis Moldenke, Phytologia 9: 99. 1963 (type from Dos La-
gunas, Pete"n, Contreras 1563).
228 FIELDIANA: BOTANY, VOLUME 24
Along roadsides or in thickets or clearings, 200 meters or less,
Pete"n. Mexico (Campeche) and British Honduras (type from Fresh-
water Creek Reserve, Corozal District, R. S. Pelly 7) .
Erect annuals or perennials, sometimes suffrutescent below, to 1 m. tall, the
stems subterete, puberulent or glabrate; leaves petiolate, the blades lance- oblong,
3-13 cm. long, 1.5-3.5 cm. wide, acute at apex, long-attenuate and usually decur-
rent to the base of the petiole, coarsely serrate, glabrate on both surfaces or puber-
ulent beneath; inflorescences slender, the rachis 2-3.5 mm. in diameter, 15-40 cm.
long, sparsely pilosulous; bracts narrowly lanceolate, 8-11 mm. long, conspicu-
ously outcurved; calyx about equalling the bracts; corolla dull red.
This appears to be closely related to S. frantzii; the bracts of the
latter are not as conspicuously divergent and its leaves are usually
pubescent on both surfaces.
TAMONEA Aublet
Erect, essentially annual, branched herbs, sometimes suffrutescent below;
leaves small, opposite, short-petiolate, dentate or incised; inflorescences spicate
or racemose, terminal and axillary, few-flowered, the flowers remote, small, sessile
or short-pedicellate, bracteate; calyx tubular in anthesis, campanulate in fruit,
subtruncate, costate, the 5 costae prolonged into short teeth; corolla blue, the tube
cylindric, slightly ampliate above, the 5 lobes spreading, oblique; stamens 4, didy-
namous, included, the filaments inserted about the middle of the corolla tube, the
2 longer ones usually bearing hump-like appendages below the anthers; anthers
ovate, the cells parallel; ovary bicarpellate with 4 almost completely separated
locules, each locule uniovulate; style terminal, stigma oblong, oblique; ovules
attached laterally near the base of the locule; fruit drupaceous, but becoming nut-
like in ours, globose or turbinate, little exserted from the accrescent calyx, obtuse or
with 4 obscure lobes at the apex, sometimes spinose at the apex, the exocarp only
slightly fleshy, the endocarp hard; endosperm none.
We use the generic name Tamonea Aublet for this verbenaceous
genus, following Miss Green in Rehder, Weatherby, Mansfeld &
Green (Kew Bull. 1935: 508. 1936). Dr. Williams has pointed out
(Fieldiana: Bot. 29: 574. 1963) that Aublet obviously changed the
name of the melastomaceous "Tamonea" while the work in which
the two "Tamoneas" was going through press and used Fothergilla
for the melastome. Aublet's obvious intent should be followed.
Perhaps four or five species in tropical America. Only one has
been found in Central America.
Tamonea spicata Aubl. PI. Guian. 660, t. 268. 1775. Ghinia
spicata Moldenke, Phytologia 1 : 169. 1935.
In savannas, about 100 meters; Alta Verapaz. British Honduras;
Nicaragua; WTest Indies; South America.
FIG. 45. Tamunea spicata. A, habit, l/2 natural size; B, fruit partially en-
closed in calyx, X 5; C, calyx opened to show pistil, X 5; D, corolla dissected to
show stamens, X 4.
229
230 FIELDIANA: BOTANY, VOLUME 24
Slender, branched herbs to 1 m. tall, the stems tetragonous, puberulent with
small, usually curved hairs; leaves on petioles 3-13 mm. long, the blades ovate to
broadly ovate, 0.5-2.5 cm. long, obtuse, truncate at the base, coarsely crenate or
incised-serrate, sparsely or rather densely pilosulous; racemes 3-8 cm. long, the
pedicels ca. 2 mm. long; calyx at an thesis 4 mm. long; corolla blue, 67 mm. long;
fruit 3-4 mm. long, glabrous, not spinose at the apex.
The Mexican T. curassavica (L.) Pers. and the West Indian T.
subbiflora Urb. & Ekm., which might be confused with ours, have
fruits which are spinose at the apex.
VERBENA L.
Reference: Lily M. Perry, A revision of the North American spe-
cies of Verbena, Ann. Mo. Bot. Card. 20: 239-362. 1933.
Annual or perennial herbs, the stems erect to prostrate, glabrous or pubescent;
leaves opposite, dentate, serrate, or variously lobate or dissected; inflorescence
spicate, the spikes terminal, densely or laxly flowered, short and headlike or elon-
gated, sometimes appearing paniculate, the flowers usually sessile, inconspicuous
or showy; calyx tubular, with 5 costae terminating in 5 unequal teeth; corolla sal-
verform or funnelform, the tube straight or curved, the limb obscurely bilabiate,
with 5 lobes; stamens 4, didynamous, inserted in the upper half of the corolla tube;
anthers ovate, the 2 cells parallel or slightly divergent; style short, the stigma bi-
lobate; ovary bicarpellate, each carpel bilocular, each locule uniovulate; fruit
usually enclosed in the calyx, dry, separating into 4 linear or oblong trigonous
cocci, each with one seed.
About 100 species, almost all American. Only the following are
known from Central America. Two of the garden verbenas, V. teu-
crioides Gill. & Hook, and V. tenuisecta Briq., both native of South
America, are widely grown in Guatemala at all elevations, but there
is no evidence of their having become naturalized.
Leaves deeply lobate, cleft, incised, or dissected.
Inflorescence short and compact, 1-2 cm. long; calyx 4-7 mm. long.
V. teucriifolia.
Inflorescence long and slender, 3-45 cm. long; calyx less than 3 mm. long.
V. menthaefolia.
Leaves crenate-serrate to serrate, but never lobate, incised, nor dissected.
Stems hirsute-hispid to pilose; leaves mostly lanceolate to broadly elliptic, the
margins coarsely crenate-serrate; bracts usually not more than half as long
as the calyx V. Carolina.
Stems glabrous or sparsely strigose; leaves mostly oblanceolate to spatulate, the
margins serrate; bracts usually equalling or two-thirds as long as the calyx
V. litoralis.
Verbena Carolina L. Syst. ed. 10. 852. 1759. V. polystachya
HBK. Nova Gen. & Sp. 2: 274. 1818. V. hirsute Mart. & Gal. Bull.
FIG. 46. Verbena Carolina. A, habit, % natural size; B, flowers, buds, and
calyces, showing subtending bracts, X 6; C, corolla dissected to show stamens,
X 8; D, calyx opened to show pistil, X 10.
231
232 FIELDIANA: BOTANY, VOLUME 24
Acad. Brux. 11 (2): 321. 1844. V. mollis Mart. & Gal. I.e. 323.
V. paucifolia Mart. & Gal. I.e. 324. Dori (Alta Verapaz, Quecchi) ;
chichavac (Chimaltenango) ; verbena (Guatemala).
Damp or dry thickets, meadows, fields, frequently a weed in cul-
tivated ground, sometimes in pine-oak forest, rarely on sand bars;
usually 1,300-3,100 meters, rarely at 85 meters; Alta Verapaz; Chi-
maltenango; Guatemala; Huehuetenango ; Izabal; Jalapa; Quezalte-
nango; El Quiche"; Santa Rosa; Solola. Arizona; Mexico; El Salva-
dor; Honduras.
Annuals or perennials, erect, usually branching, the stems hirsute-hispid to
pilose; leaves subsessile or narrowed into a short petiole, the blades lanceolate to
subelliptic, 2-8 cm. long, obtuse or acute, the margins coarsely crenate-serrate,
somewhat strigose above, hirsute-strigose beneath, especially on costa and veins;
inflorescence often irregularly disposed and appearing paniculate, 3-30 cm. long,
many-flowered, often interrupted in fruit; bracts ovate, acuminate, ciliate, usually
not more than half as long as the calyx; calyx 1.5-2 mm. long, the teeth connivent
in fruit; corolla ca. 2 mm. long, pale blue, lavender, or white; fruit ca. 1.5 mm. long.
This has been reported from Guatemala as V. ehrenbergiana
Schauer.
Verbena litoralis HBK. Nova Gen. & Sp. 2: 276, 1. 137. 1818.
V. affinis Mart. & Gal. Bull. Acad. Brux. 11(2) : 322. 1844. V. han-
senii Greene, Pittonia 3: 308. 1898. V. litoralis var. albiflora Mol-
denke, Phytologia 1: 432. 1940. Verbena; verbena cimarrona; verbena
fina; cotacam (Quecchi, fide Standley).
Wet or dry thickets, meadows, rocky slopes, often a weed in cul-
tivated ground, sometimes in pine-oak forest, 85-3,000 meters; Alta
Verapaz; Baja Verapaz; Chimaltenango; Guatemala; Huehuete-
nango; Izabal; Jalapa; Jutiapa; Pete"n; Quezaltenango; Retalhuleu;
Sacatepe"quez ; San Marcos; Santa Rosa; Totonicapan. Mexico to
Panama; South America.
Annual or perennial herbs, to 1.5 m. tall, branched above, the stems tetrago-
nous, glabrous or sparsely strigose; leaves subsessile or short-petiolate, the blades
oblanceolate to spatulate, often very narrowly so, 2-10 cm. long, acute, serrate,
sparsely strigillose on both surfaces, usually rough to the touch, prominently
veined beneath; inflorescence often appearing paniculate, the spikes slender,
densely flowered or interrupted, the flowers often congested near the end of the
rachis; bracts ovate-lanceolate, acuminate, ciliate, two-thirds as long as, or equal-
ling the calyx; calyx 2-2.5 mm. long, strigillose, the teeth minute; corolla tube
longer than the calyx, the limb 2.5-3 mm. broad, blue, purple, or white; fruit
scarcely 2 mm. long.
GIBSON: FLORA OF GUATEMALA 233
Common weeds of Central America. In Alta Verapaz the plant
has been employed as a remedy for various fevers, as well as influenza
and small-pox.
Called verbena de montana in British Honduras.
Verbena menthaefolia Benth. PI. Hartweg. 21. 1839. V. se-
tosa Mart. & Gal. Bull. Acad. Brux. 11(2): 321. 1844. Telran;
verbena.
Open fields or hillsides, 800-1,200 meters; Huehuetenango (be-
tween Nenton and Las Palmas, Steyermark 51655). Southwestern
United States; Mexico.
Perennial plants, erect or decumbent, branched, the stems sparsely and mi-
nutely hispidulous; leaves tapering at the base into a marginate petiole, the blades
1-5 cm. long, deeply lobed or incised, the divisions narrow, remotely serrate-
dentate, more or less strigillose on both surfaces; inflorescence irregularly disposed
and appearing paniculate, very slender, usually much elongated, 3-45 cm. long,
many-flowered, interrupted; bracts ovate-lanceolate, acuminate, usually shorter
than the calyx, ciliate and sparsely strigillose; calyx 2-2.5 mm. long, strigillose,
the teeth minute; corolla ca. 4 mm. long; fruit 2-2.5 mm. long.
Verbena teucriifolia Mart. & Gal. Bull. Acad. Brux. 11, pt. 2:
322. 1844. Verbena de monte (Huehuetenango).
Usually in open, grassy or stony hillsides, fields, roadsides, some-
times in scrub forest, 1,500-3,100 meters; Chimaltenango; Guate-
mala; Huehuetenango; Quezaltenango; San Marcos; Solola; Toto-
nicapan. Southern Mexico.
Annuals or perennials, prostrate, often forming small, dense mats, usually
much branched, hirtellous or glabrate; leaves 1-3 cm. long, cuneate at the base
and short-petiolate, incised-pinnatifid, the segments linear-oblong, obtuse or sub-
acute, strigillose or glabrate; inflorescence sessile or short-pedunculate, 1-2 cm.
long, few-flowered; bracts ca. half as long as the calyx, lanceolate; calyx 4-7 mm.
long, hirtellous, accrescent, in fruit often contracted at orifice with the aristiform
lobes twisted together; corolla rose-purple, the tube slightly longer than the calyx,
the limb 4-7 mm. broad; fruit 2.5-3 mm. long, the cocci reticulate-scrobiculate, the
inner face muriculate or almost smooth.
Inconspicuous plants, common in the region of Quezaltenango.
VITEX L.
Reference: Moldenke, Harold N. Materials toward a monograph
of the genus Vitex, I-X. Phytologia 5: 142-176, 186-224, 257-280,
293 336, 343 393, 404-507. 1955 1957; 6: 13-64, 70 192. 1957.
234 FIELDIANA: BOTANY, VOLUME 24
Trees or shrubs, glabrous or variously pubescent; leaves opposite, palmately
compound, petiolate, the leaflets 3-7, petiolulate, usually entire, the terminal leaf-
let largest and the 2 lower ones often greatly reduced; inflorescence cymose, the
cymes short and dense or lax and spreading, in axillary or terminal panicles, the
flowers more or less zygomorphic; calyx campanulate or cupuliform, with 5 teeth
or lobes; corolla usually blue, violet, or white, salverform, the tube short, the limb
oblique, spreading, somewhat bilabiate, the upper lip bifid, the lower and larger lip
trifid; stamens 4, didynamous, often exserted, the filaments inserted on the corolla
tube; anther cells parallel, divergent, or arcuate, attached near their apex, dehis-
cent by longitudinal slits; ovary bicarpellate, the style terminal, filiform, short-
bifid at the apex; the 4 locules of the ovary uniovulate, the ovules attached
laterally; fruit drupaceous, the endocarp hard, the exocarp juicy; seeds erect,
without endosperm.
Perhaps 250 species, in the tropics of both hemispheres. Three
occur in Guatemala and a few others are known from southern Cen-
tral America.
Leaflets 3; calyx cupuliform, subtruncate, remotely and minutely denticulate.
V. cooperi.
Leaflets 5 (rarely 7) ; calyx campanulate, lobate or dentate.
Leaflets glabrous beneath or nearly so; calyx with triangular-oblong to linear,
often reflexed lobes 1.5-2.5 mm. long V. kuylenii.
Leaflets tomentulose or velutinous beneath; calyx with acute teeth 0.5-1 mm.
long V. gaumeri.
Vitex cooperi Standl. Trop. Woods 16: 26. 1928 (nomen); Field
Mus. Bot. 4: 256. 1929.
Damp thickets, sea level to 600 meters; Izabal. Atlantic coast
of Honduras; Nicaragua; Costa Rica; Panama.
Large trees to 26 m. tall, to 75 cm. in diameter, branchlets first puberulent,
then glabrate; leaves on minutely puberulent petioles 1.3-9.5 cm. long, 3-foliate,
the petiolules 1-8 mm. long, the blades mostly 4-22 cm. long, the central blade
much larger than the lateral ones, chartaceous, broadly elliptic, elliptic-oblong, or
obovate, short-acuminate or abruptly acuminate, acute or abruptly contracted at
the base, glabrous above, puberulent beneath when young, glabrate in age; in-
florescences axillary, cymose, 3.5-14 cm. long, usually 3 or 4 times dichotomously
branched, peduncles 4-8 cm. long, finely puberulent, the flowers on puberulent
pedicels 1-3 mm. long; calyx 1-1.5(2) mm. long, cupuliform, subtruncate, puberu-
lent, remotely and minutely denticulate; corolla blue to purple, the tube 3-4 mm.
long, densely glandular-puberulent, villous in throat, the lobes 3-6 mm. long, often
minutely puberulent within; stamens glabrous except villous near base; style gla-
brous, ovary glabrous; fruit obovoid, to 13 mm. long, glabrous, black.
This has been confused with V. floridula Duchass. & Walp. of
Panama, which has larger flowers and short-pedunculate cymes. It
more closely resembles the West Indian V. divaricata Sw. which has
slightly larger flowers and glabrate pedicels and calyces.
GIBSON: FLORA OF GUATEMALA
235
FIG. 47. Vitex gaumeri. A, section of plant with inflorescences, Y^ natural
size; B, flower, X 3.5; C, corolla dissected to show stamens, X 3.5; D, calyx opened
to show pistil, X 3.5; E, fruit, X 1.
It should be noted that the only two Guatemalan collections,
both determined by Moldenke, are sterile.
Vitex gaumeri Greenm. Field Mus. Bot. 2: 260. 1907. V. lon-
ger acemosa Pittier, Contr. U. S. Nat. Herb. 20: 486. 1922 (type from
Quirigua, Izabal, H. N. Whitford & L. R. Stadtmiller 74)- Jocote de
mico (Izabal); yaxnic (Pete*n, Maya).
236 FIELDIANA: BOTANY, VOLUME 24
Damp forest, often on limestone, occasional on pine ridges and
in poor soil, sometimes in second growth, sea level to 600 meters;
Alta Verapaz; Izabal; Pete"n. Chiapas and Yucatan Peninsula of
Mexico; British Honduras; Honduras.
Large trees, sometimes 30 m. tall, with broad, spreading crowns, the trunks to
75 cm. in diameter, with brown, ridged bark, the young branchlets densely puberu-
lent, whitish; leaves on tomentulose petioles 6-16 cm. long, leaflets 5, rarely 7, on
petiolules 0.5-3 cm. long, the blades chartaceous, elliptic or elliptic-oblong, mostly
6-22 cm. long, entire, acute or acuminate, usually obtuse or rounded at the base,
puberulent or glabrate above, usually whitish or grayish beneath and densely to-
mentulose or velutinous; inflorescences axillary, cymose, in panicles 8-30 cm. long,
the flowers short-pedicellate; calyx campanulate at anthesis, patelliform in fruit,
2-3 mm. long, densely puberulent, with 5 acute teeth; corolla blue, sparsely puber-
ulent outside, sometimes minutely papillate, the tube 4-5 mm. long, 2-3 mm. wide,
sparsely villous in throat, the lobes equalling or shorter than the tube; filaments of
stamens pubescent, more densely so near base; style glabrous or nearly so, the
ovary densely pubescent; fruits somewhat fleshy, yellow, depressed- globose, mi-
nutely puberulent, 12-20 mm. broad.
A tree of this species from Pete"n was reported by H. H. Bartlett
in 1931 to have been six feet in diameter. In British Honduras, it
is sometimes called "fiddlewood," "blue blossom," and "matasano."
V. gaumeri is much like V. hemsleyi Briq. of Mexico, but differs
in its much heavier indument and smaller flowers.
Vitex kuylenii Standl. Trop. Woods 8: 6. 1926. Barabas (Izabal).
Damp forest, sometimes in pine forest, sea level to 150 meters;
Alta Verapaz; Izabal (type from Izabal, Henry Kuylen 66); Chiqui-
mula. Southern Mexico; British Honduras; Atlantic lowlands of
Honduras and Nicaragua.
Trees, 10-15 m. tall, with trunks 25 cm. in diameter or perhaps larger, the
branchlets glabrate; leaves on petioles 4-9 cm. long, leaflets usually 5 (rarely 7),
on petiolules 4-13 mm. long, the blades chartaceous, glabrous or nearly so, often
lustrous, entire, elliptic or oblong-elliptic, 4-20 cm. long, acute or acuminate, acute
at the base; inflorescence axillary, cymose, in long-pedunculate panicles, 10-15 cm.
long, the branches minutely puberulent; flowers short-pedicellate; calyx campanu-
late in anthesis, patelliform in fruit, puberulent, the tube 2-3 mm. long, the lobes
triangular-oblong to linear, subacute, 1.5-2.5 mm. long, usually becoming reflexed;
flowers fragrant, corolla light blue to purple, minutely puberulent outside, villous
in throat, the tube 6-7 mm. long, 1-2 mm. wide, the longest lobe 5-6 mm. long,
4-6 mm. broad; filaments of stamens pubescent, villous near base; style pubescent,
ovary densely pubescent to velutinous; fruits fleshy, yellow, depressed-globose,
minutely puberulent, to 15 mm. broad.
Publication 1100
UNIVERSITY OF ILLINOIS-URBANA