L I B RAR.Y
7? d. 7
FLORA OF GUATEMALA
PAUL C. STANDLEY
LOUIS 0. WILLIAMS
VOLUME 24, PART VII, NUMBER 2
CHICAGO NATURAL HISTORY MUSEUM
JANUARY 26, 1962
FLORA OF GUATEMALA
FLORA OF GUATEMALA
PAUL C. STANDLEY
Curator Emeritus of the Herbarium
LOUIS 0. WILLIAMS
Curator, Central American Botany
VOLUME 24, PART VII, NUMBER 2
CHICAGO NATURAL HISTORY MUSEUM
JANUARY 26, 1962
Library of Congress Catalog Card Number: ]>8-8076
PRINTED IN THE UNITED STATES OF AMERICA
BY CHICAGO NATURAL HISTORY MUSEUM PRESS
Families Included in Part VII, Number 2
Cactaceae. ..................... 187 Punicaceae. .
Thymelaeaceae ................. 234 Lecythidaceae . .
Lythraceae ..................... 239 Rhizophoraceae ....... - 263
Combretaceae .................. 268
LIST OF ILLUSTRATIONS
Deamia testudo 194
Epiphyllum Eichlamii 199
Epiphyllum Nelsonii 201
Epiphyllum quezaltecum 203
Hylocereus undatus 207
Pereskia aculeata 223
Rhipsalis ramulosa 229
Wilmattea minutiflora 233
Daphnopsis radiata 238
Cuphea hyssopifolia 248
Cuphea pinetorum 251
Lafoensia punicifolia 254
Lythrum vulneraria 258
Rotala dentifera 259
Cassipourea guianensis 265
Rhizophora Mangle 267
Bucida macrostachya 270
Combretum fruticosum 274
Terminalia oblonga 280
Flora of Guatemala 1
The Opuntiales, as outlined by Engler, contain a single family,
the Cactaceae. The family, with the possible exception of a few
species of Rhipsalis, is entirely American. A few cacti have become
established in other parts of the world. The Cactaceae occur in
greatest abundance, both in kinds and individuals in the dry tropics
or adjacent desert or desert-like areas in temperate regions. A few
species extend well into the cool or even cold temperate regions.
The center of greatest diversification is in Mexico and the adjacent
southwestern United States, with a secondary center outside the
wet tropics in South America.
The characters of the order are those of the single family which
CACTACEAE. Cactus Family
Succulent, perennial plants, often of bizarre forms, mostly armed with spines,
the stems various in form, plants usually terrestrial but those of the wetter tropics
often epiphytic, small and herb-like or often large and shrub-like or tree-like, fleshy
or the stems often hard and woody, simple or branched, branches often articulate,
bearing organs called areoles which may be either small or large and tuft-like or
circular and often bearing bristles and wool or hair and usually spines; leaves none
except in Pereskia and Pereskiopsis where they are broad, succulent and entire,
the leaves in some genera represented by terete or subulate scales which are soon
deciduous; spines various in arrangement and number, sometimes vaginate, often
wanting; flowers perfect or rarely unisexual, regular or somewhat irregular, small
or often very large, usually solitary, but sometimes borne in a specialized cap-like,
terminal inflorescence called a cephalium; perianth tube none or present and greatly
elongated, the segments of the limb spreading or erect and few or numerous; sepals
usually grading into the petals, but the sepals and petals sometimes unlike; stamens
sometimes few but usually numerous, short or elongate, sometimes borne in sep-
arated series; filaments inserted on the tube or the throat of the perianth, the an-
thers small, usually oblong, 2-celled; style simple, terminal, often much elongated,
lobes of the stigma 2-many, generally slender; ovary inferior, 1-celled, distinct or
sometimes immersed in the branch, ovules numerous; fruit baccate, often juicy and
edible; seeds usually numerous; cotyledons 2, accumbent, often broad or elongate;
endosperm scant or copious.
'Assisted by a grant from National Science Foundation.
188 FIELDIANA: BOTANY, VOLUME 24
Perhaps 50 genera (26, Vaupel; 124, Britton and Rose; 220,
Backeberg) and some 1,500 species. Two others of the genera
recognized by Britton and Rose, Wittia and Weberocereus, are found
in southern Central America.
The number of genera in the Cactaceae has been a matter
of considerable controversy during the past one hundred years.
Bentham and Hooker, in the Genera Plantarum (1867), were able
to "crowd" all the cacti into 13 genera. Britton and Rose, in The
Cactaceae (1919-23), perhaps the most useful work on Cactaceae
and the only one reasonably complete for Guatemala, accepted
124 genera, many of their own creation. Vaupel, in the second
edition of the Pflanzenfamilien (1925), a few years later, accounted
for the family in 26 genera. The most recent comprehensive work
on the Cactaceae is Kurt Backeberg's Die Cactaceae (1958-61, five
volumes and with perhaps one or two more to be expected) in
which it is indicated that there are 220 genera (or perhaps "micromi-
crogenera"?) in the family!
The generic names used in the following text are mostly those
used by Britton and Rose. These names are not used because we
agree with them but because Britton and Rose's publication, The
Cactaceae, is so well known to most cactus fanciers and to botanists
that the names used will be intelligible to most people who will
use this work.
If most European botanists (Backeberg excepted) have erred on
the side of conservatism in a generic concept for the Cactaceae, Brit-
ton and Rose and Backeberg have gone to the other extreme. Many
of the genera proposed or maintained by these gentlemen are based
on trivial characters. It remains for some competent systematist to
give us a taxonomically sound classification of the family.
In Central America cacti are not well represented and most of
those found belong to epiphytic groups rather than the terrestrial
ones that prevail in desert regions. The number of species is greater
in Guatemala than in any other Central American country.
Cacti are among the most fantastic and bizarre of all plants,
because of their strange forms and the curious modifications of
some of the normal plant organs. They attracted attention from
the earliest of the European invaders of America and some of them
have long been in cultivation in Europe and elsewhere. Many
exotic cacti, particularly Mexican ones, are seen now and then in
Guatemalan gardens; these we have not attempted to list here but
we have included a few of the more common of the introduced cacti.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 189
Plants with normal broad leaves, scandent or shrub-like (leaves sometimes decid-
Glochids present in the areoles; flowers pedicellate, often paniculate. .Pereskia.
Glochids none in the areoles; flowers sessile Pereskiopsis.
Plants without leaves, or the leaves reduced to small narrow scales, these soon
deciduous. Plants armed with spines or some kinds unarmed.
Areoles with glochids; young parts bearing leaves, these narrow and subulate
or scale-like, soon deciduous; perianth without a tube; plants usually abun-
dantly armed with spines.
Petals erect; stamens longer than the petals , Nopalea.
Petals spreading; stamens shorter than the petals Opuntia.
Areoles without glochids; plants without leaves; perianth with a short or much
elongate tube, rarely rotate in Rhipsalis; plants spiny or unarmed.
Plants unarmed, epiphytic.
Perianth rotate, very small, less than 1 cm. long; stems terete or com-
Perianth with an elongate tube, much more than 1 cm. long; stems various.
Plants dichotomously branched; flowers bright red, very irregular; culti-
vated plants Zygocactus.
Plants irregular branched; native plants Epiphyllum.
Plants armed with small or large spines, mostly terrestrial but sometimes
Plants globose or short-cylindric, small, mostly less than 20 cm. high.
Plants with a cap-like or cushion-like cephalium at the apex. .Melocactus.
Plants without a cephalium Mammillaria.
Plants with much elongate stems, many times as long as thick, usually large,
often vine-like or tree-like.
Plants tree-like, large, erect, usually with few or numerous thick, erect
or ascending branches, rarely simple but then tall and erect.
Branches densely covered at the ends with long white hair-like bristles.
Branches without a covering of white hair-like bristles at the ends.
Flowers 2-several at each areole; flowers about 3 cm. long.
Flowers solitary at the areoles, larger.
Corolla short-campanulate; fruit dry Pachycereus.
Corolla short-f unnelf orm ; fruit juicy Lemaireocereus.
Plants smaller, never tree-like, usually vine-like, often epiphytic.
Stems with 7-12 ribs.
Flowers short-f unnelf orm, rose-red; cultivated plants . .Aporocactus.
Flowers elongate-f unnelf orm, white; native plants.
Spines 1 cm. long or less; flowers mostly 18-20 cm. long.
Spines, at least the largest ones, 3-4 cm. long; flowers 4-7 cm. long.
Stems mostly with 3 angles or wings, rarely 4-5-angulate.
Ovary and fruit covered with large foliaceous scales, their axils with-
out spines, hairs, or bristles.
190 FIELDIANA: BOTANY, VOLUME 24
Perianth tube elongate; flowers large, 25-30 cm. long, scales naked
in the axils Hylocereus.
Perianth tube very short; flowers small, about 5 cm. long, some of
the scales with tufts of short hairs in the axils . . . Wilmattea.
Ovary and fruit not bearing large foliaceous scales, their axils spiny,
hairy, or setiferous.
Flowers red, open during the day Heliocereus.
Flowers white, mostly nocturnal.
Areoles of the ovary bearing long hairs; flowers diurnal, about
28 cm. long Deamia.
Areoles of the ovary bearing short spines; flowers 10-20 cm. long.
Perianth tube elongate, much longer than the limb; flowers
14-20 cm. long Acanthocereus.
Perianth tube much shorter than the limb; flowers about
10 cm. long Werckleocereus.
ACANTHOCEREUS Britton & Rose
Plants mostly terrestrial, the stems elongate, rather weak, articulate, at first
erect, becoming clambering, trailing, or recurved, usually deeply triangulate, some-
times 4-5-angulate; areoles bearing short wool or felt and several stout spines;
flowers nocturnal, funnelform, 1 at an areole, the tube rigid after anthesis, drying
and persistent on the ripe fruit, rather slender, dilated above, bearing a few areoles
similar to those of the branches, the areoles subtended by small scales; perianth
limb somewhat shorter than the tube, widely expanded, the outer perianth seg-
ments narrowly lanceolate to linear, acuminate, green, shorter than the white inner
segments; stamens shorter than the perianth, inserted all along the upper half of
the throat; style very slender, divided at the apex into several linear stigma lobes;
fruit spiny or naked, with a thick, dark red rind, rupturing irregularly from the
apex downward, the flesh red, very juicy; seeds small, numerous, black.
Seven species are recognized by Britton and Rose, ranging from
Mexico to Colombia. Only the following are known from Central
Stems 8-10 cm. broad, deeply crenate; spines very stout A. horridus.
Stems 8 cm. broad or narrower, low-crenate; spines slender A. pentagonus.
Acanthocereus horridus Britt. & Rose, Cactaceae 2: 122, f. 181.
1920. Cereus horribilis Berger, Kakteen 124. 1929.
Based upon cultivated plants of Guatemalan origin, collected
by Federico Eichlam, the precise locality not known; Zacapa. Mexico
Plants stout, the stems strongly triangulate, the angles wing-like, deeply undu-
late; areoles large, 3-6 cm. apart; spines brown or blackish when young, the radials
1-6, conic, less than 1 cm. long; central spine usually 1, sometimes 2, often very
stout and elongate, sometimes 8 cm. long; flowers 18-20 cm. long, the throat 4 cm.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 191
broad; outer perianth segments linear, brown or greenish, 6 cm. long, the inner
segments 3-4 cm. long; stamens white; style thick, cream-colored; fruit 3.5 cm.
long, pale red, lustrous, covered with large areoles bearing white felt, the skin thick,
splitting as the fruit ripens, the pulp red.
Acanthocereus pentagonus (L.) Britt. & Rose, Contr. U. S.
Nat. Herb. 12: 432. 1909. Cactus pentagonus L. Sp. PI. 467. 1753.
Cereus pentagonus Haworth, Syn. PL Succ. 180. 1812. Pitajaya.
Moist or dry thickets or hedges, 1,200 meters or less, most
frequent at low elevations; Pete"n; Zacapa; Jutiapa; Santa Rosa;
Retalhuleu; San Marcos; probably in most of the lowland areas.
Southern Florida and Texas; eastern Mexico; southward to Panama,
mostly near the coasts. West Indies; northern South America.
Stems erect or arching, often scandent, sometimes 7 meters long but usually
much shorter, frequently forming dense colonies or thickets; sometimes epiphytic;
joints 3-8 cm. broad, mostly triangulate but often 4-5-angulate, the angles shal-
lowly crenate; areoles 3-5 cm. apart; spines gray, acicular or subulate, the radials
6-7 and 1-4 cm. long; central spine often solitary, longer than the radials; flowers
14-20 cm. long, the ovary and perianth tube bearing conspicuous areoles with
brown felt and several subulate spines; outer perianth segments green, the inner
ones white, acuminate; fruit oval, red, large, pulpy and juicy, edible, containing
very numerous small black seeds.
Called "saite" and "pitahaya" in El Salvador; "numtzutzuy"
(Yucatan, Maya). In Yucatan, and doubtless in other regions, the
long sharp spines are sometimes employed as a substitute for pins.
Backeberg (Die Cactaceae 2: 1933-1935. 1960) uses the name
A. tetragonus (L.) Hammelinck for the plants described here, offering
not very convincing reasons for so doing.
Plants slender, vine-like, creeping or clambering, emitting aerial roots, the
stems with mostly 7-12 ribs; flowers diurnal, pink or red, one at each areole, fun-
nelform, the tube of the perianth almost straight or bent above the ovary, the limb
somewhat oblique; outer perianth segments linear, spreading or recurved, the inner
segments broad; stamens exserted, arranged in a single, somewhat one-sided clus-
ter, inserted all along the throat; perianth tube about equaling the narrow throat;
fruit small, globose, reddish, setose; seeds few, reddish brown, obovoid.
Five species, all or most of them native in Mexico.
Aporocactus flagelliformis (L.) Lemaire, 111. Hort. 7: Misc. 68.
1860. Cactus flagelliformis L. Sp. PL 467. 1753. Cereus flagelliformis
Mill. Card. Diet. ed. 8, no. 12. 1768.
192 FIELDIANA: BOTANY, VOLUME 24
Grown commonly in Guatemala for ornament, in pots or hanging
baskets or often in the open ground, chiefly at low and middle
elevations. Perhaps native in Mexico, but unknown in a wild state.
Stems at first ascending or erect, in age weak, slender, often pendent, 1-2 cm.
in diameter, the branches often prostrate and creeping; ribs 10-12, low and incon-
spicuous, slightly tuberculate; areoles 6-8 mm. apart; radial spines 8-12, acicular,
reddish brown; central spines 3-4, brownish with yellow tips; flowers 7-8 cm. long,
opening for 3-4 days, deep rose-red; inner perianth segments broader than the
outer ones, only slightly spreading; fruit globose, 10-12 mm. in diameter, red,
bristly, with yellowish pulp.
This cactus is rather frequent in cultivation also in the United
States, where it is known by the name "rat-tail cactus." The
Maya name of Yucatan is reported as "canchoh," and the Spanish
name as "flor de latigo."
Plants terrestrial, large and tree-like, with columnar trunks, erect, sometimes
simple but usually with a few heavy branches; areoles at the ends of the stems
often developing a dense mass of white wool or a pseudocephalium, or the areoles
producing long woolly hairs but not forming a pseudocephalium; stems with few
or numerous ribs, armed with short or elongate spines; flowers nocturnal, short-
campanulate or short-funnelform, straight or curved; perianth persisting on the
ripening fruit; fruit usually depressed-globose or oblong; seeds numerous, small,
black, smooth or tuberculate.
About 50 species, in tropical and subtropical America. No other
species are known in Central America.
Cephalocereus Maxonii Rose, Contr. U. S. Nat. Herb. 12:
417, t. 64- 1909. Cereus Maxonii Vaupel, Monatsschr. Kakteenk.
23: 23. 1913. Pilocereus Maxonii Berger, Kakteen 345. 1929. Pilo-
socereus Maxonii Byles & Rowley, Cact. & Succ. Journ. Gr. Brit. 19:
3, 67. 1957. Tuno; cabeza de viejo; drgano.
Dry, rocky plains and hillsides, often planted for hedges, 200-
1,000 meters or even higher; endemic; Baja Verapaz; El Progreso
(type from El Rancho, Maxon 3769); Zacapa; Chiquimula; Jutiapa;
Plants 1-3 meters high or probably even taller, simple or with a few erect
branches, glaucous or bluish green, the apices of the branches covered with soft
wool-like hairs 4-5 cm. long; ribs of the stem 6-8, the areoles small; spines about
10 at each areole, slender, yellowish, the central one 4 cm. long or less, the areoles
bearing many long soft hairs; flowers white tinged with pink (fide Clover), 4 cm.
long; ovary naked except for a few small scales; fruit about 3.5 cm. broad and
almost as long; seeds brownish, reticulate.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 193
This species is easily recognized among the segregate "genera"
of Cereus in Guatemala by the covering of long hair-like bristles
borne near the end of the stems. The cactus grows widely scattered
in the lower Motagua Valley and also in the arid valley of Sacapulas,
where it is usually found in association with Lemaireocereus. The
plant is also to be found in the Comayagua Valley in Honduras and
perhaps in other dry, hot valleys of the north coast of Honduras.
This is perhaps the species reported from Guatemala, without local-
ity, by Hemsley as Cereus senilis Salm-Dyck. This species is placed
by Backeberg in a genus described in 1957, Pilosocereus Byles &
Rowley, along with 59 other species. To suggest that before 1957
there was not a generic name proposed that can be used for 60
species of cereoid cacti seems hardly plausible. We might even
suggest a look at the genus Cereus itself.
The ends of the stems of this cactus are often cut off and carried
to distant regions by peddlers, to be used as pot plants. Those
often seen in Quezaltenango perhaps come from Sacapulas or the
lowlands of Huehuetenango.
DEAMIA Britton & Rose
Plants terrestrial or usually epiphytic, sometimes clambering over rocks or
pendent from them, usually closely adherent to the bark of trees by aerial roots,
the stems broad, normally 3-angulate, the angles broad, thin, wing-like; spines of
the areoles numerous, acicular; flowers diurnal, very large, the tube slender, elon-
gate, the throat funnelform; inner perianth segments creamy white; stamens
numerous, attached all over the throat of the perianth; scales on the ovary and
perianth tube very small, bearing 3-5 long brown bristles in their axils; stigma
lobes linear, entire.
The genus consists of a single species and is dedicated to Charles
C. Beam, who made two important collections of plants in Guate-
mala, chiefly in the Motagua Valley.
Deamia testudo (Karw.) Britt. & Rose, Cactaceae 2: 213. 1920.
Cereus testudo Karw. in Zucc. Abh. Akad. Wiss. Muench. 2: 682.
1837. D. diabolica Clover, Bull. Torr. Bot. Club 65: 570, /. 5. 1938
(type from Corozal District, British Honduras, P. H. Gentle 490).
Mostly on trees, at or little above sea level; Izabal; Zacapa;
Retalhuleu; probably in all the Pacific coast departments. Southern
Mexico; British Honduras; Honduras, and ranging southward to
Colombia. Figure 30.
FIG. 30. Deamia iestudo. 1, Stem appressed to tree trunk; X 1 A- 2, Pendent
stem; X H- 3, Flower; X M- 4, Flower receptacle, longitudinal section; X Y^
5, Stigma; X 1. 6, Ovules and funicles, much enlarged. Courtesy of Myron
Kimnach. Drawn by Mrs. M. Bios.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 195
Plants usually closely attached to tree trunks or rocks by tough aerial roots,
often spirally twisting about the trunks, the stems 3-10 cm. broad or even larger;
ribs usually 3, very thin and wing-like; areoles 1-2 cm. apart; spines spreading,
10 or more at each areole, the larger ones 1-2 cm. long, brownish; flowers fragrant,
about 28 cm. long, the slender tube about 10 cm. long; inner perianth segments
linear-oblong, acuminate, 8-10 cm. long; scales of the ovary 1 mm. long or less;
hairs of the areoles on the ovary and perianth tube 1-3 cm. long, brown.
When growing, this plant is easily recognized because of its dis-
tinctive habit. The broad green stems, closely attached to the
trunk by stout aerial roots, are suggestive of a serpent coiled about
the tree. No distinctive characters are suggested for D. diabolica
by its author, and there is nothing in its description to indicate
how it may be separated from D. testudo. There is no reason to
assume that the genus, if it be treated as such, contains more than
a single species.
The genus Echinocactus probably occurs in Guatemala, and there
have been reported on apparently no good basis E. crispatus DC.,
E. gibbosus DC., and E. cornigerus DC. E. gibbosus is referred by
Britton and Rose to the genus Gymnocalycium, a group confined
to South America. E. crispatus is a species of Hidalgo, Mexico,
and there is no reason for supposing that it grows in Guatemala.
It is placed by Britton and Rose in the genus they treat under the
weird nameEchinofossulocactus. E. cornigerus they refer toFerocactus,
as a synonym of F, latispinus (Haw.) Britt. & Rose. We have ob-
served from the train near Progreso, Guatemala (Dept. El Progreso),
a small barrel cactus that may be the plant reported from the
country under this name. It is, however, possible that the plants
there taken to be Echinocactus were actually Melocactus Ruestii.
One or more species of Echinopsis, a South American genus,
sometimes are grown in Guatemala as pot plants. They are low,
subglobose or cylindric plants with numerous sharp ribs, armed
with stout spines, the flowers white, with a long slender tube.
196 FIELDIANA: BOTANY, VOLUME 24
EPIPHYLLUM [Hermann] Haworth
Plants mostly epiphytic, the main stems terete and ligneous, the secondary
stems or branches flat, fleshy, often thin and leaf-like; leaves none; flowers medium
to large, nocturnal or diurnal, fragrant or odorless; perianth divided into a tube
and a limb, the tube often greatly elongated; stamens few to numerous, inserted
at the top of the tube and also scattered along the throat of the tube or inserted at
the top and at the base, few to many; style elongated, the stigma lobes usually free,
linear and 4-10; fruit a berry, ovoid, sometimes tuberculate or ridged, juicy, pulpy
and edible; seeds few to numerous, black.
Species perhaps 50 or more (Epiphyllum, sens. lat.). Several
others occur in Mexico and Central America.
The generic concept as used here for Guatemala includes as syno-
nyms Bonifazia Standl. & Steyerm., Chiapasia Britt. & Rose, Diso-
cactus Lindl., Marniera Backeb. (now. subnud.}, Phyllocactus Link
and Trochilocactus Lindinger (now. nud.}. It does not include, how-
ever, the very small-flowered species sometimes placed under one or
another of these generic names.
Mr. Myron Kimnach has a monograph of Epiphyllum in prepa-
ration. He apparently is maintaining E. guatemalense Britt. & Rose
[here reduced to E. strictum (Lem.) Britt. & Rose] and E. Thomasi-
anum (Schum.) Britt. & Rose [here reduced to E. macropterum (Lem.)
Britt. & Rose]. In addition an inadequate specimen from Guatemala
(Steyermark 39545) he questionably determined as E. phyllanthus
var. columbiense (Webber) Backeberg. We have not included this
taxon in the flora.
In Guatemala the names pitahaya, pitajaya or pitaya are given to
most of the following species. The fruits when large and edible are
Tube of the perianth abruptly recurved near the base.
Limb of the perianth 2-3 times as long as the tube; stamens about 50-65.
Limb of the perianth shorter than the tube.
Ovary glabrous; stamens about 35-45 E. quezaltecum.
Ovary pubescent; stamens about 13-20 E. Eichlamii.
Tube of the perianth not recurved, straight.
Tube of the perianth shorter than the limb; stamens 15 or fewer. . .E. biforme.
Tube of the perianth much longer than the limb; stamens numerous.
Ultimate branches acuminate E. oxypetalum.
Ultimate branches rounded or acute at the apex.
Secondary stems deeply crenate; thick perianth tube bearing foliaceous
scales E. crenatum.
Secondary stems shallowly crenate to almost entire, often thin; perianth
tube without foliaceous scales.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 197
Crenations of the secondary stems oblique, the sinuses between them
very broad and open; secondary stems mostly 3-4.5 cm. broad.
Crenations of the secondary stems symmetrical, not oblique, the sinuses
between them very narrow, or the secondary stems entire or nearly
so and mostly 5-8 cm. broad E. macropterum.
Epiphyllum biforme (Lindl.) G. Don in Loudon, Encycl. PL
ed. 3. 1378. 1855. Cereus biformis Lindl. Bot. Reg. 29: Misc. 51. 1843.
Disocactus biformis Lindl. Bot. Reg. 31: t. 9. 1845; Britt. & Rose,
Cactaceae 4: 202, figs. 203, 204. 1923. Paxte de polo.
In mixed forests at about 1,300 meters; Sacatepe"quez (Donnell
Smith 2486). Honduras.
Pendent epiphytic herbs to 2 m. or perhaps longer; primary stems terete, ob-
scurely winged or not, 4-6 mm. in diameter; secondary stems or branches flattened,
to about 15 cm. long and 1.5 cm. broad, fleshy, narrowly elliptic, crenate or sinuate-
dentate, areoles glabrous or nearly so; flowers borne from the areoles, about 5 cm.
long, the limb about 1 cm. broad; tepals about 10, the outer ones somewhat shorter
and narrower than the inner, linear, acute; inner tepals broader; stamens 10-12,
slightly exserted; style exceeding the stamens; stigma lobes 4, about 4 mm. long;
berry ovoid, about 1.5 cm. long.
A sterile collection from the department of Quezaltenango (Steyer-
mark 33423) may belong to this species.
Epiphyllum crenatum (Lindl.) G. Don in Loudon, Encycl. PI.
ed. 3. 1378. 1855. Cereus crenalus Lindl. Bot. Reg. 30: t. 31. 1844.
Phyllocactus crenatus Lemaire, Hort. Univ. 6: 87. 1845. Pitaya;
pitajaya; huele de noche.
Moist or wet forest, sometimes in oak forest, 1,750 meters or less;
reported from Pete"n; Izabal; Alta Verapaz; Baja Verapaz; Zacapa;
Chiquimula; Jalapa; Sacatepe"quez ; Solola; Chimaltenango; Quiche";
Huehuetenango; doubtless in most of the central departments. Hon-
duras, and probably extending into Mexico and other parts of Cen-
Often a large vine, climbing over medium-sized trees, the main stems terete
and ligneous; branches pale green or glaucescent, stiff, often 8 cm. broad, obtuse,
deeply and coarsely crenate, the crenations usually oblique, the sinuses between
them broad or narrow; areoles at the base of the stem and branches sometimes
bearing hairs or small bristles; flowers fragrant, white, about 20 cm. long, cream-
colored or greenish white outside, the limb 10-12 cm. broad; perianth tube 10-
12 cm. long, slender, bearing numerous linear scales 2-3 cm. long, the inner seg-
ments oblanceolate; filaments yellow; style white.
This species, like all or most of the others, is grown commonly
for ornament in Guatemala, planted in the ground or in pots. Large
198 FIELDIANA: BOTANY, VOLUME 24
potted plants were covered with flowers in the Gran Hotel Conti-
nental in Guatemala in late April, 1941. The fruits of all the local
species are large, handsomely colored, and full of deliciously flavored
pulp. They are very good to eat, and much prized locally, being
sometimes offered in the markets. Plants of this species are common
in the central region, especially in oak forests, but the wild plants
seldom have flowers, at least during the dry months.
Epiphyllum Eichlamii (Weingart) L. Wms. Fieldiana, Bot. 29:
378. 1962. Phyllocactus Eichlamii Weingart, Monatsschr. Kakteenk.
21: 5. 1911. Disocactus Eichlamii Britt. & Rose, Contr. U. S. Nat.
Herb. 16: 259, t. 79. 1913; Cactaceae 4: 203, /. 205. 1923; Kimnach &
Hutchison, Cact. & Succ. Journ. Am. 29: 75, /. 45. 1957. Trochilo-
cactus Eichlamii Lindinger, Beih. Bot. Centralbl. 61: 383. 1942.
Epiphyte in mixed forest, 1,000-2,800 meters; Santa Rosa; Chi-
maltenango; Quezaltenango. Endemic. Figure 31.
Epiphytic herbs; stems erect or pendent, the primary terete, sometimes nar-
rowly 2-3-angled or winged, 4-8 mm. in diameter; secondary stems or branches
becoming flattened, up to 5 cm. broad (1.5-5 cm.) and 0.2-0.4 cm. thick, mostly
oblanceolate to narrowly elliptic-lanceolate, crenate or sinuate-dentate, the areoles
with wool or occasionally 1-2 bristles; flowers from the areoles of the secondary
stems, 6-8 cm. long, the limb to 1 cm. broad; tepals 10-12 in 2 series, the outer
linear or linear-lanceolate and about 2.5 cm. long, the inner broader and a bit
longer; stamens about 25, mostly inserted at two levels, exserted; style exceeding
the stamens; stigma lobes 5; ovary globose, somewhat floccose.
Probably one of the more common Epiphyllums in Guatemala.
Epiphyllum macropterum (Lemaire) Britt. & Rose, Cactaceae
4: 193, t. 17, f. 200. 1923. Phyllocactus macroptems Lemaire, 111.
Hort. 11: Misc. 73. 1864. "Marniera macroptera" Backeberg, Die
Cactaceae 2: 736. 1959. Pitahaya; galan de noche; dama de la noche.
On rocks or trees in moist or wet forest, 1,600 meters or less; Alta
Verapaz; Izabal; Chiquimula; Suchitepe'quez; Solola; Quezaltenango;
San Marcos; doubtless also in numerous other departments. Hon-
duras; Nicaragua; Costa Rica.
Plants often very large and scandent, the main stems terete and ligneous;
branches thin and flexible, deep green, sometimes 10 cm. broad, the margins some-
what corneous, usually rather closely crenate, the crenations short, symmetrically
rounded, low, separated by very narrow, acute sinuses, the apex of the branch
obtuse, the margins sometimes almost entire; flowers about 20 cm. long, somewhat
curved; scales of the ovary very small, green, with long hairs in their axils, the
scales of the perianth tube 10-12 mm. long, spreading, acute; outer perianth seg-
FIG. 31. Epiphyllum Eichlamii. 1, 2, Stems, buds and flowers; X 4 A- 3, Pro-
liferous areole; much enlarged. 4, Stamen insertion, with areole shown attached to
base of flower; X 3. 5, Stigma lobes; much enlarged. 6, Ovules; much
enlarged. 7, Fruit; X Vs. 8, 9, Seed; X 16. Courtesy of Myron Kimnach.
Drawn by Mrs. M. Bios.
200 FIELDIANA: BOTANY, VOLUME 24
ments narrow, salmon-colored, 10 cm. long, the inner segments pure white, 8-9 cm.
long, 2-3 cm. broad; stamens lemon-yellow.
This species is one of the commonest in cultivation in Guatemala.
The senior author observed during two seasons a fine large plant on
a trellis in the patio of dona Juana Reyes in Coban. It flowered pro-
fusely in mid-April, and in a period of 10 days produced more than
150 flowers, while innumerable buds remained on the plant. The
flowers are open only at night. It is a common practice in Guate-
mala to bind the tough, wide stems of Epiphyllum around fractures.
They act as efficient bandages, hold the broken parts in place, and
are popularly believed to aid in healing them.
This species is the basis of an illegitimate generic name, Marniera,
proposed by Backeberg. Since there seems to be no need for the name
it need not be properly published.
Most of the material which we have treated as Epiphyllum ma-
cropterum has been annotated recently as E. Thomasianum (Schum.)
Britt. & Rose by Kimnach. The reason is not obvious and we prefer
not to use the name, which we consider a synonym of E. macropterum,
until reason for the use is published.
Epiphyllum Nelsonii Britt. & Rose, Contr. U. S. Nat. Herb. 16:
257. 1913. Phyllocactus Nelsonii Vaupel, Monatsschr. Kakteenk. 23:
116. 1913. Chiapasia Nelsonii Britt. & Rose, Cactaceae 4: 203, t. 206.
1923. Disocactus Nelsonii Lindinger, Bot. Centralbl. Beih. 61: 383.
1942; Kimnach, Cact. & Succ. Journ. Am. 30: 80, t. 1958.
Epiphytic on trees in moist forest, 1,300-1,500 meters; San Mar-
cos. Mexico (Chiapas). Figure 32.
Pendent epiphytic plants; primary stems rounded, bearing flattened secondary
stems from nodes, the secondary stems oblanceolate, ascending but soon pendent;
flowers mostly from the apical part of the secondary stems, strongly upcurved on
pendent stems, less so on erect stems, 9-11 cm. long, slender, the limb 5-7 cm.
broad; tepals 10-13, in two series, 4-6 cm. long and 1-2 cm. broad, the outer linear-
lanceolate, acute, the inner oblong-lanceolate, acute or obtuse; stamens many
(50-65), mostly inserted in 2 zones, declined, exserted beyond the tepals; style
9-10 cm. long, exceeding the stamens, stigma lobes 4-7, papillose, about 8 mm.
long; fruit subglobose, about 15 mm. long and nearly as broad.
This species is the basis of Britton & Rose's monotypic genus
Chiapasia, which certainly has little to recommend it. Backeberg
recently maintained the genus (Die Cactaceae 2: 761. 1959).
Epiphyllum oxypetalum (DC.) Haworth, Phil. Mag. 6: 109.
1829. Cereus oxypetalus DC. Prodr. 3: 470. 1828. C. latifrons Pfeiff.
FIG. 32. Epiphyllum Nelsonii. 1, Flowering plant; about X J^. 2, 4 and 5,
Stem, flowers and fruit; about X 2 A- 3, Cross section of flattened stem; X Vs.
6, Longitudinal section of receptacle; X 2. 7, Stigma; X 2. 8, Funicles and
ovules; much enlarged. 9 and 10, Seeds; X 16. Courtesy of Myron Kimnach.
Drawn by Mrs. M. Bios.
202 FIELDIANA: BOTANY, VOLUME 24
Enum. Cact. 125. 1837. Phyllocactus latifrons Link ex Walp. Repert.
Bot. 2: 341. 1843. Paxte de palo.
On rocks or trees, 75-2,000 meters; Alta Verapaz; Suchitepe'quez;
Quezaltenango. Southern Mexico; El Salvador; Honduras; Costa
Rica; northern South America.
Plants often several meters long and scandent, sometimes sprawling on rocks,
the stems slender, ligneous, subterete; branches thin and flexible, long-acuminate,
shallowly or deeply crenate, 12 cm. broad or less; flowers opening in the evening,
fragrant, the tube 13-15 cm. long, reddish outside, bearing distant scales about
1 cm. long; outer perianth segments reddish or amber-colored, 8-10 cm. long, the
inner segments white, oblong; stamens white; style white, the stigma lobes numer-
ous, cream-colored, entire.
Epiphyllum quezaltecum (Standl. & Steyerm.) L. Wms. Fieldi-
ana, Bot. 29: 378. 1962. Bonifazia quezalteca Standl. & Steyerm.
Field Mus. Bot. 23 : 66. 1944. Disocactus quezaltecus Kimnach, Cact.
& Succ. Journ. Am. 31: 137, t. 1959.
Epiphytic in cool mixed forests, 1,800 meters, type from near San
Martin Chile Verde and Colombo, Quezaltenango. Endemic. Fig-
Pendent epiphytic herbs; primary stems round, 5-6 mm. in diameter, the
branches or secondary stems flattened, oblong-linear to ovate-lanceolate, 14-47 cm.
long and 4-5 cm. broad, fleshy when fresh, acute or acuminate, attenuate and sub-
terete to the base, crenate, the crenations 2-3 cm. long, somewhat oblique; areoles
small, minutely felted or pilose; flowers to 9 cm. long, near the apex of secondary
stems, strongly upcurved tube longer than to twice as long as the limb; tepals
10-12, the outer lanceolate, obtuse, 15-25 mm. long and 4-6 mm. broad, the inner
about 20 mm. long and 6-8 mm. broad; stamens about 35-45, inserted on the tube
at 2 levels, exserted, style exceeding the perianth, 5-6-lobed; fruit subglobose, to
about 20 mm. long.
This species is the basis of the genus Bonifazia Standl. & Steyerm.
and was dedicated to the family of don Guillermo Bonifaz, of Que-
zaltenango. Standley spent two months in their pension and has
many cherished memories of their gracious hospitality.
There seems little reason now to maintain the genus Bonifazia;
in fact it is very close to Epiphyllum Eichlamii (Weingart) L. Wms.,
which Standley and Steyermark considered then to be a synonym of
Disocactus biformis Lindl. [= Epiphyllum biforme (Lindl.) G. Don].
This plant was considered by the senior author, whose experience
in Guatemala has been exceeded by no other botanist, to be one of
the most attractive plants of all Guatemala. The flowers, though
small, are produced in great abundance, making the plant conspic-
uous from some distance, and they are of a lovely shade of rather pale
FIG. 33. Epiphyllum quezaltecum. 1, Habit; X Y%. 2, Secondary stems and
flowers; X 2 A. 3, Cross sections of secondary stems; X 2 A- 4, Proliferous areole;
X 4. 5, Flower; X 4 A- 6, Longitudinal section of receptacle; X 2>. 7, Stigma;
X 3. 8, Funicles and ovules; much enlarged. 9, Fruit; X 2 A- 10, 11, Seeds;
X 16. Courtesy of Myron Kimnach. Drawn by Mrs. M. Bios.
204 FIELDIANA: BOTANY, VOLUME 24
reddish purple. The plants hang loosely from the tree trunks, the
flowers being abruptly reflexed at the base and pointing upward.
Epiphyllum strictum (Lemaire) Britt. & Rose, Contr. U. S.
Nat. Herb. 16: 259. 1913. Phyllocactus strictus Lemaire, 111. Hort. 1:
Misc. 107. 1854. E. guatemalense Britt. & Rose, I.e. 257, t. 78 (type
collected in Guatemala by Federico Eichlam, the locality unknown).
E. pumilum Britt. & Rose, I.e. 258 (type collected in Guatemala by
Eichlam, the locality unknown). P. gicatemalensis Vaupel, Monatsschr.
Kakteenk. 23: 116. 1913. P. pumilus Vaupel, I.e. 117. Ticrebac
(Quecchi, fide Dieseldorff).
On trees in moist or wet forest, mostly at 600 meters or less;
Pete"n; Alta Verapaz; Izabal; Santa Rosa; Escuintla; Suchitepquez;
Solola; Retalhuleu. Southern Mexico; British Honduras to Pan-
Plants often 1-2 meters long or even larger, the stems slender, subterete or
angulate; branches broadly linear, mostly 3-4.5 cm. broad, sometimes broader,
rather thick and stiff, obtuse, remotely crenate, the crenations conspicuously
oblique; tube of the flower slender, 13-15 cm. long, the few distant scales 8-12 mm.
long; outer perianth segments pinkish, the inner ones white, acuminate, 6-8 cm.
long; filaments white, the style pink or red, the stigma lobes yellow; fruit globose,
4-5 cm. in diameter; seeds small, black.
Called "Santa Rita" in Yucatan. This has been reported from
British Honduras as E. oxypetalum (DC.) Haworth.
HELIOCEREUS Britton & Rose
Contributed by MYRON KiMNACH 1
Plants epiphytic or terrestrial, stems branching basally, ascending, procum-
bent, pendent or shortly scandent, slender, usually with 3-4 subacute angles, the
spines stiff or hair-like. Flowers at or near the apex of the stems, single at an
areole, large, remaining open several days and nights, funnelform; receptacle (peri-
anth tube) with small bracts subtending short wool and longer stiff or hair-like
spines, the tepals (perianth segments) half as long as the receptacle to three times
longer, lanceolate or oblong, red (white in one Mexican species) ; stamens and style
declinate, reddish; fruit globose, spiny, green; seeds large, ovoid-reniform, minutely
This is an easily recognized genus among Guatemalan cacti be-
cause of its showy, red, spiny flowers. It is allied to Nopalxochia,
which differs in its flat stems and nearly spineless receptacle; and to
Nyctocereus, which has white nocturnal flowers. Several variable and
rather indistinct species of the genus have been described from Mex-
1 Botanical Garden, University of California, Berkeley.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 205
ico, Guatemala, El Salvador, Honduras and Nicaragua, but their
evaluation must await further field work.
Heliocereus cinnabarinus (Eichl.) Britt. & Rose, Cactaceae 2:
129. 1920; Kimnach, Fieldiana, Bot. 29: 380. 1962. Cereus cin-
nabarinus Eichlam, Monatsschr. Kakteenk. 20: 161. 1910 (living
specimen collected by Eichlam from Volcan de Agua, a type speci-
men apparently not made). H. heterodoxus Standl. & Steyerm. Field
Mus. Bot. 23: 67. 1944 (type from Guatemala, Steyermark 36291).
Epiphytic or terrestrial in wet forests, 1,800-3,800 meters; Zaca-
pa; Chiquimula; Jalapa; Guatemala; Chimaltenango; Suchitepe"quez ;
Huehuetenango; Quezaltenango; San Marcos. Mexico (Chiapas);
Stems ascending when young, later usually pendent, 5-6-angled near the base,
3-4 (2-5) -angled above, to about 6 dm. long and 2 (1-8) cm. wide, the wings
usually 1-1.5 cm. wide, on 2-angled stems 2.5-4 cm. wide, more or less crenate,
green, the new growths often reddish, the areoles 5-10 mm. apart near base and
apex of stem, on remainder of stem (1-) 3-4 cm. apart, each with a brownish to
whitish wool-mass 1-4 mm. wide and near base with up to 15 brown to white,
acicular, stiff spines about 1-1.5 cm. long, the spines on the remainder of the stem
usually fewer, about 1 cm. long and more hair-like; flowers 12-16 cm. long, the limb
about 8 cm. wide, the receptacle green, 6.5-7.5 cm. long, about 1.5 cm. wide near
the base, 1 cm. wide at the middle, about 3 cm. wide at the apex, obscurely ridged,
the lower bracts deltoid, the upper ones lanceolate to oblong, acute to obtuse-
mucronate, 10-30 mm. long and to 4 mm. wide, each subtending a cream-colored
wool-mass about 2 mm. wide and with about 15 stiff or hair-like spines which are
5-10 mm. long, white or brown, and as long and as numerous near the apex of the
receptacle as at the base; tepals diverging gradually from the tube, lanceolate to
oblong, usually abruptly acuminate, often aristate, 6-9 cm. long and 1-2.5 cm.
wide, scarlet, often yellowish near the base.
HYLOCEREUS Britton & Rose
Plants terrestrial or epiphytic, often scandent or the stems arching, elongate,
mostly triangulate, the angles often wing-like, the branches often emitting aerial
roots; areoles bearing a tuft of felt and several short spines; areoles on young
growth often bearing bristles; flowers very large, nocturnal, funnelform, the limb
as broad as long and as long as the tube or longer; ovary and perianth tube bearing
large elongate narrow foliaceous scales but no spines, felt, wool, or hairs; outer
perianth segments similar to the scales but longer; petal oid perianth segments
narrow, acute or acuminate, usually white; stamens very numerous, biseriate,
equaling or shorter than the style; stigma lobes numerous, linear, simple or
branched; fruit spineless but bearing several or numerous large persistent folia-
ceous scales, usually large and edible, juicy; seeds very numerous, small, black.
206 FIELDIANA: BOTANY, VOLUME 24
Species perhaps as many as 10 (18 recorded by Britton and Rose;
21 by Backeberg), in Mexico, Central America, West Indies, and
northern South America. One or two others are known from south-
ern Central America.
Spines 5-12 mm. long H. Ocamponis.
Spines 2-4 mm. long H. undatus.
Hylocereus Ocamponis (Salm-Dyck) Britt. & Rose, Contr.
U. S. Nat. Herb. 12: 429. 1909. Cereus Ocamponis Salm-Dyck, Cact.
Hort. Dyck. 1849: 220. 1850.
Moist thickets, about 1,350 meters; Jalapa (near Jalapa, Standley
76424) . Believed to be a native of Mexico.
Stems deeply 3-4-angulate, at first bright green, becoming glaucous, in age
dull bluish green; ribs rather deeply undulate, the margins with a corneous border;
areoles 2-4 cm. apart, borne near the base of each undulation; spines 5-8, acicular,
5-12 mm. long; flowers 25-30 cm. long and fully as broad, the outer perianth seg-
ments narrow, long-acuminate, greenish, spreading or reflexed; inner perianth
segments white, oblong, acuminate; style stout, the stigma lobes linear, entire,
green; ovary covered with imbricate ovate acute purple-margined scales.
The single Guatemalan collection is sterile, but it is improbable
that flowers would supply any characters to substantiate further its
reference to H. Ocamponis, which was known to Britton and Rose
only from cultivated plants. Although the reference of the collection
to H. Ocamponis is somewhat doubtful, because of the long spines it
seems to represent a species distinct from H. undatus.
Hylocereus undatus (Haworth) Britt. & Rose in Britton, Fl.
Bermuda 256. 1918; Cactaceae 2: 187, t. SO, f. 263. 1920. Cereus
undatus Haworth, Phil. Mag. 7: 110. 1830. Cereus trigonus var.
guatemalensis Eichlam, Monatsschr. Kakteenk. 21: 68. 1911. Hylo-
cereus guatemalensis Britt. & Rose, Cactaceae 2: 184, /. 261. 1920.
Pitahaya; pitaya; pitajaya dulce.
Epiphytic or terrestrial in thickets, hedges, on rocks or rock walls,
at 2,000 meters or usually much less; Pete"n; El Progreso; Jalapa;
Zacapa; Jutiapa; Santa Rosa; Escuintla; Guatemala; Sacatepe"quez;
Retalhuleu; Solola; Quiche". Mexico, El Salvador and the West In-
dies to South America; often cultivated in other parts of the world.
Plants terrestrial or epiphytic, when terrestrial often with arching or recurved
stems, when epiphytic more or less scandent and emitting aerial roots; ribs of the
stem generally 3, broad, thin, green or glaucous green, the margins undulate,
FIG. 34. Hylocereus undatus. A, Flower and tip of stem; X M- B, Areole;
X 2. C, Cross section of stem; X M-
208 FIELDI AN A: BOTANY, VOLUME 24
corneous; areoles 2-4 cm. apart; the spines 1-4, usually subconic or more numer-
ous and slender on young plants; flowers as much as 30 cm. long, the outer seg-
ments whitish, yellowish green, or tinged with rose, acuminate, the inner segments
white, lanceolate to oblanceolate, acute or acuminate; style elongated, to 25 cm.
long, usually yellow or yellowish, lobes of the stigma to 25; fruit 6-12 cm. long,
usually deep red when mature, covered with large foliaceous scales; seeds small,
numerous and black.
The illustration and description given and specimens seen by
Britton and Rose of H. guatemalensis seem to show a plant that
differs hardly at all from the typical forms of H. undatus. Maya
names of Yucatan, according to Gaumer, are "chacuob," "zacuob,"
"uob," "uoo" and "uo." The usual Spanish name there, as elsewhere,
is "pitahaya." The ripe fruit is juicy, sweet or acidulous and deli-
cious. It is one of the most attractive fruits seen in Guatemalan
markets, due to its brilliant coloring. It is used as a fruit out of
hand, in the preparation of cool drinks and in coloring candy and
pastry. It is cultivated occasionally in Guatemalan gardens and is
common as a wild plant in the Pacific lowlands.
LEMAIREOCEREUS Britton & Rose
Plants terrestrial, usually large and tree-like, generally with few or numerous,
erect or ascending, columnar, ribbed stems, the trunk thick and massive; areoles
rather large, felted, the spines usually stout and numerous; flowers generally di-
urnal, one at an areole, tubular-funnelform or campanulate, the short tube tardily
separating with the style from the top of the ovary; stamens numerous, in many
rows all along the inner surface of the throat; ovary more or less tuber culate,
bracteate, the bracts with tufts of felt-like hairs in the axils; areoles at first spine-
less or nearly so but soon developing a cluster of spines; fruit globose or oval, irreg-
ularly bursting in age and exposing the seeds, at first very spiny, but the spines
often deciduous in age, or at least easily separable; seeds numerous, very small,
About twenty species, ranging from Arizona to the northern coast
of South America. There are other species in Mexico and Central
Areoles conspicuously brown-felted; central spine stouter and usually much longer
than the others, normally more than 2 cm. long. . . Lemaireocereus Eichlamii.
Areoles not conspicuously brown-felted and much smaller; central spine relatively
slender and hardly longer than the others, mostly less than 1.5 cm. long.
Lemaireocereus Eichlamii Britt. & Rose, Cactaceae 2: 89,
/. 182. 1920. Cereus laevigatus var. guatemalensis Eichlam in Wein-
gart, Monatsschr. Kakteenk. 22: 182. 1912 (type from Guatemala).
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA! jj 209
L. longispinus Britt. & Rose, I.e. Cereus Eichlamii Standl. in Yuncker,
Field Mus. Bot. 9: 316 1940. Ritterocereus Eichlamii Backeberg,
Cact. & Succ. Journ. Am. 23: 121. 1951. Tuna; organo; guanocal
Dry, rocky scrub-forest plains and hillsides, 200-1,250 meters;
Baja Verapaz; El Progreso; Zacapa; Chiquimula; Baja Verapaz;
Santa Rosa; Quiche"; Huehuetenango. El Salvador; probably also
Branched organ cacti up to 6 m. or more, with a thick low trunk, usually with
many thick erect or ascending branches, deep green or the younger parts somewhat
glaucous; ribs 6-10; the areoles large, brown-felted, borne on the tops of the undu-
lations; spines 4-10 at each areole, spreading or reflexed, acicular, the central spine
much longer than the others, often to 8 cm. or more, gray; flowers 6-7 cm. long,
white inside, dark red outside, outer perianth segments serrulate; filaments white;
ovary tuberculate, each tubercle bracteolate; areoles of the ovary with brown felt
but no spines.
This is apparently the abundant and characteristic plant of the
plains of the lower Motagua Valley, frequently in association with
other organ cacti. The plants are much used for hedges, which are
one of the typical sights of many parts of Mexico and Central Amer-
ica. The branches are separated from the wild plants and set side
by side. Their planting involves a great deal of labor, but once set
the hedges are good for many years. The pulp of the fruit is very
good to eat. The fruits (called tunas*) ripen in March and April or
even later, and at that season they are a common article in some of
Cereus Yunckeri Standl. in Yuncker, Field Mus. Bot. 9: 316,
/. 7. 1940; L Wms. Fieldiana, Bot. 29: 384. 1962.
Dry, hot valleys in scrub forest, 600-900 meters or perhaps less;
Huehuetenango (Steyermark 51341). Honduras. Organo.
Much-branched organ cacti up to 10 m. tall, the trunks as much as 35 cm. in
diameter toward the base, the branches erect or ascending, thick, the broadly tri-
angular costae (on available specimens) 9, intervals broad but acute; areoles
8-15 mm. apart on the ridges, tomentum brownish, obscure or almost none; spines
mostly about 7 at each areole, the central one usually longest, from about 3 to
15 mm. long, grayish. Flowers and fruits unknown.
Among the many "microgenera" currently accepted by cactus
specialists this might possibly go to Pilosocereus. It was probably
placed by Standley in the correct genus and until we know more
about the plant it seems unwise to make a combination to one of the
210 FIELDIANA: BOTANY, VOLUME 24
Plants small, globose or short-cylindric, tuberculate, the tubercles arranged in
somewhat spiral rows, terete, angulate, or compressed, usually bearing wool or
hairs and sometimes bristles, the sap milky or watery; spines in clusters at the apex
of the tubercle, sometimes all alike, or the central ones very different from the
radials, all straight or some of the central spines uncinate; flowers, so far as known,
diurnal, arising from the axils of old tubercles, more or less campanulate, small,
variously colored but mostly pink or red; perianth segments narrow, spreading;
stamens numerous, inserted on the base of the perianth tube, short, included;
style about equaling the stamens, the stigma lobes linear; fruit small, usually
clavate, naked, scarlet, rarely white or greenish; seeds brown or black.
Britton and Rose recognize 150 species, as Neomammillaria, rang-
ing from western United States to Nicaragua. Most of them are
Tubercles not emitting milk when cut, the milk tubes, if any, only in the stem.
Tubercles emitting milk freely when cut.
Flowers yellow M . woburnensis.
Flowers pink or red, or with dark red stripes.
Wool in the axils of the tubercles yellow M. Eichlamii,
Wool in the axils of the tubercles white M. Praelii.
Mammillaria Eichlamii Quehl, Monatsschr. Kakteenk. 18: 65.
1908. Neomammillaria Eichlamii Britt. & Rose, Cactaceae 4: 94,
/. 91. 1923. Chile.
Dry plains and hillsides, 300-1,600 meters, El Progreso; Zacapa;
Baja Verapaz; described from Sabanetas (possibly Dept. Guate-
mala), the material collected by Federico Eichlam. Honduras.
Plants solitary or caespitose, the plant body globose or short-cylindric, 15 cm.
long or less, yellowish green, the tubercles only slightly angulate, with copious
milky sap; axils filled with dense, yellowish or whitish wool and longer white
bristles; radial spines 7-8, ascending, white with short brown tips; central spines
usually 1, sometimes 2, stouter; flower buds covered with long wool; outer perianth
segments narrow, acuminate, with a dark red stripe down the middle, otherwise
cream-colored, the inner segments acuminate, cream-colored or light lemon-yellow;
style longer than the stamens; stigma lobes 4-6, yellow.
The local Mammillarias are common pot plants in Guatemala,
especially in the cooler regions, and frequently are offered for sale
Mammillaria Praelii Muehlenpf. Allg. Gartenz. 14: 372. 1846.
M. viridis var. Praelii Salm-Dyck, Cact. Hort. Dyck. 1849: 16.
1850. M. viridis Salm-Dyck, I.e. 116. 1850. M. inclinis Lemaire,
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 211
111. Hort. 5: Misc. 9. 1858. Cactus Praelii Kuntze, Rev. Gen. 261.
1891. C. viridis Kuntze, I.e. Pinuela; toniboc; huevos de coyote.
Based on plants of Guatemalan origin; we refer here with much
doubt several collections from exposed limestone rocks, Huehuete-
nango, at 800-2,500 meters.
Plants globose or short-cylindric, light green, depressed at the apex, densely
spiny, the axils of the tubercles lanate and setose; tubercles somewhat tetragonous;
areoles villous; radial spines 4, forming a cross, the uppermost and lowermost
much elongated; flowers rose-red or red-purple.
Mammillaria Ruestii Quehl, Monatsschr. Kakteenk. 15: 173.
1905. M. Celsiana var. guatemalensis Eichlam, Monatsschr. Kak-
teenk. 19: 59. 1909 (type collected near Guatemala by Federico
Eichlam). Corazdn de piedra.
Exposed rocky places, 700-1,600 meters; Chiquimula; Jalapa;
Guatemala. Described from Honduras; Nicaragua.
Plants mostly short-cylindric, as much as 20 cm. high and 4-6 cm. in diam-
eter, light green, almost hidden by the dense spines; axils of the tubercles lanate,
at least when young; radial spines 20 or more, white, 5-6 mm. long, spreading, the
central spines usually 4, sometimes 5, much stouter than the radials, pale yellow,
7-8 mm. long; flowers often almost hidden by the spines, 8 mm. long; inner peri-
anth segments acute, pale purple or rose-red; stigma lobes 4, linear, elongate; fruit
clavate, red, the seeds brown.
Mammillaria woburnensis Scheer, Lond. Journ. Bot. 4: 136.
1845. Cactus woburnensis Kuntze, Rev. Gen. 261. 1891. M. chapi-
nensis Eichlam & Quehl, Monatsschr. Kakteenk. 19: 1. 1909. Neo-
mammillaria woburnensis Britt. & Rose, Cactaceae 4: 100. 1923.
Dry, rocky, open slopes, 200-1,250 meters; El Progreso (based at
least in part on material collected at El Rancho by Eichlam); Za-
capa; Chiquimula; Jutiapa; Quiche 1 ; the species was described from
material of Guatemalan origin grown in England.
Stems usually several and forming small clumps, globose or cylindric, dull
green, giving off copious sap when cut; tubercles angulate, lanate in their axils;
radial spines 5-9, yellowish or white; central spines 1-8, often elongate, reddish
or yellow; flowers yellow, about 1 cm. long; fruit red, clavate, 2.5 cm. long or
shorter; seeds minute, brown.
MELOCACTUS Link & Otto
Plants solitary or cespitose, globose or short-cylindric, conspicuously many-
costate, bearing clusters of spines on the ribs; inflorescence a compact cap-like
mass of hairs and bristles, these forming a cephalium borne at the top of the plant,
212 FIELDIANA: BOTANY, VOLUME 24
this often large and sometimes elongate; flowers small, pinkish, arising from the top
of the cephalium, tubular-sal verform, the perianth segments few, spreading; sta-
mens inserted near the apex of the slender perianth tube; style slender, the stigma
lobes linear, few; fruit clavate, naked, usually red; seeds small, black.
The genus occurs in Mexico, Central America and the West In-
dies, extending to northern South America. No other species in
Melocactus Ruestii Schumann, Verzeichn. Kult. Kakt. 26.
1896. Cactus Maxonii Rose, Smithson. Misc. Coll. 50: 63. 1907.
Melocactus guatemalensis Giirke, Monatsschr. Kakteenk. 18: 93. 1908.
M. Maxonii Giirke, I.e. 93. Cactus Ruestii Britt. & Rose, Cactaceae
Dry, rocky, open or brushy plains or hills, 200-700 meters; Baja
Verapaz; El Progreso and doubtless other departments. Honduras.
Plants depressed-globose, 10-15 cm. high or larger, dull blue-green, the ribs
11-15, broad-based; radial spines 7-11, spreading or recurved, pale red or rose-
colored, 1-1.5 cm. long, stout and hard; central spine usually solitary, porrect or
ascending; cephalium small; flowers small, deep rose-red, opening in the afternoon;
fruit deep rose-red, clavate, the small seeds black and lustrous.
The plant is widely distributed in the lower Motagua Valley but
is of rather sparse occurrence. Plants often are grown in pots in
Guatemala, and we have seen them offered for sale by vendors
around the Parque Central in Guatemala City. The fruit is sweet
The types of Melocactus Maxonii and M. guatemalensis were both
collected in Guatemala; that of M. Ruestii in Honduras. The limited
material which is available, in addition to field observation, indicates
that the three names represent only a single species.
Called "cabeza de viejo" and "barba de viejo" in Honduras.
Plants large, usually with a well-defined trunk and few or numerous, erect or
ascending, thick, ribbed branches, all the areoles bearing uniform spine clusters;
flowers diurnal, small, several at an areole, sometimes as many as 9, with a very
short perianth tube, the segments widely spreading; ovary bearing a few minute
scales with tufts of wool in their axils, spineless; fruit small, globose, fleshy and
edible; seeds very small, black.
Four species are described; the others are Mexican.
Myrtillocactus Eichlamii Britt. & Rose, Cactaceae 2: 180,
/. 256. 1920.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 213
Based upon material collected by Federico Eichlam in 1909 in
Guatemala, the locality unknown, but possibly the lower Motagua
Stems strongly 6-angulate, deep green or slightly glaucous, the ribs obtuse;
areoles 2 cm. apart, large, circular, with grayish wool at an thesis; radial spines 5,
bulbose at the base; central spine 1, slightly longer than the radials; flower buds
dark purple, the outer perianth segments greenish, with red tips; inner perianth
segments about 10, creamy white, spreading at almost a right angle to the tube;
stamens numerous, the style white, slightly longer than the stamens; flowers
fragrant; fruit 6 mm. in diameter, globose, wine-colored, naked except for a few
This species was sent by Eichlam to Britton and Rose, who de-
scribed it, and so far as we know, it has not been seen or collected by
any botanist in the half century since that time. The description
above is taken from the original.
Plants shrub-like, much-branched, generally with definite short cylindric
trunks; branches (joints) compressed, thick and succulent, broad or rather nar-
row; glochids present in the areoles, abundant or few; spines solitary or clustered
in the areoles, not vaginate, large or small; leaves small, subterete, early deciduous;
areoles bearing white wool, glochids, and often spines; flowers arising in the areoles,
mostly at or near the edges of the joints; sepals ovate, erect; petals red or pink,
erect, appressed against the stamens and style; filaments and style slender, much
exceeding the petals; ovary somewhat tuberculate, naked or armed with spines,
with a very deep umbilicus; fruit a juicy berry, red, edible, usually without spines;
seeds numerous, flat, covered by a hard osseous aril.
About 7 species, in Mexico and Guatemala; possibly native far-
ther southward but probably only naturalized there.
Joints spineless or nearly so, sometimes with a few scattered, very short spines.
Joints armed with numerous elongate spines.
Joints broadly obovate N. guatemalensis.
Joints oblong or narrowly oblong.
Areoles 2-2.5 cm. apart N. lulea.
Areoles mostly 3-4 cm. apart or more remote N. dejecta.
Nopalea cochenillifera (L.) Salm-Dyck, Cact. Hort. Dyck.
1849: 64. 1850. Cactus cochenillifer L. Sp. PI. 468. 1753. Tuna;
tunal; chuh (Poconchi) ; tuno; tuno de Castilla.
Probably not native in Guatemala, but much planted at low and
middle elevations; thoroughly naturalized in many places at middle
elevations, up to 1,500 meters or even higher, perhaps the relics of
214 FIELDIANA: BOTANY, VOLUME 24
former commercial plantations; often planted for hedges; Alta Vera-
paz; Baja Verapaz; Izabal; Zacapa; Jutiapa; Guatemala; Sacatepe"-
quez; Chimaltenango; Suchitepe'quez ; Retalhuleu; Quezaltenango;
Huehuetenango; probably to be found in all departments. Mexico;
planted for ornament commonly in many parts of tropical America.
Plants shrub-like, commonly 2-4 meters high, the trunk sometimes 20 cm. in
diameter; joints oblong-obovate, sometimes 50 cm. long, green or at first bright
green; spines none, or very small ones sometimes developing on old joints; glochids
numerous, caducous; leaves very small, subulate, early deciduous; flowers arising
at the tops of the joints, usually numerous, about 5 cm. long; ovary subglobose,
2 cm. long, tuberculate and bearing many glochids; sepals broadly ovate, bright
red, acute; petals similar to the sepals but longer, erect; stamens pink, much ex-
serted; stigma lobes 6-7, greenish, longer than the stamens; fruit red, about 5 cm.
long; seeds hard, 5 mm. long.
The Maya name in Yucatan is "pacam." The Nahuatl name,
"nopal," is applied to the plant in some parts of Central America
and in Mexico but is little used in Guatemala. There the name
"tuna," given in Mexico and elsewhere to the fruit, is applied to the
plant, which is often called "tuno" or "tunal," the names used com-
monly for all the Opuntia and Nopalea species. This species has been
of great economic importance in the past, as the host of the cochineal
insect, from which a handsome dye was obtained. The plants were
cultivated on a large scale in Mexico, but the principal source of the
dye was the Canary Islands, which in 1868 produced more than
6,000,000 pounds, valued at four million dollars. In Guatemala co-
chineal was grown on a smaller scale, but it is said that great areas
of land about Antigua and Amatitlan were devoted to it. In the early
1880's neglected nopaleras were still to be seen about Antigua, and
in 1883 as much as 184 cwt. were exported from Guatemala. The
dye was much used for coloring the local textiles, and some may be
produced at the present time about Zacapa, Salama and Amatitlan.
At many places in the central region there are extensive thickets of
Nopalea on the drier hills, and they probably are the remains of
former cultivated fields. The plants are grown commonly in many
places for hedges. Cochineal dye was in use by the original inhabi-
tants of Mexico and Central America, and immediately after the
Conquest it was exploited by the Spaniards. It was long one of the
chief articles of tribute to the crown. Today it has been almost
wholly displaced by synthetic dyes. The cochineal insects were
"planted" upon the branches of the plants, where they multiplied
rapidly. When mature, they were brushed off into bags, then dried,
and in this form they were exported. They are very small and it is
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 215
almost incredible that millions of pounds could have been collected
in a single year.
Nopalea dejecta Salm-Dyck, Cact. Hort. Dyck. 1849: 64. 1850.
Opuntia dejecta Salm-Dyck, Hort. Dyck. 361. 1834. Tuna.
Moist or dry thickets, often in coastal thickets, mostly on the
Pacific coastal plain, 1,350 meters or less; Santa Rosa; Retalhuleu;
Huehuetenango (Cuilco); probably occurring in all or most of the
Pacific departments. Southern Mexico; El Salvador.
Plants erect, rather sparsely branched, shrub-like, or sometimes a small tree
of 5 meters with a definite trunk, strict, bright green; spines numerous, sometimes
4 cm. long, slender, at first pale yellow or pinkish, in age gray; joints narrow, 10-
20 cm. long and 3.5-5 cm. broad or sometimes longer; flowers about 5 cm. long, the
sepals obtuse; petals erect, dark red; stamens dark red, long-exserted.
The fruit, as in other species, is edible, but of inferior quality.
This plant is found wild only, and almost always in medium or dense
shade under low trees. It is common about Champerico.
Nopalea guatemalensis Rose, Smithson. Misc. Coll. 50: 330,
tt. ?, 42. 1907. Tuna.
Dry, rocky plains and hillsides, 400-900 meters; Zacapa; El Pro-
greso (type from El Rancho, Maxon 3774). Possibly also in Hon-
Plants tree-like, 5-7 meters high, or when young low and shrub-like, much-
branched, with a dense crown and a definite thick trunk; joints bluish green, obo-
vate to broadly oblong, 15-20 cm. long; areoles numerous, filled with white wool;
spines 5-8 at each areole, or often very numerous, most of them soft, hair-like, and
flexuous, nearly or quite porrect, unequal, white or sometimes pink, the longest
2.5-3 cm. long; leaves small, linear, reflexed; flower 5-6 cm. long; sepals ovate, the
petals red or rose; fruit 4-5 cm. long, clavate, red, somewhat tuberculate, deeply
umbilicate at the apex, without conspicuous glochids; seeds 4 mm. broad.
The characters by which this and some of the Mexican species
are separated in the key of Britton and Rose's The Cactaceae do not
appear very convincing.
Nopalea lutea Rose, Contr. U. S. Nat. Herb. 12: 405, t. 58. 1909.
On sandbars or rocky hillsides, 300-1,100 meters; El Progreso
(type collected near El Rancho, Kellerman 7046) ; Chiquimula; Juti-
apa. Honduras; Nicaragua.
Plants tree-like, 5 meters high or less, with a short distinct trunk and several
large branches; joints oblong to obovate-oblong, 10-22 cm. long, pale green and
216 FIELDIANA: BOTANY, VOLUME 24
slightly glaucous; areoles 2-2.5 cm. apart, filled with short brown wool; spines
weak, yellow, acicular or bristle-like, the longest 4 cm. long; flowers 5 cm. long, the
petals red, 2 cm. long; ovary with numerous prominent areoles filled with yellow
bristles; fruit red, 4 cm. long; seeds 4-5 mm. in diameter.
NYCTOCEREUS Britton & Rose
Plants erect, clambering, or procumbent, rather slender, sparsely branched or
simple, the stems cylindric, ribbed, the ribs rather numerous and low; areoles each
bearing a tuft of short white wool and small radiating bristles or weak spines;
flowers large, white, nocturnal; ovary bearing small scales, short or long wool, and
clusters of weak spines or bristles; perianth funnelform, bearing scales and tufts
of weak bristles below the middle, above the middle bearing distant, narrowly
lanceolate scales; inner perianth segments widely spreading, obtuse or subacute;
stamens numerous, shorter than the perianth; style about equaling the stamens;
fruit fleshy, scaly, with tufts of spines or bristles; seeds large, black.
Five species are recognized, based upon characters that do not
appear convincing. Two other Central American ones are reported,
both from Nicaragua.
Nyctocereus guatemalensis Britt. & Rose, Contr. U. S. Nat.
Herb. 16: 240, it. 70, 71. 1913. Cereus guatemalensis Vaupel, Mo-
natsschr. Kakteenk. 23: 86. 1913.
Dry, brushy plains, 200-300 meters; endemic; El Progreso (type
from El Rancho, Maxon 8510) ; Zacapa.
Stems erect, subscandent, or recurved and arching, sometimes prostrate, 1-2
meters long or more, 3-6 cm. in diameter, very densely spiny; ribs 8-12, low;
radial spines about 10; central spines 3-6, usually much longer than the radials,
the longest ones 3-4 cm. long; flowers fragrant, white, 4-7 cm. long; ovary some-
what tuberculate, each tubercle tipped by an areole bearing a cluster of pinkish or
brownish spines; outer perianth segments brownish, the inner ones lanceolate,
acute; stamens much shorter than the perianth, attached all along the surface of
the wide throat; style stout, 3 cm. long; fruit 2 cm. long, spiny; seeds black, lus-
trous, 3 mm. in diameter.
A common plant on plains about Zacapa and elsewhere in the
lower Motagua Valley. This species may prove to be the same as
N. Hirschtianus (Schum.) Britt. & Rose, described from Nicaragua.
OPUNTIA Miller. Prickly pear
Plants low and branched from the base or often shrub-like or tree-like with
definite trunks, branched, the branches (joints) usually compressed and flattened,
succulent, sometimes cylindric or globose, often with a woody skeleton; areoles
axillary, bearing spines, barbate bristles (glochids), hairs, flowers, and sometimes
glands; leaves small, terete, caducous; spines solitary or fasciculate, terete or com-
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 217
pressed, vaginate or naked, variously colored; glochids usually numerous, borne
above the spines; flowers mostly 1 at an areole; ovary inferior, 1-celled, many-
ovulate, bearing scale-like leaves, the areoles often with spines and glochids; sepals
green or colored, grading into the petals; petals mostly red or yellow, sometimes
green, spreading; stamens much shorter than the petals, sensitive; style thick, the
stigma lobes short; fruit baccate, dry or juicy, often edible, spiny or naked; seeds
covered with a hard osseous aril, whitish, compressed; embryo curved; cotyle-
dons 2, large.
Species about 240, all American and most numerous in Mexico,
but distributed from southern Canada to Patagonia. Probably a
few species besides those listed here are found in other parts of Cen-
tral America, but the plants are not plentiful in southern Central
America except in restricted areas. This is the largest genus of the
Cactaceae, but many of the species are poorly understood and sep-
arated by no very definite characters, and the real number of species
is decidedly uncertain. It is evident from their key to the groups
and species that Britton and Rose had no very clear idea of the rela-
tionships of many of the species, and perhaps the genus can never
be divided clearly into groups and specific units. Backeberg, Die
Cactaceae 1: 119-628. 1958, gives more than 350 species belonging
to Opuntia and the segregate opuntioid genera. The work is not so
useful as that of Britton and Rose.
Some of the species of Opuntia recorded from Guatemala are none
too well marked. While all Opuntias are native to the Americas,
some were introduced into Spain and other parts of the Mediter-
ranean region soon after the time of discovery. Some have become
thoroughly naturalized there and are now as much a part of the
landscape as they are in Mexico and Guatemala. Opuntias were in-
troduced into Australia and soon became a terrible pest there, ruin-
ing large areas of cultivated land. Biological control in recent years
has reduced the number of plants in Australia almost to the vanish-
ing point. In their native homes Opuntias usually are not trouble-
some. While abundant in many areas, they can be destroyed by
cutting and burning, and show little tendency to spread. Further-
more, in some areas, mostly in Mexico, they are of great economic
importance. The plants are, of course, very offensive where they
are abundant, sometimes forming impenetrable thickets, for the
longer spines can inflict severe wounds that heal slowly. If the spe-
cies is one in which the spines are encased in a loose sheath, this
remains in the flesh when the spine is removed, often causing fester-
ing wounds. More troublesome than the spines are the innumerable
short bristle-like glochids, which adhere in great numbers to the skin
218 FIELDIANA: BOTANY, VOLUME 24
if a plant is brushed carelessly. In some regions of the southwestern
United States the plants have been found to be good forage for stock
in time of drought. The spines may be burned off with a blow torch,
but if cattle are starving for lack of grass, they will not hesitate to
eat the joints, spines and all. In Mexico the young, tender joints are
peeled to remove all the spines and glochids, then cooked and eaten
as a vegetable. They may be eaten thus in Guatemala, but we have
not observed any use made of them. Very often the plants are grown
in Guatemala as hedges, a purpose for which they are fairly effective,
although stock will push through them. There are many miles of
Opuntia hedges in the Occidente, especially in the highlands about
Quezaltenango. Sometimes the "tunos" are planted along the tops
of adobe walls, thus keeping out at least human marauders, an effect
sometimes obtained equally well by setting broken bottles or other
glassware along the walls.
In parts of Mexico tunas are an important food for man at their
harvest time, constituting for some weeks a large part of the diet.
The best fruits are large, handsomely colored, and full of richly col-
ored, red juice of agreeable flavor, with rather abundant pulp. The
glochids and spines can be removed with a brush when the fruit is
ripe. There always is some danger of getting glochids in the mouth,
but persons habituated to eating the fruit seem to pay little attention
to them. The pulp and juice are used in Mexico commonly for color-
ing food, also for preparing sirups and a sort of marmalade called
"queso de tuna," which may be bought at almost any season of the
year. Tunas are little eaten in Guatemala, principally because fruit
of most local plants is inferior in quality. In many wild species the
fruit is almost dry and in no way edible. The seeds are numerous
and large, hard and quite indigestible, and if eaten with the pulp,
as they often are, they cause constipation. It is well known that
some Indians of southern California and Baja California, at least in
times when food was scarce, collected the hard seeds after they had
passed through the body, ground them into a coarse meal, and re-
used them as food. There may well be in Guatemala some species
of Opuntia not listed here, particularly among the cultivated plants,
whether grown for ornament or for their fruit. Many of the culti-
vated plants are spineless or nearly so, and such a form is not enu-
merated on the following pages. They may be 0. Ficus-indica (L.)
Mill., a species whose native habitat is unknown, but which is found
in cultivation in many parts of tropical America. The term "nopal"
is of Nahuatl origin, but the word "tuna" is said to come from the
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 219
Antilles. Two Indian names are recorded for the plants from Guate-
mala, "nuxtil" (Baja Verapaz) and "noxtie" (Quiche 1 ); the Quecchi
name is "tun," a corruption of the Mexican "tuna."
Joints of the stems subterete, small, 3-7 cm. long; flowers lemon-yellow, drying
red O. pubescens.
Joints strongly compressed, flat.
Joints puberulent or pubescent.
Plants low and spreading, mostly 50 cm. high or less; joints small, mostly
5-7 cm. broad O. decumbens.
Plants large, 1-2 meters high; joints 9-16 cm. broad.
Joints rounded obovate O. Guilanchi.
Joints mostly oblong or obovate-oblong O. tomentella.
Spines very slender and usually short, mostly about 1 cm. long, sometimes
2 cm. long 0. guatemalensis.
Spines mostly stout or very stout, chiefly 2-5 cm. long or even larger.
Large spines generally only 1-2 at an areole, relatively slender, mostly 3 cm.
long or shorter O. Eichlamii.
Large spines generally 3 or more at a node, stout, mostly 3-6 cm. long.
Opuntia Deamii Rose, Contr. U. S. Nat. Herb. 13: 309, t. 65.
1911; Britt. & Rose, Cactaceae 1: 187, /. 229. 1919.
On open rocky hillsides or in moist or dry, oak forest, 1,000-
1,700 meters; endemic; Baja Verapaz; Jalapa; El Progreso; Guate-
mala (type from Fiscal, Deam 6228) .
Plants erect, about a meter high, sparsely branched, green or bright green,
glabrous, often or usually with a very short, cylindric trunk; joints mostly very
large, obovate or oblong-obovate, 25-35 cm. long, as much as 20 cm. broad; are-
oles rather remote, commonly 4-5 cm. apart, relatively small; spines 2-6, generally
4, white or dull yellow, stout, somewhat compressed, spreading or porrect, 3-6 cm.
long; flowers about 7 cm. long, reddish; fruit oblong, 6 cm. long, naked except for
a few spines near the apex, wine-red inside and outside; seeds small, 3 mm. broad.
Opuntia decumbens Salm-Dyck, Hort. Dyck. 361. 1834. Tuno;
lengua de vaca; arpdn.
On dry, brushy, often rocky plains or hillsides, sometimes in oak
forest, 200-700 meters; El Progreso; Zacapa; Chiquimula; Jalapa;
Jutiapa. Western and southern Mexico.
Plants erect or usually spreading, 50 cm. high or less; joints few or numerous,
10-20 cm. long, obovate or rounded-obovate, finely, softly, and inconspicuously
pubescent, grass-green; areoles small or rather large, bearing yellowish wool
and very numerous yellow glochids; spines sometimes wanting, usually solitary,
sometimes numerous, slender or rather stout, 5.5 cm. long or shorter, yellow or
220 FIELDIANA: BOTANY, VOLUME 24
gray; flowers small, about 4 cm. long; petals deep yellow; fruit deep purple, very
juicy; seeds 4 mm. broad.
A common plant at many localities in the dry Oriente, sometimes
forming large colonies among shrubbery or in open places.
Opuntia Eichlamii Rose, Contr. U. S. Nat. Herb. 13: 310, t. 66.
1911; Britt. & Rose, Cactaceae 1: 187, /. 230. 1919. Tuna; tuno.
Chiefly on dry brushy hillsides or in dry or moist thickets, often
planted to form hedges, 250-2,200 meters; endemic; El Progreso;
Zacapa; Jutiapa; Guatemala (type collected near Guatemala, on the
road to Mixco, Federico Eichlam); Huehuetenango; Quezaltenango.
Plants tree-like when fully grown and often 5 meters high or more, many
of the plants, however, only a meter high or even less, glabrous, the large branches
few; joints often very numerous, 15-20 cm. long or frequently smaller, broadly
obovate to orbicular, somewhat glaucous or bright green; areoles small, commonly
3-3.5 cm. apart; larger spines 4-6 or fewer, very unequal, pinkish at first, becoming
white, or sometimes blackish in age, 3 cm. long or shorter, spreading, the largest
ones somewhat compressed; glochids brown; flowers 3.5 cm. long, the petals red;
stigma lobes 8-11, bright green; fruit 4 cm. long, strongly tuberculate, scarcely
This is a common plant in the central region, doubtless occurring
also in Sacatepe"quez and Chimaltenango and in some of the other
central and western departments. Generally the plants are about a
meter high, and it is only under favorable conditions that they be-
come tree-like. It is suspected that most of the tree-like plants have
been destroyed, since they are a pest and would not be permitted to
remain about dwellings or cultivated ground. It is altogether pos-
sible that more than a single species is represented by the Guate-
malan material we have referred here. It is presumably this species
that is planted so commonly for hedges about Quezaltenango, and
it is frequent in Huehuetenango and elsewhere in the Occidente. It
is also the arborescent Opuntia so plentiful on the dry, rocky hills
along the road between Guatemala and Amatitlan.
Opuntia guatemalensis Britt. & Rose, Cactaceae 1: 218, /. 285.
1919. Tuna; tuno.
Dry, rocky plains and hillsides, often in thickets, 2,300 meters
or less; endemic; type collected by Glover B. Wilcox in 1909 in some
unknown part of Guatemala, probably along the Pacific coast; Za-
capa; Jutiapa; Santa Rosa; Huehuetenango; Quezaltenango.
Plants erect or low and spreading, about 60 cm. high, glabrous; joints few or
rather numerous, often lustrous, green or pale green, sometimes with purplish
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 221
blotches below the areoles; areoles small, filled with brown wool; spines 1-3 at
each areole, small and inconspicuous, terete, acicular, white with a blackish tip
when young, in age gray, generally deflexed, commonly 1 cm. long or shorter but
sometimes 2 cm. long; flower buds reddish; flowers small, the petals lemon-yellow,
2.5 cm. long; stigma lobes cream-colored; fruit subglobose, deep red, very juicy,
edible; seeds 4 mm. in diameter.
Opuntia Guilanchi Griffiths, Kept. Mo. Bot. Card. 19: 265.
Dry slopes, 2,300-2,500 meters; San Marcos (near Tajumulco,
Steyermark 36555). Mexico (Zacatecas).
Plants shrub-like, 1.5-2 meters high, often with a short distinct trunk, grayish
green; joints broadly obovate or rounded-obovate, 20-25 cm. long, 14-16 cm.
broad, finely but inconspicuously pubescent; areoles rather large; spines 2-3 at
an areole, slightly compressed, at first whitish, becoming yellowish, 2.5 cm. long
or shorter; glochids pale yellow; fruit subglobose, 4 cm. in diameter, finely pubes-
cent, green turning rose, the pulp rose-red.
The Guatemalan locality is far remote from the Mexican range of
the species, but such a range is not improbable or inconsistent in this
genus, in which too many species seem to have been separated on
little else than their "widely different ranges." The specific name is
taken from a Mexican vernacular name of the fruit.
Opuntia pubescens Wendland in Pfeiffer, Enum. Cact. 149.
In thickets, about 1,350 meters; Baja Verapaz; Huehuetenango.
Plants erect and 60-90 cm. high, often much lower, much-branched; joints
easily detached, subterete, glabrous or pubescent, 3-7 cm. long; spines numerous,
short, brick-brown or buff -brown; flowers lemon-yellow, drying red; filaments
greenish; style white, the stigma lobes cream-colored; fruit small, only 2-2.5
cm. long, red, somewhat spiny; seeds 3 mm. in diameter.
The spines are barbed, and thus the joints easily become attached
to passing animals or objects and are transported from one locality
to another. In Guatemala, however, the species is apparently rare
Opuntia tomentella Berger, Monatsschr. Kakteenk. 22: 147.
1912; see also Federico Eichlam, Monatsschr. Kakteenk. 20: 81. 1910.
Tuna; tuna de monte.
Pastures, plains, or open hillsides, sometimes in oak forest, 1,300-
2,300 meters; endemic; Jalapa; Guatemala (type collected by Federico
222 FIELDI ANA: BOTANY, VOLUME 24
Eichlam, probably between Guatemala and Mixco); Sacatepe"quez;
Plants shrub-like, about a meter high, often with a short trunk; joints few or
numerous, obovate or oblong-obovate, 20-30 cm. long, 9-15 cm. broad, light green,
often somewhat lustrous, puberulent; areoles 1.5-3 cm. apart, small; spines 1-2,
acicular, white or yellowish, mostly 7-10 mm. long, porrect, sometimes wanting;
glochids few; flowers numerous, orange-red, 5-6 cm. long; petals obovate; filaments
yellowish green; style rose-colored, the stigma lobes white; ovary somewhat
tomentose, armed with numerous black glochids; fruit oblong, red within and
outside, acid, scarcely edible.
Eichlam states that the plant often is infested with cochineal
PACHYGEREUS Britton & Rose
Plants usually very large and tree-like, with few or numerous thick heavy
branches and a definite trunk, the branches ribbed, armed with clusters of stout
spines; flowers diurnal, with a rather short tube, the outer perianth segments short,
spatulate; stamens numerous, included, inserted along the throat of the perianth;
style included; ovary and perianth tube covered with small scales, these bearing
felt and bristles in their axils; fruits large, bur-like, dry, usually densely covered
with clusters of deciduous spines and bristles; seeds large, black.
About 10 species; all the others are Mexican.
Pachycereus lepidanthus (Eichlam) Britt. & Rose, Cactaceae
2: 76. 1920. Cereus lepidanthus Eichlam, Monatsschr. Kakteenk. 19:
Based on material collected by Federico Eichlam near El Rancho,
Chiquimula; El Progreso.
Plants simple or with a few stout branches, light green, the ribs 7-9, rather
low, separated by broad rounded intervals; areoles 1 cm. apart, small; radial spines
about 10, slender, 1.5 cm. long or the longer ones sometimes 5 cm. long; central
spines stouter, somewhat flattened, 3-6 cm. long; flowers 7 cm. long, 2.5 cm. broad;
perianth segments 3-4-seriate, 2.5 cm. long, 8 mm. broad, red below, sepia-brown
above, persistent on the fruit; ovary and perianth tube covered with membranous
scales; fruit dry.
Shrubs or trees, often scandent, branched and bearing normal green fleshy
entire leaves; spines binate or fasciculate in the leaf axils, neither vaginate nor
barbate; glochids none; leaves alternate, deciduous in age; flowers solitary, corym-
bose or paniculate, rotate, white, red or yellow; stamens numerous; stigma lobes
linear; fruit globose or very broadly turbinate, red or yellow, often bearing small
leaves, fleshy and juicy; seeds black, lustrous.
224 FIELDIANA: BOTANY, VOLUME 24
Species about 20, in tropical America. Some of the species are
used commonly as stocks on which to graft other cultivated cacti.
The fruits are eaten in some regions but are not very good ; we have
no information to the effect that they are eaten in Central America.
Plants scandent; branches with a pair of short recurved spines in each areole.
Plants erect shrubs or trees.
Spines long and straight P. autumnalis.
Spines acicular P. Conzattii.
Pereskia aculeata Mill. Card. Diet. ed. 8. 1768. Cactus Pereskia
L. Sp. PL 469. 1753. P. Pereskia Karst. Deutsch. Fl. 888. 1882.
Often planted for ornament, especially in patios. Probably native
of West Indies and tropical South America, but cultivated in other
regions. Figure 35.
A large slender vine, sometimes 10 meters long; spines of the older stems
solitary or 2-3 together, slender, straight; spines of the leaf axils binate or ternate,
2-4 mm. long, strongly recurved; leaves short-petiolate, lanceolate to oblong-
elliptic, mostly less than 10 cm. long, acute or acuminate, acute or obtuse at
the base; flowers paniculate or corymbose, white, pale yellow or tinged with pink,
2.5-4.5 cm. broad; ovary bearing small leaves and often also spines; fruit smooth
at maturity, subglobose, 1.5-2 cm. in diameter or somewhat larger; seeds black,
4-5 mm. broad.
The leaves are cooked and eaten in some parts of tropical Amer-
ica. The finest vine we have seen in Guatemala was one in the patio
of the hotel in Cuilapa. Its owner said that a friend had presented
it to her as a yellow Bougainvillea! In Chiapas (Mexico) the com-
mon name Buganvilla blanca is reported.
Pereskia autumnalis (Eichlam) Rose, Contr. U. S. Nat. Herb.
12: 399, tt. 52-54- 1909. Pereskiopsis autumnalis Eichlam, Mo-
natsschr. Kakteenk. 19: 22. 1909. Rhodocactus autumnalis F. M.
Knuth in Backeb. & Knuth, Kaktus-ABC 96. 1935. Manzanote;
Abundant in the lower Motagua Valley, on dry rocky plains and
hillsides, also in the Oriente, 200-900 meters; Baja Verapaz; Zacapa;
El Progreso; Jutiapa; Guatemala (Fiscal and lower); probably in all
the departments of the Oriente. El Salvador; Honduras; Nicaragua.
A large shrub or a tree as much as 9 meters high, the trunk low and thick,
often 40 cm. or more in diameter, very spiny, the crown more or less rounded
and spreading, often very dense; younger branches reddish brown; spines in the
leaf axils usually solitary, sometimes ternate, slender, 3-4 cm. long, rarely 16
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 225
cm. long; leaves thick and fleshy, oblong to orbicular, 2-8 cm. long, rounded to
subacute at the apex and apiculate, rounded or obtuse at the base; flowers solitary
near the ends of the branches, short-pedunculate; ovary covered with leaf -like
scales; flowers 4-5 cm. broad, orange-red; stamens numerous; fruit globose,
4-5 cm. in diameter, glabrous, bright yellow; seeds black, lustrous, 4 mm. long.
Sometimes called "mateare" in El Salvador and Nicaragua. The
name "manzanote" is the prevalent one in the lower Motagua Valley,
but in Jutiapa the tree is called "matial." Los Manzanotes is an
aldea of Zacapa, Manzanotal is a caserio of El Progreso, and both
names probably are very appropriate ones. This is one of the most
abundant trees about Zacapa, forming stands of great extent on the
plains, in association with spiny Leguminosae and other shrubs and
trees. At a short distance the Pereskia trees remind one strongly
of apple trees, being about the same size and form, and the resem-
blance is even greater when the manzanote trees are covered with the
yellow fruits, as they are in late April, when the trees are mostly
devoid of leaves. The thick trunks are densely covered with long
stout spines. The trees are quite useless except as hedge plants.
We have not noted Pereskia hedges in Guatemala, but in El Salvador
there are many miles of them; some of the country roads are shut in
on both sides with them. They are certainly successful for this pur-
pose, so far as obstructing large animals is concerned, but the hedges
are dangerous to man and probably also to domestic animals. The
minute irritating glochids, which can do great damage to the eyes,
are produced in myriads, and when there is a breath of wind these
are scattered through the air. The fruits may be edible in theory,
but no one would risk the glochids that cover them. No domestic
animals eat the plant, and even the wood is unsuitable for fuel, be-
cause in handling it one will become covered with the glochids, which
cause intense itching and irritation of the skin. The trees are con-
spicuous and somewhat ornamental when covered with their abun-
dant orange or orange-red flowers, about October.
Pereskia Conzattii Britt. & Rose, Cactaceae 1: 24. 1919.
Dry, hot plains at low elevations; Mexico and probably Guate-
mala (Eichlam, without locality) .
Tree 8-10 m. tall; bark of stems and branches brown and smooth; leaves
orbicular to obovate, acute, 1-2.5 cm. long; areoles small, with short white wool
and a few long hairs; spines 2-6 on young branches, 10-20 on main stem, acicular,
2-2.5 cm. long, at first yellowish brown, dark brown in age; flowers not known;
ovary bearing small scales; fruit naked, pyriform, more or less stalked at the
base, 3-4 cm. long; seeds black, glossy, 3 mm. long, with a small white hylum.
226 FIELDIANA: BOTANY, VOLUME 24
PERESKIOPSIS Britton & Rose
Shrubs or small trees, the older trunks consisting of a solid woody cylinder
covered with bark and resembling ordinary dicotyledonous tree trunks, the branches
sometimes elongate and more or less pendent or subscandent; areoles circular,
usually with spines and also bearing hairs, wool, and glochids; leaves broad and
flat, entire, unarmed; flowers similar to those of Opunlia, yellow or red; ovary
sessile, usually with leaves at the areoles; fruit red, juicy; seeds very hard, osseous,
few, covered with matted hairs.
About 10 species in tropical America, mostly Mexican. Only the
following is known from Central America.
Pereskiopsis Kellermanii Rose, Smithson. Misc. Coll. 50: 332.
In dry, often rocky thickets, 200-1,250 meters; endemic; Zacapa;
Chiquimula; Quiche"; type from Trapichito (probably in the Depart-
ment of Guatemala), Kellerman 6025. Honduras.
Stems glabrous, sometimes as much as 5 meters long and scandent, about 2
cm. in diameter; spines slender, acicular, brown, several or only 1, sometimes
absent, the glochids numerous, brown; young branches green and fleshy, the
older ones sometimes red; areoles on young stems bearing many long white hairs,
brown glochids, and often several brown spines; spines on older branches usually
solitary, almost black, and 2-3 cm. long; leaves fleshy, glabrous, lustrous, elliptic
to oblong-elliptic or suborbicular, acute or subacute, narrowed to the base, sessile;
flowers unknown; fruit red, glabrous, 3-6 cm. long, bearing small leaves, the areoles
with numerous brown glochids; seeds covered with matted hairs.
Specimens collected by Eichlam in Guatemala were placed by
Rose in Pereskia Conzattii Britt. & Rose. The material (U. S. Nat.
Mus.) is sterile but could belong to this species.
Slender epiphytes, usually pendent from the branches of trees, often much
branched and forming dense clumps; stems terete, angulate, or complanate and
leaf -like; leaves none or represented by minute bracts; areoles borne on the mar-
gins in flat-stemmed plants, along the ridges or irregularly scattered in other forms,
small, usually bearing hairs, wool, bristles, and flowers, never spines; flowers small,
usually solitary, nocturnal or diurnal; perianth segments distinct, few, sometimes
only 5, usually spreading, sometimes reflexed; stamens few or numerous, slender,
erect, inserted in 1-2 rows on the outer margin of the disc; style erect, the stigma
lobes 3 or more, usually slender and spreading; ovary small, sometimes depressed
or sunken in the stem; fruit globose or oblong, juicy, white or colored, generally
naked, sometimes bearing a few scales; seeds small, few or numerous.
A genus of about 50 or perhaps more not very clearly defined spe-
cies, all American and tropical with the exception of a few species
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 227
which are to be found from West Africa across to Madagascar, and
on islands of the Indian Ocean as far as Ceylon. These may have
been transported from America by birds or other agents.
Other species are described from Mexico and Central America.
Rhipsalis micrantha and R. ramulosa are similar vegetatively to
species of Epiphyllum. The latter has recently been placed in a seg-
regate of Epiphyllum but would seem to go better into Rhipsalis.
Stems compressed and often leaf-like, 8 mm. broad or broader, sometimes trigonous.
Stems mostly 1.5-3 cm. broad R. ramulosa.
Stems mostly less than 1 cm. broad R. micrantha.
Stems terete, usually less than 4 mm. in diameter.
Areoles filled with dense masses of tawny hairs 2-3 mm. long . . . .R. Bartlettii.
Areoles naked or when young bearing a few white bristles R. Cassutha.
Rhipsalis Bartlettii Clover, Bull. Torr. Bot. Club 65: 567,
/. 6, 7. 1938.
Known in Guatemala only from the type, Uaxactun, Pete*n,
Bartlett. Mexico (Veracruz).
Plants pendent, as much as 2 meters long, the branches slender, dichotomous
or sometimes verticillate, light green, the terminal joints sometimes less than 1 mm.
in diameter; areoles prominent, filled with dense masses of tawny hairs 2-3 mm.
long; flowering areoles not lanate, the ovary sunken in the stem; fruit white, glo-
bose, 3-4 mm. in diameter; seeds black, reniform, less than 1 mm. long, reticulate
We have seen no material of this species, but it is probably a
synonym of R. Cassutha; the dense masses of hairs that are supposed
to characterize the "species" may be of the nature of insect galls,
such as have been found in other species of Rhipsalis.
Rhipsalis Cassutha Gaertn. Fruct. & Sem. 1: 137. 1788. Tatache
(Pete"n, Maya) ; bejuco de quebradura.
On branches of trees, 600 meters or less; Pete"n; Alta Verapaz;
Santa Rosa; Escuintla; Suchitepe"quez ; Retalhuleu; Huehuetenango;
probably in all the lowland departments. Southern Florida; Mexico;
British Honduras to Panama; West Indies; South America; Ceylon;
Plants often forming dense masses a meter long or more, much-branched;
stems fleshy, terete, when young bearing several white bristles in the areoles,
naked when old, mostly 2-4 mm. in diameter, rather pale green, the branches
generally in pairs but often verticillate; flowers lateral, solitary, white; petals
228 FIELDIANA: BOTANY, VOLUME 24
2 mm. long or somewhat longer; fruit naked, white or pink, translucent, ripening
in only a few days after flowering, globose, 5 mm. or less in diameter.
In cultivation often called "mistletoe cactus," the white berries
and naked stems suggesting some of the Loranthaceae. The plant
is often grown in hanging baskets in Guatemala. When wild it grows
most plentifully on large trees, drooping in great masses from the
highest branches. The flowers persist for several days. At mid-day
they are almost rotate, but they close in late afternoon. Said to be
used in fractures of bones of men and horses, hence bejuco de que-
Rhipsalis micrantha (HBK.) DC. Prodr. 3: 476. 1828. Cactus
micranthus HBK. Nov. Gen. et Sp. 6: 65. 1823. R. Tonduzii Weber,
Diet. Hort. Bois. 1046. 1898. Pitahaya.
Epiphyte in forest, 1,000-1,300 meters, San Marcos (Volcan Taju-
mulco, Steyermark 37591). Costa Rica to Peru.
Pendent epiphytic plants, much-branched; primary stems terete or with 2,
sometimes 3 narrow wings or angles; secondary stems or branches flat, with 2
or sometimes 3 wings or angles, up to 1 cm. broad, the secondary stems occa-
sionally borne in groups of 3; areoles 2-4 cm. apart on the wings, with or without
short bristles or scales; flowers small, white, 5-7 mm. long including the ovary;
tepals 2-3 mm. long, linear-oblong, obtuse; fruit a globose, pulpy berry to about
10 mm. long.
The available material of this species is inadequate. The incom-
plete description is drawn from the Guatemalan material cited.
Rhipsalis ramulosa (Salm-Dyck) Pfeiffer, Enum. Cact. 130.
1837. Cereus ramulosus Salm-Dyck, Hort. Dyck. 340. 1834. R. cori-
acea Polak. Linnaea 41 : 562. 1877. Disocactus ramulosus Kimnach,
Cact. & Succ. Journ. Am. 33: 14, t. 1961. Guacamayo.
Epiphytic in moist forest, from near sea level to about 1,500 me-
ters; Pete"n; Alta Verapaz; Escuintla; Suchitepe*quez; Solola; Hue-
huetenango; San Marcos. West Indies; Mexico; British Honduras,
Central America and Panama; south to Brazil and Bolivia. Figure
Pendent epiphytes, branched, a meter long or more; stem terete, slender, some-
what woody; branches or secondary stems flat and leaf -like, up to 15 cm. long and
to 4 cm. broad, remotely crenate; areoles about 1.5 cm. apart and without bristles
when mature; flowers relatively small, single at the areoles, to about 12 mm. long;
tepals 10 or fewer, to about 15 mm. long, the outer ones oblong, the inner lanceo-
late; stamens 12-30, inserted in a single area at the top of the receptacle, exserted;
style exceeding the stamens or about the same length, stigma 3-4-lobed; fruit
baccate, ovoid, up to 8 mm. long.
FIG. 36. Rhipsalis ramulosa. 1, Habit; X ^g. 2, Stems with buds, flowers
and immature fruits; X 4 A- 3 and 4, Flower from side and front; X 2J^.
5, Longitudinal section of receptacle; X 6. 6, Stigma; X 12. 7, Fruit; X 2^-
8, 9, Seeds; X 16. Courtesy of Myron Kimnach. Drawn by Mrs. M. Bios.
230 FIELDIANA: BOTANY, VOLUME 24
SELENICEREUS Britton & Rose
Plants slender, trailing or scandent, the stems elongate, ribbed, with generally
7-10 ribs, often with aerial roots; areoles small, sometimes elevated on small knobs,
usually bearing small spines; flowers large, often very large, nocturnal, the perianth
tube elongate, often somewhat curved; scales of the ovary and perianth tube small,
usually with long felt, hairs, and bristles in their axils; upper scales and outer peri-
anth segments similar, narrow, the inner perianth segments broad, white, usually
entire; stamens numerous, in 2 separated clusters, one cluster forming a circle at
the top of the tube, the other scattered over the long slender throat, the filaments
much elongate, weak; style elongate, thick, often hollow, the stigma lobes numer-
ous, slender, entire; fruit large, reddish, covered with clusters of deciduous spines,
bristles, and hairs.
A small "genus" of perhaps a half dozen not very distinct species
(Britton & Rose enumerate 16 and Backeberg lists 24!). It is ques-
tionable that the two species enumerated below are distinct.
Hairs of the flower areoles ferruginous or fulvous S. grandiflorus.
Hairs of the flower areoles bright white S. hondurensis.
Selenicereus grandiflorus (L.) Britt. & Rose, Contr. U. S. Nat.
Herb. 12: 430. 1909; Cactaceae 2: 197, t. 33. 1920. Cactus grandi-
florus L. Sp. PI. 467. 1753. Cereus grandiflorus Mill. Card. Diet,
ed. 8, no. 11. 1768.
Thought to be grown as an ornamental and perhaps escaped in
the country; native in the West Indies.
Plants usually terrestrial but sometimes epiphytic, the stems often much elon-
gate and scandent, about 2.5 cm. in diameter, green or bluish green; ribs usually
7-8, sometimes fewer, low, separated by broad rounded intervals; spines acicular,
1 cm. long or less, brown or yellowish brown, in age grayish, intermixed with numer-
ous whitish hairs; flower buds densely covered with brown or fulvous hairs; flowers
about 18 cm. long, the outer perianth segments narrow, salmon-colored, the inner
segments white, acute, entire; style often longer than the inner perianth segments;
fruit ovoid or subglobose, whitish or pink, juicy, about 8 cm. long.
This is the plant most often known in cultivation under the name
"night-blooming Cereus." It long has been a favorite cactus for
cultivation in the United States. In the tropics it often grows lux-
uriantly, covering walls and small trees.
Selenicereus hondurensis (Schum.) Britt. & Rose, Contr. U. S.
Nat. Herb. 12: 430. 1909; Cactaceae 2: 199, /. 275. 1920. Cereus
hondurensis Schum. in Weingart, Monatsschr. Kakteenk. 14: 147.
On trees or rocks, near sea level, Izabal. Honduras.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 231
Stems scandent or clambering, 1.5 cm. in diameter, green; ribs 7-10, low; are-
oles 6-10 mm. apart; spines rather short, 5-7 mm. long, stout, usually surrounded
by much longer, white hairs or bristles, these most conspicuous on young branches;
flowers 20 cm. long or larger; outermost perianth segments linear, brownish, acu-
minate, yellow, the inner segments pure white, 10 cm. long, 1-1.5 cm. broad; scales
of the ovary and perianth tube linear, bearing numerous long bristly hairs in the
There are in our collections several specimens and photographs
of Selenicereus hondurensis. We are not sure how these may be dis-
tinguished from S. grandiflorus, to which probably most of the spe-
cies of Selenicereus might be reduced.
WERCKLEOCEREUS Britton & Rose
Plants epiphytic, scandent, the stems triangulate, emitting numerous aerial
roots; areoles bearing short bristles or very weak spines and a tuft of felt; flowers
short-funnelform, the tube rather broad; ovary and perianth tube bearing many
areoles, each with several almost black, acicular spines and a tuft of short black
felt, subtended by minute scales; outer perianth segments lanceolate, subacute,
narrow, the inner segments broader; stamens numerous; style about equaling the
longer stamens, the stigma lobes several, linear; fruit globose, its areoles spiny.
Two species are recorded for the genus, the other from Costa
Rica. The genus is dedicated to Carlos Werckl (see Standley: Car-
los Werckle, in Science 63: 221-223. 1926), who lived for many years
in Costa Rica, devoting the greater part of his life to the study of
the flora of that country.
Werckleocereus glaber (Eichlam) Britt. & Rose, Addisonia 2:
13, t. 47. 1917; Cactaceae 2: 216, t. 39. 1920. Cereus glaber Eichlam,
Monatsschr. Kakteenk. 20: 150. 1910 (described from cultivated
plants obtained from the Pacific coast of Guatemala) ; Sacatepe"quez.
Stems slender, triangulate, about 2 cm. thick, pale green and slightly glaucous,
scandent by aerial roots; ribs somewhat nodose, the areoles borne on the upper
part of the elevation, small, 3-4 cm. apart; spines 2-4 at each areole, 1-3 mm. long,
acicular but with enlarged bases; flowers 10 cm. long or more, the ovary and peri-
anth tube bearing clusters of yellow or brown, acicular spines; inner perianth seg-
ments white, oblanceolate, acute, somewhat serrate; style pale yellow, the stigma
WILMATTEA Britton & Rose
Plants usually epiphytic, scandent, very slender, the stems emitting aerial
roots, armed with few very small spines, 3-angulate; flowers small for the group,
generally solitary at the areoles, nocturnal, with a narrow limb and a very short
232 FIELDIANA: BOTANY, VOLUME 24
tube; ovary covered with ovate imbricate reddish scales, each subtending a small
areole filled with felt and sometimes 1 or more bristles; filaments and style short.
The genus consists of a single species named for Mrs. T. D. A.
Cockerell (Wilmatte P. Cockerell), who made a collection of plants
Wilmattea minutiflora Britt. & Rose, Cactaceae 2: 195, t. 32,
f. 272. 1920. Hylocereus minutiflorus Britt. & Rose, Contr. U. S.
Nat. Herb. 16: 240, t. 69. 1913. Cereus minutiflorus Vaupel, Mo-
natsschr. Kakteenk. 23: 86. 1913.
At or little above sea level; Izabal (type collected near Lake
Izabal, R. H. Peters) . British Honduras; Atlantic coast of Honduras.
Plants slender, as much as 9 m. long, the stems 2.5 cm. or usually much less
in diameter, deep green, the angles acute, the areoles 2-4 cm. apart; spines usually
1-3, minute, brownish; flowers about 5 cm. long, very fragrant, the tube 1 cm. long
or less; outer perianth segments linear, red on the costa and apex, 3-4 cm. long, the
inner segments very narrow, white, acute; stamens white, 1 cm. long, inserted at
the base of the inner perianth segments; scales of the ovary oblong to ovate, purple
or greenish at the base; style white, 2 cm. long, the lobes white.
The specific name is a most inappropriate one, since a flower two
inches long can scarcely be considered minute.
ZYGOCACTUS Schumann. Christmas cactus
Plants epiphytic, small, the stems dichotomously much-branched, compressed,
divided into short joints; flowers terminal, irregular; ovary terete, smooth, bearing
minute scales above; perianth tube abruptly bent just above the ovary; stamens
slender, white, arranged in 2 groups; style slender, purple, as long as the stamens,
the stigma lobes linear, purple, erect, adherent to one another; fruit purple, turgid,
terete, pulpy, the skin thin; seeds dark brown or almost black, lustrous.
The genus consists of a single species, and perhaps is not really
distinct from Epiphyllum.
Zygocactus truncatus (Haw.) Schum. in Mart. Fl. Bras. 4,
pt. 2: 224. 1890. Epiphyllum truncatum Haworth, Suppl. PL Succ.
Native of the mountains of Rio de Janeiro, Brazil, but cultivated
as a pot plant in tropical and temperate regions; common as a house
plant in Guatemala.
Plants lustrous dark green, the joints about 3 cm. long, sharply serrate, with
2 conspicuous teeth at the truncate apex; terminal areole filled with brown wool
FIG. 37. Wilmattea minutiflora. 1, Stems with buds and flower; X 1. 2, Flower;
X 1. 3, Flower, longitudinal section; X 1. 4, Floral bract and areole; X 4.
5, Nectaries and stamen bases; X 3. 6, Stigma; X 2. 7, Ovules and funicles;
greatly enlarged. Courtesy of Myron Kimnach. Drawn by Mrs. M. Bios.
234 FIELDIANA: BOTANY, VOLUME 24
and bristles; flowers 6-7 cm. long, the tube 2 cm. long; inner perianth segments
bright red or white, oblong, obtuse to acute, reflexed; fruit obovoid, 1.5-2 cm.
This is one of the best of all cacti for house culture since it grows
luxuriantly and in the United States generally blooms freely about
Christmas; hence the common name in use for it.
The Myrtiflorae is a large order divided by Engler and Diels
(Syllabus der Pflanzenfamilien, ed. 11, 1936) into 23 families. Ten
of these families occur in Guatemala, two of them introduced and
eight with native species. The Myrtaceae and the Melastomaceae
are the two largest families in Guatemala as they are throughout
most of the rest of the American tropics. The representatives of the
families of this order in Guatemala are mostly woody. There are
many fine large forest trees in the order; many of the deep green
seashore forests, the mangroves, belong here. Plants of economic
importance are found in several of these families: fruits in the
Myrtaceae and Punicaceae; exotic nuts in the Lecythidaceae; tans
and dyes of considerable importance in the Rhizophoraceae; timbers
of some importance in the Combretaceae, and woods used locally
for construction or fuel in most of the families. Ornamentals are
also found in most of these families.
Leaves usually opposite but sometimes alternate, simple. There
is a tendency from perigyny to epigyny in the order. The flowers
are cyclic and the development of a hypanthium (calyx tube) upon
which the stamens and petals usually are inserted, is diagnostic; the
hypanthium sometimes is completely adnate to the ovary.
Trees or shrubs, rarely herbs, the bark often separating into meshed fibers;
leaves opposite or alternate, entire, mostly penninerved; stipules none; flowers per-
fect or by abortion polygamous or unisexual, regular, capitate, umbellate, short-
racemose, or rarely solitary, the inflorescences pedunculate or sessile, terminal or
axillary; perianth inferior, usually with a slender tube, the limb 4-5-lobate, the seg-
ments imbricate, spreading in anthesis, equal or the 2 inner ones slightly smaller;
scales as many as the perianth lobes or twice as many, affixed within the tube; sta-
mens as many or twice as many as the perianth segments, affixed at the middle of
the tube or higher, the filaments filiform, short; anthers erect or dorsifixed near the
base, 2-celled, the cells parallel, dehiscent by longitudinal slits; hypogynous disk
annular, cupular, composed of 4-5 scales, or none; ovary sessile or short-stipitate,
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 235
entire, 1-celled or 2-celled; style short or elongate, usually eccentric in the 1-celled
ovary, central in the 2-celled ovary, the stigma terminal, capitate or subdiscoid;
ovules 1 in each cell, laterally attached near the apex of the cell, anatropous,
pendulous; fruit indehiscent, nut-like, drupaceous, or baccate; seed pendulous
or laterally affixed, usually with crustaceous testa; endosperm copious and carnose,
or sometimes scant or none; embryo straight, the cotyledons carnose, the radicle
About 40 genera, mostly in Australia and South Africa, but sev-
eral occur in South America, and one in the United States and Can-
ada. No other genus is found in Mexico or Central America, but
Schoenobiblus occurs in Panama.
DAPHNOPSIS Martius & Zuccarini
Reference: Lorin I. Nevling, Jr., A revision of the genus Daphnop-
sis, Ann. Mo. Bot. Gard. 46: 257-363, illus. 1960.
Dioecious shrubs or small trees; leaves alternate, exstipulate, membranaceous
or subcoriaceous; inflorescences axillary or terminal, racemose, umbellate, some-
times fasciculate or rarely the flowers solitary; flowers tetramerous, perigynous,
unisexual; calyx usually urceolate or campanulate, the lobes equal or not, im-
bricate; petals 8, 4 or none, often connate into a papilliform or squamiform ring
in the throat of the calyx tube; stamens in staminate flowers 8, inserted in two
series on the calyx tube, the upper series opposite the calyx lobes, the lower alternate
with them, sessile, subsessile or with short filaments, introrse; pistillate flowers
usually smaller than the staminate, with 4 or 8 or no staminodia, pistil 1, superior
and borne on a gynophore, with a single pendulous ovule; style terminal; fruit a
Species 46, according to Nevling. Others are known from Mexico
and Central America. The genus is limited to tropical America.
Two sides of leaves very unlike in color; sericeous-pubescent below.
Two sides of leaves essentially concolorous; glabrous below or if pubescent then
the two sides concolorous.
Secondary peduncle deciduous with the flowers, the peduncle terminated with
a ball-like torus D. americana.
Secondary peduncles persistent on the torus of the primary peduncle.
Adult leaves pubescent beneath D. malacophylla.
Adult leaves glabrous or nearly so.
Leaf blades small, 3-5.5 (-11) cm. long, somewhat narrowed to an obtuse
tip D. ficina.
Leaf blades large, 7-25 cm. long, very acute or acuminate.
Leaf blades oblong-elliptic, mostly 8-11 cm. long; umbellate inflorescences
mostly 5-7-flowered D. Tuerckheimiana.
Leaf blades mostly 17-25 cm. long, narrowly oblong-oblanceolate; um-
bellate inflorescences usually more than 7-flowered D. radiata.
236 FIELDIANA: BOTANY, VOLUME 24
Daphnopsis americana (Mill.) J. R. Johnston, Contr. Gray
Herb. 34: 242. 1909. Laurus americana Mill. Gard. Diet. ed. 8,
Lauras 10. 1768. Daphne Bonplandiana Kunth, Syn. PL Aequin. 1 :
447. 1922. Daphnopsis Lindenii Meissn. in DC. Prodr. 14: 523. 1857.
D. Bonplandiana Standl. Contr. U. S. Nat. Herb. 23: 1013. 1924.
D. americana ssp. guatemalensis Nevling, Ann. Mo. Bot. Gard. 46:
312. 1960. Coralillo; coralillo bianco; camaman; capulincito; chaca-
Moist, wet or dry thickets or scrub forest; Zacapa; Jutiapa;
Santa Rosa; Guatemala; Escuintla; Huehuetenango. Mexico.
Shrubs or trees to 10 m., the branches reddish, glabrous or nearly so; leaves
elliptic, lanceolate or oblanceolate, acute or acuminate, coriaceous, shining, gla-
brous, 5-15 cm. long, 1-4 cm. broad, cuneate to the base into a short petiole 1
cm. or less long, nerves inconspicuous, 15-20 pairs, veins many, prominulous;
inflorescence umbellate, the flowers sessile on the torus or nearly so and dehiscent
from it, pubescent when young; calyx of pistillate flowers obconic, about 2 mm.
long, those of staminate flowers longer; fruit about 1 cm. long.
There is some question as to the proper name for the tree here
described. Nevling's D. americana ssp. americana contains at least
two "units": one has broad obovate leaves which would seem to
accord with Miller's description; the other is difficult to distinguish
from Nevling's subspecies guatemalensis, as also from D. Bonplandi-
ana and D. Lindenii. We have seen no authentic material of Laurus
americana Mill, and apparently Nevling had not.
Daphnopsis ficina Standl. & Steyerm. Field Mus. Bot. 22: 254.
1940 (type from Dept. Guatemala, Garcia Salas 1442) . Chilillo.
Wet forest or cloud forest; Jalapa; Zacapa; Baja Verapaz; Guate-
mala; Quiche". Mexico (Chiapas).
Shrubs or small trees to 5 m. tall, the young branches reddish, sparsely pubes-
cent or glabrous; leaves small, elliptic to oblong-elliptic, acute or acuminate,
rarely obtuse, contracted or decurrent into a short petiole at the base, blades
3-11 cm. long, 1-4 cm. broad, glabrous above, sparsely pubescent or glabrous
below, nerves and veins prominent on both sides; inflorescences terminal or lateral,
sericeous when young, umbelliform; calyx tube of pistillate flowers urceolate,
1.5-3 mm. long, that of staminate flowers narrower, sericeous; fruit subglobose,
nearly glabrous, about 1 cm. long.
Nevling maintains this species in his monograph, yet it looks
suspiciously like D. Tuerckheimiana Donn.-Sm.
Daphnopsis malacophylla Standl. & Steyerm. Field Mus. Bot.
23: 68. 1944 (type from Huehuetenango, Steyermark 49104).
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 237
Wet, cool cloud forest, 1,500-3,000 meters; endemic; Huehue-
A shrub or tree 4.5-6 meters high, the branches ferruginous, the young ones
densely hirtellous or short-hirsute with fulvous, spreading or ascending hairs;
leaves chartaceous, on petioles 4-8 mm. long, oblong-elliptic or oblong-obovate,
7-16 cm. long, 2.5-6.5 cm. broad, acute or acuminate with an obtuse tip, acute at
the base, sparsely pilose above with slender, spreading or subappressed hairs,
in age glabrate, densely and softly pilose beneath with spreading or subappressed
hairs, the lateral nerves about 7 pairs, irregular, divergent at a rather wide angle,
the veins prominulous, laxly reticulate; peduncles terminal and extra-axillary,
1.5-4.5 cm. long, densely hirtellous, the umbels dense, many-flowered, the short
pedicels 3 mm. long or less; staminate perianth 6 mm. long, appressed-hirtellous,
the lobes scarcely 1 mm. long; pistillate perianth 5 mm. long, strigose, the lobes
rounded-ovate, spreading; style short-exserted ; fruit ovoid, 7 mm. long, sparsely
Daphnopsis monocephala Donn.-Sm. Bot. Gaz. 47: 261. 1909
(type from El Progreso, Kellerman 5714). D. retifera Standl. &
Steyerm. Field Mus. Bot. 22: 254. 1940 (type from Jutiapa, Steyer-
Moist or dry, brushy slopes or plains, 650-1,100 meters; endemic;
El Progreso; Chiquimula; Jutiapa; Guatemala.
A shrub 1.5-3 meters high, the branches stout, densely covered with long and
short, subappressed or spreading, brownish or fulvous hairs; leaves short-petiolate,
the petioles stout, 5-8 cm. long, often marginate to the base, the two sides distinctly
bicolored, the blades subcoriaceous, elliptic-oblong to lance-oblong or oblanceolate-
oblong, 7-17 cm. long, 2-6.5 cm. broad, very obtuse or rounded at the apex, usually
somewhat narrowed toward the apex, cuneate-attenuate to the base, glabrous
and lustrous above, pale beneath, when young densely sericeous with very long
hairs, in age densely and softly short-pilose with chiefly spreading hairs, the
lateral nerves about 8 pairs, very oblique, the veins prominent, closely and con-
spicuously reticulate; peduncles subterminal, solitary, simple, 2-3 cm. long, the
flowers all sessile or nearly so; staminate perianth with a tube 4.5 mm. long, the
lobes 1.5 mm. long.
This is doubtless the best marked of all the North American
species of the genus.
Daphnopsis radiata Donn.-Sm. Bot. Gaz. 14: 30. 1889 (type
from Alta Verapaz, Tuerckheim 1163). D. Selerorum Gilg, Verh.
Bot. Ver. Brandenb. 48: 153. 1917 (type from Guatemala, Seler 2866).
Damp or wet, mixed forests, 900-2,000 meters, endemic; Alta
Verapaz; Quezaltenango; Huehuetenango. Figure 38.
Dioecious shrubs or small trees to 8 m., the young branches sparsely pubescent
or usually glabrous; leaves elliptic, elliptic-lanceolate or elliptic-oblanceolate,
FIG. 38. Daphnopsis radiata. A, Habit; X 1 A. B, Pistillate flower; X 5.
STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 239
acuminate, cuneate to the base into a short petiole, obscurely pulvinate or usually
entirely glabrous, 7-25 cm. long and 2-10 cm. broad; inflorescence terminal or
usually so, umbellate, pubescent to somewhat canescent; peduncle of pistillate
inflorescence to 6 cm. long, or usually shorter, of the staminate about the same,
secondary peduncles as much as 1 cm. long, flower dehiscent at apex of secondary
peduncle; calyx tube of pistillate flowers urceolate, 4-4.5 mm. long, sparsely
sericeous outside, the tube of staminate flowers narrow, about 6 mm. long; ovary
ovoid, glabrous or slightly pubescent, the style about 2.5 mm. long, as long as
or exceeding the calyx; fruit ovoid, 7-9 mm. long.
Nevling has maintained D. radiata and D. Selerorum as distinct
species but we find no difference in them. D. malacophylla, placed
by Nevling as a synonym of D. Selerorum, seems to be as distinct as
most species in this difficult genus.
Daphnopsis Tuerckheimiana Donn.-Sm. Bot. Gaz. 16: 13.
1891 (type from Pansamala, Tuerckheim 1039).
Wet mixed forests, 1,200-2,500 meters, endemic; Alta Verapaz;
A shrub of 1.5 meters or a small tree, the branches sparsely sericeous at first;
leaves short-petiolate, subcoriaceous, glabrous or when young sparsely sericeous
beneath, mostly oblong-elliptic and 8-11 cm. long, 2.5-4 cm. broad, acute or
acuminate, acute at the base; umbellate inflorescences lateral and terminal,
mostly 5-7-flowered, the staminate secondary peduncles short, the pistillate 7
mm. long or less, sericeous; staminate perianth 7-8 mm. long, the lobes short;
fruit black, with thin flesh, glabrous, ovoid, 7 mm. long, short-rostrate at the obtuse
Daphnopsis flavida Lundell (Phytologia 2: 3. 1941), described
from Matuda 4159, which was collected on Mount Ovando in Mex-
ico near the Guatemalan border, probably belongs here as a syn-
The family Elaeagnaceae is represented by a few species in North
America, but none are native in tropical America. Elaeagnus philip-
pinensis Perr., native of the Philippine Islands, has been planted at
Bananera, Izabal and perhaps elsewhere. It is a shrub or small tree
with alternate short-petiolate entire leaves which are green and gla-
brous on the upper surface, covered beneath with a dense coat of
whitish and brown, closely appressed scales.
240 FIELDIANA: BOTANY, VOLUME 24
LYTHRACEAE. Loosestrife Family
Reference: E. Koehne, Lythraceae, Pflanzenreich IV. 216. 1903.
Annual or perennial herbs, shrubs, or trees; leaves generally opposite, rarely
verticillate or scattered, entire; stipules none, or 2-10 or more and subulate, rarely
2 and inserted with the leaves; flowers actinomorphic or rarely zygomorphic,
usually perfect, 3-16-parted, mostly 4-6-parted, axillary and solitary or cymose,
rarely paniculate, in Cuphea mostly extra-axillary or interpetiolar; pedicels mostly
bibracteolate; calyx broad or tubular, the lobes in bud usually valvate, appendages
often alternate with the lobes; petals inserted in the throat of the calyx between
the lobes, regular or zygomorphic, often crumpled in bud, sometimes fugacious,
rarely none; stamens inserted at different heights on the calyx tube (hypanthium),
as many as the calyx lobes, or fewer, or more numerous; anthers generally dorsifixed;
ovary free, sometimes stipitate, completely or incompletely 2-6-celled, a hypogy-
nous disk sometimes present; style simple or none; stigma small, capitate or
punctiform, rarely 2-lobate; ovules numerous or sometimes only 2, anatropous,
ascending; fruit capsular, dehiscent or indehiscent, dry; embryo straight, without
endosperm, the cotyledons flat or rarely convolute, often auriculate-cordate.
About 22 genera and 450 species or more, most numerous in trop-
ical regions. Other genera represented in Central America are Heimia
and Grislea, the former of which should be found in Guatemala.
Calyx tubular, slightly curved or else calcarate or gibbous at the base; herbs or
small shrubs; flowers usually zygomorphic Cuphea.
Calyx not tubular or, if so, neither calcarate nor gibbous at the base.
Leaves conspicuously dotted with small black glands; shrubs Adenaria.
Leaves not black-dotted.
Plants herbaceous, rarely somewhat suffrutescent at the base.
Calyx tubular Lythrum.
Calyx campanulate or hemispheric.
Capsule septicidally dehiscent; leaves acute at the base Rotala.
Capsule irregularly ruptured; leaves auricula te at the base. .Ammannia.
Plants woody throughout, trees or large shrubs.
Flowers 12-16-parted; calyx about 3 cm. long; native trees Lafoensia.
Flowers 4-7-parted; calyx much smaller; cultivated trees or large shrubs.
Stamens 8; leaves opposite Lawsonia.
Stamens 15-many; leaves alternate Lager stroemia.
Shrubs or small trees, black-glandular throughout, the young branches 4-
angulate; flowers mostly 4-parted, small, imperfectly unisexual, the inflorescences
axillary, umbelliform, the outer pedicels bracteolate at the base; calyx turbinate,
in fruit semiglobose, not nerved, villous within above the stamens, the lobes
equaling or shorter than the tube, without appendages; petals white or pale yellow-
ish; stamens 7-12, equal or unequal; ovary turbinate-globose, stipitate or rarely
sessile, generally 2-celled, usually hirtellous at the apex; style filiform, the stigma
large, 2-lobate; fruit indehiscent, 2-celled, subcoriaceous; seeds cuneate-obovoid.
The genus consists of a single species.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 241
Adenaria floribunda HBK. Nov. Gen. & Sp. 6: 188, *. 549. 1823.
Dry or moist thickets, 600 meters or less; Santa Rosa (near El
Molino) ; Retalhuleu (between Nueva Linda and Champerico) . West
Indies; southern Mexico; Honduras; Nicaragua; Costa Rica; Pan-
ama; widely distributed in South America.
A shrub or small tree, 4.5 meters high or less, often densely branched, the
branches slender, puberulent; leaves short-petiolate, membranaceous, oblong-
lanceolate or ovate-oblong, 5-12 cm. long, long-acuminate to acute, rounded to
subacute at the base, green above, glabrous, paler beneath, puberulent on the
nerves, densely black-punctate; flowers white, densely puberulent, densely black-
punctate; calyx 3 mm. long; petals longer than the calyx; stamens long-exserted;
capsule 4 mm. in diameter.
In some parts of Central America, particularly in the Canal Zone
where it is known as "fruta de pavo," this is a common shrub in second
growth, but in Guatemala it seems to be very rare.
Annuals, glabrous or nearly so, the stems more or less 4-angulate; leaves
opposite, narrow, sessile; flowers very small, axillary or cymose; calyx campanulate
to globose or ovoid, 4-angulate, 4-dentate, often with small appendages in the
sinuses; petals 4, deciduous; stamens 4-8, inserted on the calyx tube, the filaments
short or elongate; ovary enclosed in the calyx tube, subglobose, 2-4-celled, rupturing
Species about 20, mostly in tropical regions. Only the following
are known in Central America. Ammannia latifolia L. is known in
Flowers pedicellate, in small axillary cymes A. auriculata.
Flowers sessile or nearly so, 2-3 together in the leaf axils A. coccinea.
Ammannia auriculata Willd. Hort. Berol. 1:7,*. 7. 1803.
Muddy margin of a small waterhole, 200 meters; Zacapa. Cen-
tral and southern United States; Mexico; El Salvador; Honduras;
Cuba; South America; Asia; Africa.
Plants erect, 5-25 cm. high, often widely branched; leaves sessile, linear-
lanceolate or oblong, acute or subacute, auriculate at the base, 1-3.5 cm. long;
flowers in small axillary short-pedunculate few-flowered cymes, the pedicels 1-3
cm. long; calyx green, 2 mm. long; petals small, purple; stamens exserted; capsule
usually enclosed in the persistent calyx.
Ammannia coccinea Rottb. PI. Hort. Havn. Descr. 7. 1773.
Wet open soil at or little above sea level; Pete"n; British Hon-
duras. Eastern and central United States; Mexico; Honduras; Pan-
242 FIELDIANA: BOTANY, VOLUME 24
ama; West Indies; South America; also on several island groups in
Oceania. Possibly adventive in Iran.
Plants erect glabrous, 15-40 cm. tall, often much branched below somewhat
succuent laves'linear-lanceolate, 2.5-7 cm. long, 2-6 mm broad acute or
acurmna e dilated and auriculate-clasping at the base; flowers 1-5 in each leaf axil,
Se or nearly so; calyx 2 mm. long; petals small, purple, fugacious; style elongate
Annual or perennial herbs, rarely shrubs; leaves opposite or rarely verticillate;
flowers zygomorphic, 6-parted, small or large, the flowers racemose, the racemes
oftenTafy the flowers often appearing to be all axillary, alternate or opposite,
rarefy vert dilate; bractlets 2 or none; calyx tubular, often calcarate at the base;
petals 6 rarely 2 or none, very rarely 4; stamens 11, rarely 9 or 6; ovary sessile,
usuaUy w?th a basal, dorsal or rarely cupuliform disk, or the disk sometimes
absent; ovules numerous or few, often 3, very rarely only 2; capsule finally de^s-
cent and the calyx cleft by the emergent reflexed placentae; seeds lentiform,
usually narrowly winged.
Species about 200, all American and nearly all tropical. A very
few additional ones are found in southern Central America,
genus is better represented in Guatemala than in any other part of
Central America, and Mexico has 75 species or more. There is one
other species in Guatemala but the material is not adequate for c
Flowers small, the calyx less than 1 cm. long.
Leaves linear or oblanceolate; plants stiffly erect, suffrutescent, almost always
growing on rocks in the edges of streams.
Leaves linear; pedicels much shorter than the subtending leaves.
Leaves mostly oblanceolate.
Pedicels almost as long as or longer than the adjacent leaves; stems and
leaves pubescent to glabrous C. utnculosa.
Pedicels much shorter than the adjacent leaves; stems and leaves hispid-
ulous C - *top>*-
Leaves broader than oblanceolate, usually much broader, never linear.
Leaves long petiolate; plants herbaceous, annual C. secundiflora.
Leaves sessile or on very short petioles; annual or perennial, sometimes lig-
Calyx glabrous, gland-dotted C. mimuloides.
Calyx pubescent, usually hispidulous, at least on the nerves.
Leaf blades acute at the base.
Plants herbaceous or essentially so, sometimes suffrutescent at the
b ase C. carthagenensis.
Plants ligneous throughout.
Calyx sparsely hispidulous, otherwise glabrous; leaves glabrous ex-
cept on the margins C. flarisetula.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 243
Calyx glandular-hispidulous and puberulent; leaves pubescent on
both surfaces decandra.
Leaf blades obtuse or rounded at the base.
Leaves glabrous on the upper surface; plants suffrutescent.
Leaves hispidulous on the upper surface; slender annuals.
Stamens about equaling the calyx tube C. Wrightii.
Stamens much shorter than the calyx tube.
Leaves sessile or nearly so C . micrantha.
Leaves petiolate c SderL
Flowers large, the calyx more than 1 cm. long, often much longer.
Petals purple or violaceous, sometimes almost black, sometimes very dark red
but drying purple.
Stems usually hirsute with long spreading hairs C. aequipetala.
Stems puberulent or hirtellous with short hairs.
Interior wings of the calyx retrorse-hirtellous; petals 2 C. cyanea.
Interior wings of the calyx glabrous; petals usually 6 C. pinetorum.
Petals scarlet or bright red, retaining this color when dried, rarely none.
Flowers all or partly in terminal bracteate racemes C. infundibulum.
Flowers never racemose, borne in the axils or near the axils of large leaves.
Calyx bright red; petals none or minute c. Heydei.
Calyx green throughout or nearly so; petals usually present and conspic-
Appendages of the calyx conspicuously aristate C. aristata.
Appendages of the calyx not aristate.
Leaves pilose on the upper surface with spreading hairs. . .C. Nelsonii.
Leaves scabrous or glabrate on the upper surface.
Calyx and stems with a dense covering of uniform short spreading
viscid hairs C. sanguinea.
Calyx thinly hispid with long spreading hairs, often also scabrous or
stngose, sometimes with very long hairs near the base only
| stems scabrous or thinly hispid.
Pedicels axillary or arising between the internodes; leaf blades
mostly obtuse or rounded at the base C. axilliflora.
Pedicels inserted between the petioles (not in axils); leaf blades
mostly long attenuate at the base C. appendiculata.
Cuphea aequipetala Cav. Icon. PI. 4: 57, t. 382, /. 2. 1797.
C. aequipetala var. laevicaulis Koehne, Bot. Jahrb. 2: 411. 1882. C.
aequipetala var. hispida Koehne, I.e. Granadita; coralillo; clarincillo;
pegajosa (fide Aguilar) .
Dry to wet soil, fields, thickets, or pine-oak forest, sometimes in
hedges or cultivated ground, 1,500-2,500 meters; Alta Verapaz; Baja
Verapaz; Jalapa; Guatemala; Sacatepequez; Chimaltenango; Solola;
Quich^ ; Huehuetenango. Mexico; Honduras.
244 FIELDIANA: BOTANY, VOLUME 24
Plants perennial, generally from a hard woody root, usually much branched,
stems a meter long or less, generally prostrate or procumbent, sometimes sprawling
over small shrubs, usually densely hispid with long spreading purplish hairs, some-
times glabrate; leaves sessile or nearly so, ovate or lance-ovate, mostly 1-3 cm.
long and 5-15 mm; broad, acute to obtuse at the apex, rounded or acute at the base,
glabrate or scabrous above, sometimes sparsely hispid, hispid or glabrous beneath
with spreading hairs; flowers axillary, on pedicels 1-5 mm. long, the pedicels
bracteolate at the apex; calyx 13-23 mm. long, purple or purplish, sparsely hispid;
appendages small and inconspicuous; petals deep purple or rose-purple, half as
long as the calyx or longer; stamens exserted.
Cuphea appendiculata Benth. PL Hartweg. 61. 1839.
Moist thickets or cliffs, sometimes in pine forest, 1,000-2,300
meters; Chiquimula; Solold; Quezaltenango. Southern Mexico.
A shrub 1.5 meters high or less, usually sparsely branched, the branches rather
slender, when young thinly hispid with spreading yellowish hairs, soon glabrate,
sometimes also strigillose; leaves short-petiolate, lanceolate or ovate-lanceolate,
5-14 cm. long, 1.5-4.5 cm. broad, narrowly long-acuminate, usually long-
attenuate to the base, scabrous above and sometimes with a few spreading hairs,
hispidulous beneath on the veins, elsewhere scaberulous or glabrous; flowers sub-
tended by large leaves, the pedicels 2-4 mm. long, bracteolate above the middle;
calyx 27-33 mm. long, green, strigillose or scabrous and usually also hispid with
pale hairs; appendages linear; petals bright red, 7-9 mm. long, obovate; disk
almost semicupular; ovules 8.
Cuphea arista ta Hemsl. Diagn. PI. Mex. 51. 1880.
Moist or wet, pine or mixed forest, 800-1,500 meters; Pete"n
(Camp 35, British Honduras boundary) ; Zacapa (Sierra de las Mi-
nas); type from the Motagua Valley, Salvin & Godman, probably
from Sierra de las Minas. British Honduras.
A slender shrub a meter high or less, the stems puberulent; leaves on very short
petioles, oblong-lanceolate, 3-4.5 cm. long, attenuate-acuminate, rounded to
attenuate at the base, rough on the upper surface and slightly scaberulous, paler
beneath and glabrous or sparsely hispidulous; pedicels interpetiolar, solitary,
3-10 mm. long, bracteolate at the apex; calyx 20-30 mm. long, greenish, very
sparsely setulose and also scaberulous; appendages conspicuously 1-2-aristate;
petals scarlet, the 2 dorsal ones 15-17 mm. long, the 4 ventral ones 4-5 mm. long;
This has been reported from British Honduras as C. axilliflora
Cuphea axilliflora Koehne, Pflanzenreich IV. 216: 170. 1903.
C. appendiculata var. axilliflora Koehne, Bot. Jahrb. 2: 412. 1882.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 245
Mostly in wet pine forest, 1,250-1,450 meters; Huehuetenango-
Alta Verapaz (type from Coban, Tuerckheim 172); Quiche*. British
Plants herbaceous or frutescent, 1-2 meters high, erect or often subscandent
or sprawling over shrubs, the stems strigose or sometimes hispidulous with yellowish
hairs; leaves short-petiolate, ovate or oblong-ovate, 5-10 cm. long 2 5-5 cm
broad, acuminate, usually rounded at the base but sometimes acute, scabrous
above and very rough to the touch, scabrous beneath and sometimes hispidulous
on the veins; floral leaves little smaller than the others, the pedicels 4-8 mm
long, bracteolate at the apex; calyx 28-33 mm. long, greenish, strigose or scaberulous
and sometimes also hirsute with eglandular yellowish hairs; appendages lanceolate
or oblong; petals flame-red, about 10 mm. long, broadly obovate, the 4 ventral
ones sometimes rudimentary and subulate; disk very thick, subglobose, deflexed-
stamens exserted; ovules 11.
Cuphea calophylla Cham. & Schlecht. Linnaea 2: 361. 1827
C. microstyla Koehne in Mart. Fl. Bras. 13, pt. 2: 224. 1877 (type
from Guatemala, Skinner). C. calophylla var. orthodisca Koehne
Bot Jahrb. 2: 138. 1881. C. calophylla var. microstyla Koehne,'
Reported by Koehne as collected in Guatemala by Bernoulli and
by Skinner, the localities not indicated. Southern Mexico; British
Honduras to Panama; southward to Brazil.
Plants perennial, usually erect and rigid, commonly much branched and 35
cm. high or less, often suffrutescent below, the older stems ferruginous, the young
ones puberulent and usually also hispidulous; leaves numerous, sessile or nearly
so ovate-oblong to lanceolate, 1-5 cm. long, acute, usually rounded or very obtuse
at the base, thinly hispidulous or glabrate above, hispidulous beneath and some-
times strigose or scabrous; flowers mostly at the ends of the branches, the floral
leaves generally much reduced, the pedicels 2-6 mm. long, bracteolate at the
apex; calyx 6 mm. long, hispidulous, the teeth subequal; petals purple or violet-
stamens included; ovules mostly 6-8.
Called "hog weed" and "hog bush" in British Honduras. The
plant often invades cultivated ground, and because of the large roots
and the tough hard stems, it is difficult to eradicate. It is rather
strange that there are no Guatemalan specimens at hand of this
weedy plant, which is common almost throughout the Atlantic low-
lands of Central America. There can be no doubt that it does grow
Cuphea carthagenensis (Jacq.) Macbride, Field Mus. Bot. 8:
124. 1930. Lythrum carthagenense Jacq. Stirp. Amer. Hist. 148. 1763
C. balsamona Cham. & Schlecht. Linnaea 2: 363. 1827. Pica-mano
(fide Aguilar); caqui mesU (Alta Verapaz); chichibe (Pete'n).
246 FIELDIANA: BOTANY, VOLUME 24
Mostly in moist or wet soil, meadows, thickets, open banks, sand-
bars along streams, often a weed around dwellings or in waste and
cultivated ground, 1,800 meters or less; Pete"n; Alta Verapaz; Baja
Verapaz; Izabal; El Progreso; Zacapa; Chiquimula; Jalapa; Jutiapa;
Santa Rosa; Escuintla; Guatemala; Chimaltenango; Quiche"; Hue-
huetenango; Suchitepe"quez; Retalhuleu; Quezaltenango; San Mar-
cos. Mexico; British Honduras to El Salvador and Panama; West
Indies; South America. Introduced in Hawaii.
Plants essentially annual but probably persisting for a longer time, commonly
erect and 50 cm. high or less, rarely suffrutescent, often much branched, the stems
puberulent and more or less glandular-pilose; leaves short-petiolate, obovate to
ovate or lance-oblong, 2-5 cm. long, acute or obtuse, generally acute at the base,
more or less scabrous on both surfaces and rough to the touch, often sparsely
hispidulous, when young sometimes strigose; flowers small and inconspicuous,
usually subtended by large leaves, the pedicels very short, bracteolate at the apex;
calyx 4.5-6 mm. long, sparsely hispidulous or sometimes glabrous in age, pale
green; petals small, pale purple; stamens 11, included; ovules 4-8.
This is a very common and variable weedy plant almost through-
out Central America except at high elevations.
Cuphea cyanea DC. Prodr. 3: 85. 1828. C. cyanea var. hirtella
Koehne, Bot. Jahrb. 2: 417. 1882. Clarincillo.
Moist or dry thickets or pine-oak forest, 1,900-3,700 meters;
Jalapa; Chimaltenango; Solola; Huehuetenango. Central and south-
A slender shrub 1-1.5 meters high, the stems pubescent or hispidulous; leaves
on petioles 1.5 cm. long or less, ovate or oblong-ovate, 2-8 cm. long, 1-5 cm.
broad, acuminate or long-acuminate, usually rounded or subcordate at the base,
scabrous on both surfaces and sometimes hispidulous; inflorescence often paniculate,
the leaves all small and bract-like, the pedicels 10 mm. long or less, bracteolate
near the apex; calyx 16-23 mm. long, red and yellow, viscid-hispidulous, the spur
usually large and well developed; petals purple, spatulate or obovate; stamens
short-exserted; ovules 5-6.
Cuphea decandra Ait. Hort. Kew. ed. 2. 3: 3, 151. 1811. Lyth-
rum ciliatum Swartz, Prodr. Veg. Ind. Occ. 76. 1788. C. ciliata
Koehne, Bot. Jahrb. 1: 454. 1881, not C. ciliata Ruiz & Pavon, 1794.
A species of the Greater Antilles and Colombia, represented in
continental North America by the following variety:
Cuphea decandra var. Purpusii (Brandeg.) Bacigalupi, Contr.
Gray Herb. 95: 9. 1931. C. Purpusii Brandeg. Univ. Calif. Publ.
Bot. 4: 378. 1913.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 247
At 270 meters; Alta Verapaz (Chama, Harry Johnson 521).
A slender shrub 20-40 cm. high, woody throughout or nearly so, usually
much branched, the branches densely glandular-hirtellous, brownish; leaves on
very short petioles, obovate or oblong, 1.5-4 cm. long, acute or obtuse, acute at
the base, pubescent on both surfaces, usually densely so; floral leaves much reduced
and bract-like, the flowers in distinct racemes, the pedicels 2-5 mm. long, not
bracteolate; calyx 7-11 mm. long, glandular-hirtellous and puberulent; petals
purple or lavender, oblong, the ventral ones 4-5 mm. long, the dorsal ones slightly
smaller; stamens not exserted, the style in age short-exserted; ovules 10-20.
Cuphea flavisetula Bacigalupi, Contr. Gray Herb. 95: 8. 1931.
Dry open rocky slopes, or often in pine forest, sometimes on ser-
pentine or on rocks in the edges of streams, 1,400-2,500 meters;
Jalapa; Huehuetenango. Western and southern Mexico; Honduras.
An often densely branched shrub, woody throughout or nearly so, 30-60
cm. high, sometimes procumbent, the stems puberulent and sometimes hispidulous,
in age often almost wholly glabrous; leaves sessile or nearly so, lance-elliptic to
elliptic or broadly obovate, mostly 1-2 cm. long, acute to rounded at the apex,
acute at the base, hispid-ciliate; flowers in terminal racemes, the floral leaves
reduced and bract-like, the pedicels not bracteolate; calyx 7-8 mm. long, hispidulous
with yellowish hairs; petals oblanceolate, the 2 largest 2.5 mm. long; stamens 9,
subexserted; ovules 12-18.
Cuphea Heydei Koehne ex Donn.-Sm. Bot. Gaz. 19: 256. 1894.
Dry or moist forest, 1,800-2,100 meters; endemic; Quiche" (type
from Nebaj, Heyde & Lux 4480).
An erect shrub a meter high, or the stems sometimes elongate and trailing,
rooting at the nodes, as much as 3.5 meters long, reddish, when young densely
hirsute; leaves short-petiolate, elliptic or lanceolate, mostly 5-9 cm. long and 2-3.5
cm. broad, narrowly long-acuminate, acute or attenuate at the base, glabrous
above or nearly so, paler beneath and sparsely hispid or almost glabrous; flowers
few, the floral leaves scarcely reduced, the pedicels 1-2 cm. long, bracteolate at
the apex; calyx scarlet, 22-28 mm. long, sparsely hispid; stamens exserted; petals
none or minute; ovules 9-14.
The species is known from three collections, all made at Nebaj.
An attractive plant, perhaps worthy of cultivation.
Cuphea hyssopifolia HBK. Nov. Gen. & Sp. 6: 199. 1823.
C. hyssopifolia f. subrevoluta Koehne, Bot. Jahrb. 2: 153. 1881 (type
from Coban, Tuerckheim 17). Hierba de burrioncillo (fide Aguilar).
Usually on rocks in the edges of swift streams, sometimes in wet
forest, 1,500 meters or less; Alta Verapaz; Izabal; Quiche"; Huehue-
tenango. Mexico; Honduras. Figure 39.
FIELDIANA: BOTANY, VOLUME 24
FIG. 39. Cuphea hyssopifolia. A, Habit; XI. B, Flowers, one partially dis-
sected; X 2.
An erect shrub 25-50 cm. high, usually with very numerous suberect slender
branches, the stems pubescent and sometimes appressed-hispidulous; leaves sub-
sessile, crowded, linear or nearly so, 1-3 cm. long, suberect or spreading, glabrous
or with a few scattered hairs on the costa, 1-nerved; floral leaves not reduced, the
pedicels 2-7 mm. long, bracteolate at the apex; calyx glabrous or with a few
scattered short hairs, 6-8 mm. long; petals pale purple or white; ovules 5-8.
This is one of several Central American plants, species of Cuphea,
Aster, Eupatorium, Lindenia and a few other groups, whose habitat
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 249
is almost wholly confined to large rocks along the edges of usually
swift streams. These rocks project above the mean level of the
stream, but during times of heavy rain the plants often are covered
by rushing water. Most of these plants, obviously, have tough stems
that are able to withstand the debris carried by the often rapid cur-
rents of water.
Cuphea infundibulum Koehne in Mart. Fl. Bras. 13, pt. 2:
In forest, or on brushy rocky slopes, 800-1,900 meters; Chi-
quimula; Huehuetenango. Mexico; Costa Rica; Panama.
An erect shrub or herb, usually a meter high or less, the slender stems strigose
or scabrous, often also glandular-hirsute; leaves short-petiolate, lanceolate to ovate-
oblong or ovate, 5-15 cm. long, 1.5-6 cm. broad, generally acuminate at each
end, strigose-scabrous, sometimes densely strigose when young, sometimes hispidu-
lous beneath on the veins; flowers mostly in terminal paniculate racemes, the
floral leaves usually reduced to small bracts, the pedicels 2-4 mm. long, bracteolate
above the middle; calyx 2-3 cm. long, green or tinged above with red, strigose
and more or less hirsute; petals bright red; disk semicupular; ovules about 7.
Cuphea micrantha HBK. Nov. Gen. & Sp. 6: 196. 1823. At
750 meters; Santa Rosa (Rio de los Esclavos, Heyde & Lux 3764).
Honduras; Greater Antilles; South America.
Plants 40 cm. high or less, the stems pubescent and glandular-hirsute; leaves
oblong or lance-linear, 1.5-6.5 cm. long, acute or acuminate, acute to rounded
at the base, hispid; floral leaves little reduced, the pedicels 1-2.5 mm. long, brac-
teolate at the apex; calyx 4-7 mm. long, hispidulous; petals violaceous or purple,
Cuphea Valerii Standl. & L. Wms. from Honduras is a synonym
of this species. Closely allied to C. carthagenensis.
Cuphea mimuloides Cham. & Schlecht. Linnaea 5: 570. 1830.
Maja hypericoides Klotzsch in Schomb. Fl. & Faun. Guian. 1191.
1848, without diagnosis. C. mimuloides var. guianensis Koehne in
Engl. Bot. Jahrb. 1: 446. 1881. C. mimuloides var. hypericoides
Koehne, Pflanzenreich IV. 216: 101. 1903.
In bogs or wet places, 900-1,400 meters; Jutiapa. Mexico; Brit-
ish Honduras; Honduras; West Indies; northern South America.
A slender erect annual, usually about 20 cm. tall, sometimes to 50 cm.,
branched, the stems glabrous or sparsely puberulent; leaves short-petiolate or
subsessile, ovate-oblong to narrowly oblong or oblanceolate, 1-2 cm. long, obtuse,
acute or attenuate at the base, glabrous or nearly so; floral leaves reduced, the
slender pedicels 5-9 mm. long, spreading; calyx 4-7 mm. long, glabrous or minutely
250 FIELDIANA: BOTANY, VOLUME 24
glandular, distended in age by the capsule; petals 1.5-2 mm. long, white to pink;
stamens scarcely equaling the calyx, the style included; ovules about 100.
Cuphea Nelsonii Rose, Contr. U. S. Nat. Herb. 5: 137, t. 15.
Moist forest, 1,600-2,500 meters; endemic; Alta Verapaz; Baja
Verapaz; El Progreso; Huehuetenango (type collected between Jacal-
tenango and San Martin, E. W. Nelson 3600).
Erect and suffrutescent, 1.5 meters high or less, or sometimes larger and
subscandent, the stems ferruginous, densely viscid-hirsute; leaves short-petiolate,
lanceolate to ovate, 3-6 cm. long, acute or acuminate, obtuse or acute at the
base, rather densely scabrous and hirsute on both surfaces; flowers few, the floral
leaves greatly reduced, the pedicels bracteolate at the apex; calyx 20-30 mm. long,
greenish, densely pilose with long spreading purplish hairs; petals bright red;
Koehne gives the habitat of the species erroneously as Mexico.
Cuphea pinetorum Benth. PI. Hartweg. 74. 1839. C. Hookeri-
ana Walp. Repert. Sp. Nov. 2: 107. 1843. Pititos morados; pegajosa
rosada; leoncillo; boca de murcielago; clavo rojo (fide Aguilar).
Moist or dry thickets or forest, common in pine-oak forest,
1,000-2,400 meters; Alta Verapaz; El Progreso; Zacapa; Jalapa;
Chiquimula; Santa Rosa; Escuintla; Guatemala; Sacatepe*quez; Chi-
maltenango; Solola; Quiche"; Huehuetenango; Totonicapan; Quezal-
tenango (type from San Ramon, Hartweg 529) ; San Marcos. Central
Mexico; Honduras; Nicaragua. Figure 40.
Plants perennial, slender, usually erect and a meter high or less, the stems
brittle, usually sparsely branched, sometimes suffrutescent below but essentially
herbaceous, scabrous; leaves subsessile, mostly lanceolate or linear-lanceolate,
3-11 cm. long, long-attenuate, acute or rounded at the base, scabrous; floral leaves
reduced to linear bracts, the inflorescence thus terminal and racemose, the pedicels
4-7 mm. long, interpetiolar, bracteolate below the apex; calyx 17-20 mm. long
or sometimes shorter, pinkish, viscid-hirtellous, the spur very short or well de-
veloped; petals from nearly white to dark red or almost black, the dorsal ones
half as long as the calyx or longer, the others much reduced; stamens exserted;
Koehne treated C. Hookeriana as distinct from C. pinetorum, re-
porting both species from Guatemala. He separated the two on the
basis of the size and shape of the smaller petals, certainly not a prac-
tical character, and one that probably has no systematic importance,
especially since, when kept distinct, the two species have the same
ranges. In spite of its rather large flowers, this is not an attractive
plant, being rather harsh and faded in appearance. The petals are
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 251
FIG. 40. Cuphea pinetorum. A, Terminal part of a plant; X 1. B, Receptacle
showing attachment and exserted series of stamens; X 3. C, Large petal. D, Small
often almost black. This is a characteristic species of mixed pine-oak
forest in the mountains of Guatemala, abundant in many regions
and quite variable.
Cuphea platycentra Lemaire, sometimes called "cigarette
plant," and in Guatemala "cigarrito," is planted occasionally in
gardens for ornament. It is a suffrutescent or herbaceous, almost
glabrous plant native of southern Mexico; the calyx is bright red
and about 2 cm. long.
Cuphea sanguinea Koehne, Bull. Herb. Boiss. 7: 565. 1899.
Chocshdn (Huehuetenango) .
In moist thickets or open forest, mostly in pine forest, 1,200-
3,000 meters; endemic; Huehuetenango (type collected above Jacal-
252 FIELDIANA: BOTANY, VOLUME 24
tenango, Seler 2629); San Marcos (volcanoes of Tajumulco and
Plants herbaceous or suffrutescent, erect and about a meter high, or the
stems sometimes supported on other vegetation and as much as 3 meters long,
densely viscid-hispidulous with reddish or purple hairs; leaves on petioles 4-7
mm. long, ovate to ovate-oblong, 3-7 cm. long, 2-3.5 cm. broad, acuminate,
rounded or obtuse at the base, very scabrous above, scabrous beneath or along
the nerves densely short-hirtellous; floral leaves little if at all reduced, the pedicels
4 mm. long, bracteolate; calyx 20-25 mm. long, greenish, densely viscid-hirtellous;
petals bright red, 12 mm. long; stamens short-exserted.
Cuphea secundiflora Sesse" & Moc. ex DC. Prodr. 3: 84. 1828.
C. leptopoda Hemsl. Diagn. PL Mex. 52. 1880 (type collected between
Esquipulas and Jupilingo, Chiquimula, Bernoulli 747).
At about 900-1,300 meters; Chiquimula; Santa Rosa; Quiche";
Huehuetenango. To central Mexico.
Annual, erect, 30-40 cm. high, soft-stemmed, little branched, the stems
pubescent and glandular-hirtellous with pale hairs; lower petioles 1-3.5 cm. long,
the upper ones shorter; leaf blades ovate or lance-ovate, 3.5-7 cm. long, 2-3.5 cm.
broad, acute or short-acuminate, acute at the base, thin, rather densely scabrous,
sparsely setulose above; inflorescences terminal, spike-like or head-like, the flowers
sometimes secund, the branchlets short, 1-few-flowered, the pedicels very short;
calyx hispidulous, green, 6-7 mm. long; 2 dorsal petals deep purple, cuneate-oblong,
the 4 ventral petals much smaller or absent; stamens and style included.
Cuphea Seleri Koehne, Bull. Herb. Boiss. 7: 565. 1899.
Known only from the type, Uaxacanal, Huehuetenango, C. & E.
Stems about 20 cm. long, simple, densely glandular-hirtellous; leaves on
petioles 5-8 mm. long, ovate, about 3 cm. long and 1.5 cm. broad, narrowed to the
subobtuse apex, rounded at the base, minutely strigillose, more or less setulose
above and beneath along the nerves; inflorescence simple; calyx 7-8 mm. long,
hispidulous; petals probably purple, the 4 ventral ones about one-third as wide
as the 2 dorsal ones and very acute; disk almost horizontal.
Cuphea utriculosa Koehne in Mart. Fl. Bras. 13, pt. 2: 452.
1877. C. utriculosa var. panamensis Koehne, I.e. C. utriculosa var.
Donnell-Smithii Koehne, Bot. Gaz. 18: 203. 1893. Parsonsia utricu-
losa Standl. Contr. U. S. Nat. Herb. 23: 1017. 1924 (type from So-
lold, Shannon 403). Nido de anguila (fide Aguilar).
Almost always on large rocks in or at the edge of swift streams,
1,500 meters or lower; Pete"n; El Progreso; Izabal; Zacapa; Chiqui-
mula; Jalapa; Jutiapa; Santa Rosa; Sacatepe"quez ; Guatemala; Chi-
STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 253
maltenango; Quiche"; Huehuetenango; Suchitepe"quez; Retalhuleu.
Mexico; British Honduras to Panama.
Plants perennial, stiff and tough, erect, usually 30-40 cm. high, densely
branched above; branches suberect, ferruginous, suffrutescent below, pubescent
or glandular-hirtellous, sometimes glabrate; leaves very numerous, sessile or
subsessile, oblanceolate or linear, 1-3.5 cm. long, 3-7 mm. broad, obtuse or sub-
acute, attenuate to the base, glabrous, sometimes glandular-ciliolate; floral leaves
usually much smaller than the lower ones, the flowers usually numerous, the slender
pedicels 6-15 mm. long, not bracteolate; calyx 4-7 mm. long, usually glabrous,
greenish; petals pale purple, obovate or cuneate-oblong, the ventral ones 3-4
mm. long; stamens included, the style sometimes exserted; ovules 25-75.
The species varies greatly in quality and quantity of pubescence.
Cuphea Wrightii Gray, PI. Wright. 2: 56. 1853.
Moist or rather dry thickets, open banks, fields, often on rocky
or grassy slopes in pine-oak forest, 800-2,800 meters; Chiquimula;
Jalapa; Jutiapa; Santa Rosa; Guatemala; Sacatepe"quez; Chimalte-
nango; Huehuetenango. Southwestern United States; Mexico; Hon-
duras; Costa Rica; Panama.
An erect annual, usually branched, 10-40 cm. high, the stems slender, glandular-
hispidulous with mostly purplish hairs, usually also pubescent, sometimes retrorse-
pilose below; leaves small, on petioles 6-14 mm. long, lanceolate to oblong or
ovate, 1.5-4 cm. long, 5-25 mm. broad, narrowed to the subacute apex, rounded
to acute at the base, strigose-scabrous and sparsely hispidulous, rarely glabrate;
floral leaves similar to the others but mostly much smaller, the branchlets 1-few-
flowered, the pedicels 2-7 mm. long, bracteolate near the apex; calyx 5-6 mm.
long, greenish, hirsute with short purplish hairs; petals purple, the 2 dorsal ones
obovate or almost orbicular, the ventral ones much smaller, cuneate-oblong;
stamens not exserted; style short-exserted in age.
Closely allied to Cuphea carthagenensis.
Heimia salicifolia (HBK.) Link should occur in the Pacific foot-
hills of Guatemala. It is common and widely distributed in Mexico,
and has been collected at San Vicente, El Salvador. It is a low gla-
brous shrub with linear or narrowly lanceolate leaves and rather large
and showy, bright yellow, axillary flowers.
Glabrous trees or shrubs; leaves opposite, coriaceous, lustrous, penninerved,
produced into a broad acumen with an obtuse, often deflexed tip bearing at the
apex a conspicuous subterminal pore; flowers large, 8-16-parted, racemose or
FIELDIANA: BOTANY, VOLUME 24
FIG. 41. Lafoensia punicifolia. A, Fruiting branch; X ^. B, Under surface
of leaf tip showing pore; X 5. C, Copy of Sesse and Mocino's sketch of a flower
for their L. mexicana; X 1.
subpaniculate, the bracts foliaceous but smaller than the leaves; bractlets large,
covering the buds; calyx campanulate or semiglobose, coriaceous, the limb plicate,
the lobes caudate, implicate; appendages of the calyx none; petals large, erose;
stamens 16-32 and uniseriate, inserted at or below the middle of the tube, long-
exserted, spirally contorted in bud; anthers narrowly oblong or linear, recurved;
ovary subsessile or stipitate, the placenta globose or disciform; ovules numerous;
style very long, the stigma subcapitate; fruit capsular, hard and woody, 1-celled,
loculicidally 2-4-valvate; seeds complanate, broadly winged.
Ten species, in tropical America.
Only one reaches North
Lafoensia punicifolia DC. Me"m. Soc. Phys. Geneve 3, pt. 2:
86, t. 1. 1826. Palo de culebra.
Moist forest, 600-1,300 meters; Jalapa; Santa Rosa; Huehuete-
nango. Southern Mexico; El Salvador; Costa Rica; Panama; north-
ern and western South America. Figure 41.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 255
A tree of 12-24 meters, the young branchlets obscurely tetragonous; leaves
on petioles 3-6 mm. long, coriaceous, usually yellow-green when dried, lustrous,
oblong or lanceolate, 5-11 cm. long, obtuse or acuminate, acute at the base or
obtuse, the nerves many pairs, conspicuously elevated beneath; inflorescence
racemose or subpaniculate, the pedicels 2-3.5 cm. long; flowers 12-16-parted;
calyx campanulate, 2.5-3 cm. long, rounded at the base; petals yellowish green
or yellow, turning red in age, 3-3.5 cm. long; filaments as much as 12 cm. long,
the style 13 cm. long; capsule broadly ovoid or ellipsoid-ovoid, apiculate, 3-5
cm. long, terete; seeds, with the wing, oblong, about 3 cm. long and 1 cm. broad.
Called "trompillo" and "cuyapo" in El Salvador. The tree is
common in Guatemala at some places along the road from Cuilapa
to Chiquimulilla. In this genus the heartwood is bright greenish
yellow or olive, of high luster, hard and heavy or moderately so,
rather fine in texture; it is not difficult to work, taking a smooth
finish and high polish. It is used for miscellaneous purposes in some
regions but has little or no commercial importance. In Brazil the
bark serves as the source of a yellow dye.
LAGERSTROEMIA L. Crape myrtle
Trees or shrubs, the leaves generally alternate; stipules binate, minute, decidu-
ous; flowers often large and showy, mostly 5-8-parted, racemose or cymose and
often paniculate, the bracts and bractlets small or minute, the pedicels articulate
at the insertion of the bractlets; calyx semiglobose or turbinate, coriaceous, terete
or costate, the lobes usually caudate; calyx appendages none or very small; petals
usually unguiculate; stamens 15-200, in 1-many series, the anthers broadly elliptic
or orbicular; ovary globose, or elongate at the apex, glabrous or tomentose, 3-6-
celled, the ovules numerous; style slightly exceeding the longest stamens, the
stigma scarcely thicker than the style; capsule ellipsoid or oblong, ligneous, 3-6-
valvate; testa of the seeds produced at the base into an appendage, produced
above into a wing.
About 30 species, in eastern and southern Asia and Australia.
Lagerstroemia indica L. Sp. PI. ed. 2. 734. 1762. Jupiter;
Planted commonly for ornament at low and middle elevations,
rarely above 1,500 meters. Native of eastern and southern Asia,
but grown as an ornamental plant in most warm regions.
A shrub or small tree, the bark smooth, gray, the young branchlets tetragonous,
glabrous; leaves sessile or short-petiolate, oblong-elliptic to rounded, 2-7 cm. long,
acute to rounded at the base, short-acuminate to emarginate at the apex, sub-
coriaceous, glabrous or nearly so, often hirtellous beneath on the costa; flowers
white, pink, or purple, usually 6-parted, the panicles 5-20 cm. long, many-flowered,
the branches usually short-hirtellous, the pedicels 3-15 mm. long; calyx 7-10 mm.
long, glabrous, the lobes erect; petals 12-20 mm. long; stamens 36-42; capsule
ellipsoid-globose, 9-13 mm. long, half included in the calyx.
256 FIELDIANA: BOTANY, VOLUME 24
Crape myrtle is a popular garden shrub in the lower parts of
Guatemala. Although it seldom or never is planted in the higher
mountains, it could be grown there, since in the United States it
withstands the winter as far north as the Potomac Valley.
LAWSONIA L. Henna
Glabrous shrubs or small trees, the branchlets often indurate and spinescent,
the young branchlets tetragonous; leaves small, opposite, short-petiolate, penni-
nerved; stipules conic, minute, whitish; flowers 4-parted, in terminal pyramidal
panicles, small, the pedicels bracteolate at the base or middle; calyx broadly
turbinate, subcoriaceous, terete, the lobes slightly longer than the tube, ovate-
triangular, without appendages; petals short-unguiculate, reniform, corrugate;
stamens 8, the filaments thick, subulate, exserted; ovary sessile, 2-4-celled, the
style stout; fruit globose, indehiscent or irregularly ruptured; seeds thick, trigonous-
pyramidal, the testa spongious at the apex.
The genus consists of a single species.
Lawsonia inermis L. Sp. PI. 349. 1753. L. alba Lam. Encycl. 3 :
106. 1789. Reseda; ricidrdn (Pete"n, fide Lundell).
Planted commonly for ornament at low and middle elevations
and as high as 2,100 meters, or probably higher; often growing in
hedges where apparently not planted by man, and more or less nat-
uralized in some regions, especially on the Pacific plains. Native
probably in eastern Africa and Asia; now grown for ornament or for
its sweet-scented flowers in most tropical regions.
A low tree, usually 6 meters high or less, with somewhat spreading crown;
leaves oblong or obovate, 1-2.5 cm. long, mucronate-acuminate, narrowed at the
base; panicles mostly 5-20 cm. long; calyx 3-5 mm. long; petals 4-6 mm. long,
pale yellow; fruit 4-6 mm. long.
Henna is not a handsome tree, and in Central America apparently
it is planted because of the agreeable fragrance of the otherwise un-
attractive flowers. The odor is similar to that of mignonette. In the
Orient the leaves are much used for staining the nails, hands, and
feet yellow, and also for dyeing the hair and beard. If a paste of
the leaves is applied to the hair or beard, it soon produces a bright
red color, which is much admired by some classes of Mohammedans.
If after this treatment an indigo paste is applied, the hair becomes
jet black. The plant yields a dull red dye for cloth. A perfume often
is extracted from the flowers. No use is made of the plant in Central
America, although the use of henna upon women's hair is not un-
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 257
LYTHRUM L. Loosestrife
Annual or perennial herbs with slender, often wiry stems; leaves small, opposite,
verticillate, or alternate; flowers usually small, mostly 4-6-parted, sometimes
obscurely zygomorphous, often dimorphous, solitary in the leaf axils or forming
terminal spikes or racemes; calyx tubular, usually with evident appendages;
stamens 4-12, the ventral ones usually inserted much higher than the dorsal ones;
ovary sessile or nearly so, the style generally developed; capsule 2-valvate, the
valves often 2-lobate, cartilaginous or submembranaceous; seeds 8-many, small.
About 24 species, widely distributed in temperate and warm re-
gions. Only the following are found in Central America. The species
are poorly marked, for the most part. Those of Mexico and Central
America and of the rest of North America for that matter are not
well understood; they are in need of thorough revision.
Calyx about 6 mm. long, or shorter L. acinifolium.
Calyx 8-12 mm. long L. vulneraria.
Lythrum acinifolium Sesse" & Mocino ex Koehne, Bot. Jahrb.
1: 322. 1881. L. acinifolium Sess & Moc. ex DC. Prodr. 3: 81. 1828,
Wet forest or thickets or on open wet banks, 2,200 meters or less;
Pete"n; Baja Verapaz; Quiche"; Huehuetenango. Mexico.
Plants erect, 1-2 meters high, herbaceous or somewhat woody, often much
branched, the stems slender, the older ones brown; leaves opposite, subsessile,
oblong or elliptic-oblong, 1-3 cm. long, obtuse, at the base rounded or obtuse;
flowers solitary in the leaf axils (as in other Guatemalan species), the calyx 4-6
mm. long; petals purple; annulus at the base of the ovary very narrow; style
much exceeding the stamens.
Perhaps not distinct from the following species.
Lythrum vulneraria Schrank, PI. Rar. Hort. Monac. t. 27. 1819.
L. Kennedyanum HBK. Nov. Gen. & Sp. 6: 194. 1823. Clavo de tierra
Moist or wet thickets, sometimes along streams, 1,400-2,000 me-
ters; Alta Verapaz; Sacatepe"quez; Quiche"; Huehuetenango. Central
and southern Mexico. Figure 42.
Plants erect, a meter high or less, or sometimes procumbent, the stems usually
several, simple or branched above, brown; leaves oblong or ovate-oblong, sessile,
1-3 mm. long, 3-15 mm. broad, obtuse, rounded or cordate at the base; pedicels
1-3 mm. long, bracteolate at the base; calyx 8-12 mm. long, purplish; flowers
dimorphous, the dolichostylous ones with stamens two-thirds as long as the calyx,
the style conspicuously exserted, the brachystylous ones with stamens about
reaching the apex of the calyx, the style reaching only to the middle of the calyx;
petals purple, sometimes 1 cm. long.
FIELDIANA: BOTANY, VOLUME 24
FIG. 42. Lythrum vulneraria. A, Portion of stem; X IK- B, Leaves and
flower; X 3. C, Flower dissected; X 5.
There are included here specimens collected by Salvin and by
Donnell-Smith that were determined as L. maritima HBK. by the
Annual or perennial herbs, growing in wet soil or in water, glabrous; leaves
opposite or verticillate, rarely alternate, sessile or subsessile; flowers regular,
3-6-parted, sometimes dimorphous, small, solitary in the leaf axils, usually sessile,
sometimes in terminal spikes or racemes, 2-bracteolate; calyx campanulate, 3-6-
lobate, usually not nerved, with or without appendages; petals persistent or
caducous, sometimes none; stamens 1-6, inserted upon the calyx lobes; ovary
sessile or substipitate, incompletely 2-4-celled, the ovules few or numerous; style
elongate or none; capsule septicidally 2-4-valvate, cartilaginous, the walls densely
transverse-striate; seeds minute.
Species about 20, chiefly in tropical regions of both hemispheres,
mostly in tropical Asia and Africa, only 3 in America.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA
Calyx not appendaged R. mexicana.
Calyx with tooth-like appendages between the teeth.
Appendages equaling or shorter than the calyx teeth R. ramosior.
Appendages 3 times as long as the calyx teeth R. dentifera.
Rotala dentifera (Gray) Koehne, Bot. Jahrb. 1: 161. 1880.
Ammannia dentifera Gray, PI. Wright. 2:55. 1853.
FIG. 43. Rotala dentifera. A, Branch; X 1. B, Inflorescence and leaves;
X 1 1 A- C and D, Hypanthium, natural position and dissected; X 5.
Wet banks or meadows, often about seasonal ponds, 250 meters
or less; Izabal; Zacapa; Retalhuleu. Mexico; British Honduras; El
Salvador; Nicaragua; Panama. Figure 43.
Plants annual or perhaps sometimes more enduring, erect or procumbent,
often much branched from the base, somewhat succulent, the stems 40 cm. long
or less; leaves opposite, oblanceolate or linear-oblanceolate, 1-4 cm. long, 3-7 mm.
broad, obtuse, attenuate to the base, sessile or short-petiolate, 1-nerved; bractlets
mostly longer than the calyx; calyx about 3-4 mm. long, the appendages twice
as long as the teeth or usually longer; petals pink, little longer than the calyx
teeth; capsule almost wholly enclosed in the calyx.
Rotala mexicana Cham. & Schlecht. Linnaea 5: 567. 1830.
260 FIELDIANA: BOTANY, VOLUME 24
In water in marshy meadows or on mud, 900-1,800 meters; Chi-
quimula; Jutiapa. Mexico; Honduras; Panama; Cuba; South Amer-
ica; Africa and Asia.
Plants repent or aquatic and partly or wholly submerged, the stems very
slender, branched; leaves in whorls of 3-5 or sometimes opposite, linear, 3-13 mm.
long, or the emersed leaves lanceolate or oblong, obtuse; flowers usually 4-5-parted,
the bractlets equaling or shorter than the calyx; calyx scarcely 1 mm. long, in
fruit semiglobose; stamens 2-3; capsule globose, 2-3-valvate.
Rotala ramosior (L.) Koehne in Mart. Fl. Bras. 13, pt. 2: 194.
1877. Ammannia ramosior L. Sp. PL 120. 1753.
In marshes or ditches, often at the edges of swamps or on sandy
stream beds, 200-1,000 meters; Zacapa; Jutiapa; Retalhuleu; Quiche".
United States; Mexico; El Salvador; Honduras; Nicaragua; Panama;
West Indies; South America; Philippine Islands.
Plants annual or perhaps sometimes more enduring, erect or procumbent,
often much branched, the stems mostly 30 cm. long or shorter; leaves opposite,
oblanceolate or linear-oblanceolate, 1-4 cm. long, obtuse, 1-nerved, attenuate
to the sessile or subsessile base; bractlets equaling or shorter than the calyx;
calyx in anthesis 2.5-3 mm. long, in fruit 4-5 mm. long, the appendages equaling
or shorter than the teeth; petals equaling or slightly longer than the teeth, pink
or white; capsule 3-4-valvate.
PUNICACEAE. Pomegranate Family
Shrubs or small trees, the branchlets sub terete, often spinescent; leaves
opposite or subopposite, often fasciculate, entire; flowers perfect and regular,
short-pedicellate, axillary, solitary or subfasciculate, large, red; calyx persistent,
thick-coriaceous, the tube adnate below to the ovary, turbinate, ampliate above
the ovary, 5-7-lobate; petals 5-7, inserted in the throat of the calyx, lanceolate,
corrugate; stamens very numerous, inserted in numerous series in the calyx throat,
the filaments filiform, incurved, the anthers ovate, versatile; ovary inferior, many-
celled, the cells biseriately superposed; style filiform, flexuous, swollen at the base,
the stigma capitate; ovules multiseriately crowded on the placentae, these adnate
to the septa and to the walls of the cells; fruit baccate, inferior, globose, large,
crowned by the persistent calyx limb, the cells many-seeded, the septa mem-
branaceous; seeds large, angulate, the testa coriaceous, imbedded in juicy pulp;
cotyledons foliaceous, spirally convolute, biauriculate at the base, the radicle very
A single genus.
PUNICA L. Pomegranate
The genus consists of a single species. The generic name is de-
rived from the Latin punicus, that is, Carthaginian, in reference to
ancient Carthage, the present-day Tunis.
STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 261
Punica Granatum L. Granado (the plant); granada (fruit);
Native of the Mediterranean region, but cultivated for its fruit
in most tropical and subtropical regions, or even in warm-temperate
lands. Planted commonly in Guatemala, at almost all elevations
except the highest, but only in small quantities.
A shrub or small tree, 6 meters high or less, usually branched from the ground,
sometimes with a short trunk, the bark thin, brownish gray; leaves short-petiolate,
elliptic to oblong or oblanceolate, 2-6 cm. long, obtuse, attenuate at the base,
glabrous; petals obovate to suborbicular, 1.5-2.5 cm. long, bright red; fruit 5-10
cm. in diameter, the pulp white or pink.
The pomegranate apparently is little esteemed in Central Amer-
ica, being unable to compete in quality with numerous excellent trop-
ical and temperate fruits. The many large seeds are an objectionable
feature. The wood is hard, close-grained, and light yellow. It has
been used as a substitute for boxwood (Buxus) in making engravings.
The bark and the rind of the fruit are astringent, and in some regions
are utilized for tanning and dyeing leather. The bark of the stem
and root its active properties due to an alkaloid pelletierine is an
efficient vermifuge, especially in the case of tapeworms. The large
flowers are brilliantly colored and decorative. The pomegranate is
by no means an exclusively tropical plant, for it survives winter in
the United States as far north as Washington, D.C., although prob-
ably it does not produce fruit at that latitude.
LECYTHIDACEAE. Brazil-nut Family
References: H. Pittier, The Lecythidaceae of Central America,
Contr. U. S. Nat. Herb. 26: 1-14, tt. 1-12. 1927. Reinhard Knuth,
Lecythidaceae, Pflanzenreich IV. 219a: 1-146. 1939.
Large shrubs or trees, often very tall; leaves alternate, often very large,
simple, without stipules, pinnately nerved; flowers usually large and showy,
perfect, solitary or racemose; sepals 2-5; petals 4-6, adnate to the stamen tube,
imbricate in bud, sometimes none; stamens numerous, united at the base, curved
in bud; anthers versatile, dehiscent by longitudinal slits; disk within the stamens,
annular; ovary inferior, 2-20-celled, with 2 or more inverted ovules in each cell;
style simple; fruit baccate and indehiscent or a pyxis, then often ligneous and
opening by an apical lid; seeds large or very large, often oily.
About 15 genera in tropical America, mostly in Amazonian South
America. Six genera are known from southern Central America,
chiefly in Panama. Most important member of the family is the
Brazil-nut, Bertholletia excelsa Humb. & Bonpl., well known for its
262 FIELDIANA: BOTANY, VOLUME 24
richly flavored nuts, which sometimes reach the shops of Guatemala
and other parts of Central America. The tree is a native of the Ama-
zonian forests and is cultivated at Lancetilla, in Honduras, and per-
haps elsewhere in Central America.
Medium-sized trees, usually with few thick branches; leaves very large, narrow,
generally crowded at the ends of the branches, entire or sinuate-dentate; flowers
large, borne on the trunk, pedicellate or subsessile, fasciculate; calyx tube turbinate,
not produced above the ovary, the limb cupular or cyathiform, at first subentire,
finally irregularly cleft into 2-4 lobes; petals usually 4, spreading; stamens very
numerous, inserted in numerous series on a thick subcupular disk, the inner ones
smaller, the filaments thick, connivent and involute; anthers small, the cells
distinct; ovary 4-celled; style none or short and conic, the 4 stigmas cruciately
radiate; ovules 2-4 in each cell, pendulous; fruit fleshy, more or less ovoid, crowned
by the calyx limb; seeds usually 1, pendulous, with a thick testa.
About 10 species in tropical America. The wood is yellow, of
medium weight and density, rather coarse- textured, easy to work,
not durable. Apparently little or no use is made of it.
Calyx lobes 5 mm. long; leaves with apex rounded or acuminate, margins not
glandular G. integrifolia.
Calyx lobes 3-4 mm. long; leaves long-acuminate with glandular margins.
Grias Gentlei Lundell, Wrightia 2: 122. 1961. Bombowood; wild
Known only from the type collection from British Honduras,
"Tree, 20 cm. in diam., glabrous. Leaves large, sessile, narrowly oblanceo-
late, up to 80 cm. long, 20 cm. wide, with glands along the subentire margin,
these becoming reddish-black with age; apex attenuate-acuminate, base attenuate
and cuneate with age, essentially spathulate, the midrib thick and prominent on
both surfaces, the lateral nerves 20 to 30. Flowers usually 3 to 7, fasciculate on
old wood, the basal bracts ovate-deltoid, 2 to 3 mm. long, acute. Pedicels, including
hypanthium, scarcely 1 cm. long, glabrous. Calyx entire, or essentially so in bud,
splitting at anthesis into two or more segments 3 to 4 mm. long. Petals 4, some-
times 5, thick, glabrous, pellucid-punctate, inaequilateral, asymmetrically elliptical,
up to 18 mm. long, 12 mm. wide, rounded at apex. Androecium about 8 mm.
long, the stamens numerous, the anthers about 0.5 mm. long, longitudinally dehis-
cent, scarcely thicker than the filaments. Ovary 4-celled, stigma 4-lobed."
We have not seen material of this species. Dr. Lundell, whose
description appears above, separates it from G. integrifolia on the
basis of its "smaller calyx and receptacle, long acuminate leaf blades
and glandular margins, fewer leaf veins."
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 263
Grias integrifolia (Standl.) Knuth, Pflanzenreich IV. 219a: 30.
1939. Gustavia integrifolia Standl. Field Mus. Bot. 4: 240. 1929.
Wet mixed forest, often in wooded swamps, at or little above sea
level; Izabal. Mexico; British Honduras; Honduras; Nicaragua (type
from Bragman's Bluff, Englesing 225).
A tree 4-10 meters, with few thick branches, or sometimes unbranched, the
trunk 15 cm. or more in diameter; bark light brown or greenish gray, rather
smooth, separating in thin flakes; plants glabrous throughout, the young branches
very thick, densely leafy at the ends; leaves huge, sometimes a meter long and
35 cm. broad, oblong-oblanceolate or somewhat spatulate, acuminate or almost
rounded and abruptly acuminate, long-attenuate to the base, sessile or nearly
so, entire or obscurely undulate, subcoriaceous, the lateral nerves as many as 40
pairs; inflorescences short, umbelliform, 3-5-flowered, the pedicels 1 cm. long;
flowers 3 cm. broad, the receptacle turbinate, 5 mm. long; calyx lobes 2, broadly
rounded at the apex, 5 mm. long; petals 4, creamy white, obovate-oblong; fruits
large, yellowish green, fleshy.
Called "genip" in British Honduras; "jaguillo," "irayol" (Hon-
duras); "morro cimarron" (Oaxaca). It is rather strange that in
British Honduras and Honduras the tree is associated or confused
with Genipa, and in Oaxaca with Crescentia, neither of which it much
resembles. The tree is a conspicuous one because of its great bunches
of huge leaves. The youngest ones often or usually are deep purplish
red. It is stated that the Lecythidaceae do not have stipules. In this
tree the new leaves are subtended by stipule-like, deep red organs that
are oblong or lance-oblong, as much as 15 cm. long, and caducous.
If these are not stipules, we do not know what they should be called.
The North American trees of this genus are represented by few speci-
mens, and the status of the various species is rather uncertain. It is
not yet established that G. integrifolia is distinct from G. Fendleri
Seem, of Panama and Costa Rica, or they from G. cauliflora L. of
Jamaica. This tree is common in the swampy forests of the north
coast of Guatemala.
RHIZOPHORACEAE. Mangrove Family
Trees or shrubs, usually glabrous, the branchlets terete; leaves opposite and
stipulate, rarely alternate and without stipules, petiolate, coriaceous, entire,
serrulate, or sinuate-crenate; stipules interpetiolar, often elongate, caducous;
flowers mostly perfect, axillary, cymose, paniculate, spicate, or racemose, rarely
congested or solitary, small or large, bibracteolate or ebracteolate; calyx tube
more or less adnate to the ovary, rarely free, the limb produced beyond the ovary
and cleft into 3-14 lobes, these valvate and persistent; petals as many as the calyx
segments and usually shorter, inserted at the base of the calyx limb, mostly con-
cave or involute and embracing the stamens, sessile or unguiculate, emarginate,
264 FIELDIANA: BOTANY, VOLUME 24
bifid, or lacerate at the apex, rarely entire, convolute or inflexed in bud; stamens
2-4 times as many as the petals or rarely of the same number, inserted on the
margin or base of a perigynous or epigynous disk, the lobes of the disk sometimes
produced as staminodia; filaments short or elongate, sometimes capillary; anthers
short or elongate, Tsasifixed or dorsifixed, 2-celled, longitudinally dehiscent; ovary
usually inferior, 2-5-celled, rarely 6- or 3-celled, or the septa obscure and the ovary
1-celled; style simple, the stigma simple or lobate; ovules usually 2 (rarely 4 or more)
and collateral in each cell, attached to the axis above its middle; fruit usually
coriaceous, crowned by the persistent calyx limb, indehiscent or sometimes tardily
and septicidally dehiscent, 1-celled and 1-seeded or 2-5-celled and with 1-seeded
cells; seeds pendulous, the endosperm carnose or none, with or without an aril,
the testa coriaceous or membranaceous.
About 15 genera, with few species, widely distributed in tropical
regions; only the following genera in North America.
Calyx 4-parted; fruit 1-seeded; leaves very thick, obtuse Rhizophora.
Calyx campanulate, 4-5-lobate; fruit 3-seeded; leaves relatively thin, usually
Shrubs or trees, glabrous or nearly so; leaves opposite, subcoriaceous or almost
membranaceous, entire or sinuate-crenate or dentate, penninerved; stipules short,
caducous; flowers axillary, solitary or fasciculate, pedunculate or subsessile, whitish;
calyx ebracteolate, campanulate, shallowly 4-5-lobate, the lobes triangular, erect,
valvate; petals 4-5, inserted at the base of a cupular crenate disk, spatulate, un-
guiculate, fimbriate-lacerate at the apex; stamens 15-30, the filaments filiform,
the anthers oblong; ovary globose or ovoid, free, sessile, 3-4-celled, the style filiform,
the stigma 3-4-lobate; ovules 2 in each cell, pendulous; fruit ovoid, fleshy-
coriaceous, tardily 3-4- valvate, the cells 1-seeded; seeds pendulous, arillate, the
testa coriaceous; endosperm carnose, the embryo straight, the cotyledons plane.
Species perhaps 50 in tropical regions of America, Africa, Asia
and Australia; no other species in Central America.
Cassipourea guianensis Aubl. Hist. PI. Guian. Fr. 1: 529, t. 211.
1775; L. Wms. Fieldiana, Bot. 29: 369. 1961. Legnotis elliptica Sw.
Prodr. 84. 1788. Cassipourea elliptica Poir. Encycl. suppl. 2, 131.
1811. C. podantha Standl. Field Mus. Bot. 4: 241. 1929 (type from
Panama). C. macrodonta Standl. I.e. 242 (type from Panama). C.
belizensis Lundell, Bull. Torr. Bot. Club 66: 598. 1939 (type from
British Honduras, Gentle 2749).
Wet forest from sea level up to some 2,000 meters; Izabal; Suchi-
tepe"quez. Mexico; Central America except El Salvador; Panama;
West Indies; South America to Brazil and Peru. Figure 44.
A shrub or tree, sometimes 18 m. tall with a trunk 35-45 cm. in diameter
but usually lower, the branchlets strigillose or glabrate; leaves on petioles 6-8
STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA
mm. long, oblong-elliptic to lance-oblong, rarely rather broadly elliptic or ovate-
elliptic, mostly 8-12 cm. long and 3.5-4.5 cm. broad, gradually or abruptly acumi-
nate or long-acuminate, gradually or abruptly narrowed at the base and acute,
at first sparsely strigillose but in age glabrous, entire or obsoletely sinuate-serrate,
the lateral nerves slender, 6-7 pairs; pedicels solitary or fasciculate and few,
FIG. 44. Cassipourea guianensis. A, Habit; X Yz- B, Flower; X 4. C, Flower
partly dissected; X 5. D, Fruit; X 4.
sessile to 3-4 mm. long, strigillose; calyx glabrous, 3.5-4 mm. long, abruptly
contracted at the base into a short thick stipe, the lobes acute, sericeous within;
petals white, densely short-pilose, deeply laciniate; ovary sericeous or glabrate;
style sericeous, exserted from the calyx; fruit oval or obovoid, 7-8 mm. long,
rounded at the apex, glabrate.
Called "waterwood" in British Honduras and "goatwood" in
Panama. The thick sap wood is yellowish, the heartwood pale brown ;
moderately hard, heavy, tough, strong, splintery, and rather fine-
textured; finishes smoothly and is fairly durable. In British Hon-
duras it is used for railway crossties and house frames.
266 FIELDIANA: BOTANY, VOLUME 24
RHIZOPHORA L. Mangrove
Reference: F. M. Salvoza, Rhizophora, Nat. & Appl. Sci. Bull.
Univ. Philippines 5: 179-237. 1936.
Glabrous trees with thick branchlets; leaves opposite, petiolate, thick-
coriaceous, entire; stipules large, interpetiolar, caducous; peduncles axillary,
dichotomously or trichotomously branched, few-flowered, the flowers large, cori-
aceous, sessile or pedicellate; calyx subtended by 2 bractlets, these connate into
a cupule, the calyx tube short, adnate at the base to the ovary, the limb 4-parted,
the lobes lanceolate, valvate; petals 4, inserted at the base of a fleshy entire disk;
stamens 8-12, inserted with the petals, the filaments short; anthers elongate,
acuminate, connivent, at first multilocellate, finally bivalvate; ovary semi-inferior,
2-celled, produced above the calyx as a fleshy cone; style conic at the base, subulate,
the stigma bidentate; ovules 2 in each cell; fruit coriaceous, ovoid or obconic,
surrounded above the base by the reflexed calyx limb, 1-celled, 1-seeded; seed
pendulous, without endosperm, the radicle elongate-clavate, perforating the apex
of the fruit upon the tree and then falling upright into the mud.
Perhaps three or more species widely distributed along tropical
seashores of the world. One other, R. samoensis (Hochr.) Salvoza,
is said to occur from Mexico to perhaps Ecuador or beyond along the
Pacific coast of America, then out into Oceania. We have not been
able to distinguish it among our material. Still another species is re-
corded from Panama, R. Harrisonii Leechman (R. brevistyla Salvoza).
Rhizophora Mangle L. Sp. PI. 443. 1753. Mangle; mangle
Abundant along both seacoasts, at least in many localities, often
forming very dense and extensive stands, usually in association with
Conocarpus, Laguncularia, and Avicennia. Southern Florida; from
Baja California and Tamaulipas southward in Mexico, and along the
whole Central American coast; West Indies; South America; Oceania.
A tree, sometimes 25 meters tall but usually smaller, the trunk rarely a meter
in diameter, the bark thin, brownish gray, shallowly furrowed, red within; leaves
petiolate, very thick and leathery, obovate or elliptic, 5-15 cm. long, obtuse, entire,
deep green above, paler beneath, the nerves obsolete; stipules 2.5-4 cm. long;
peduncles mostly 2-3-flowered; calyx 1 cm. long; petals yellow, 7-8 mm. long, vil-
lous inside, chiefly below the apex; stamens 8, about 5 mm. long; fruit 2.5-3.5 cm.
Called "red mangrove" in British Honduras; "tapche," "tabche"
(Yucatan, Maya) . The mangrove and its associates form an impor-
tant species association mangrove swamp that characterizes many
parts of the shore line of all tropical America. These swamps, often
reaching to the water but sometimes separated by sandbars, contain
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 267
FIG. 45. Rhizophora Mangle. A, Tip of a branch; X H- B, Flower; about
X 2. C, Longitudinal section of flower; about X 2. D, Sepal, inner face; about
X 2. E, Petal; about X 2. F, Anther, inner face; about X 2. G, Fruit, nearly
mature; X 3 A-
but little vegetation other than the several "mangroves," although
a few other shrubs and occasionally some herbaceous plants often are
associated with them. Their branches often bear a small number of
epiphytic plants. These trees are confined to salt or brackish water,
and the swamps usually are flooded at high tide. Rhizophora is espe-
cially adapted to an aquatic habitat by its large hard stilt or prop
roots that rise far above the soil and are somewhat bowed out, like
an arc of a circle. Mangrove swamps are especially well developed
about the mouths of streams, whence they spread rapidly seaward,
taking advantage of the silt and debris lodged there by the streams.
In this manner they are often important agents in extending the land
area. The seeds often take root on small islets or in shoals, where
they form small islands that gradually increase their area. The floor
of these mangrove thickets is one vast slimy tangle of prop roots over
which it is all but impossible to make one's way. The roots are often
covered with oysters and other marine animals, and are much fre-
quented by spider crabs, spiders, and large grasshoppers. Seen from
within, nothing could be less attractive than a mangrove forest.
268 FIELDIANA: BOTANY, VOLUME 24
Viewed from a distance, however, it is beautiful, because of its per-
manent fresh green coloring. Entering by ship the bay at Puerto
Barrios, one has a comprehensive view of large areas of such swamps.
The wood is dull red or reddish brown, sometimes purplish, uni-
form or with darker stripes; sap wood rather thick and grayish; very
hard and heavy, the specific gravity about 1.15; grain variable, from
straight to very irregular, fine-textured ; hard to cut, rather harsh and
splintery, takes a good polish, is strong and durable. The timber is
used in some regions for rafters, beams, knees and ribs of boats, and
miscellaneous construction, also for posts, piling, and railway ties.
In Central America it is much used for charcoal, that obtained from
mangrove being said to be the best of all for the kitchen. The bark
contains 20-30 per cent of tannin and is much used locally for tanning
skins, especially where oak bark is not available. Large amounts of
the bark or its extract are exported from tropical America to the
United States and Europe. The young shoots are used for dyeing;
they give red, olive, brown, or slate colors, according to the salts used
in association with them.
The method of propagation is peculiar. While still attached to
the tree, the seed develops a radicle about twice as thick as a lead
pencil and 30-60 cm. long, which when detached falls like a dart and
sticks upright in the mud, ready to put forth leaves and roots. If
the seeds are carried away by currents they float upright until they
reach a lodging place. Oviedo stated that the fruits, perhaps the
radicles, were sometimes eaten by the Indians, "when they can find no
better fruit (for it is somewhat bitter), and they say it is wholesome."
Trees or shrubs, often scandent, unarmed or bearing spines, the branchlets
terete, with few or no lenticels, the pubescence sometimes lepidote; leaves oppo-
site, alternate, or rarely verticillate, coriaceous or membranaceous, simple, petio-
late, entire; stipules none; flowers mostly perfect and spicate or racemose, rarely
paniculate or capitate, small and green or sometimes rather large, showy, and
bright-colored, bracteate; calyx tube adnate to the ovary, terete or angulate,
sometimes tubular, the limb with 4-5 lobes or segments, usually campanulate,
persistent or deciduous, the lobes valvate in bud; petals none or 4-5, imbricate or
valvate; stamens 4-5 or 8-10, inserted on the limb or base of the calyx, sometimes
biseriate, the filaments subulate or filiform, inflexed in bud; anthers small, versatile,
didymous, sagittate, or oblong, dehiscent by longitudinal slits; epigynous disk
sometimes present, lobate; ovary wholly adnate to the calyx, 1-celled; style simple,
filiform, the stigma simple, acute or obtuse; ovules 2-6, usually suspended from
the apex of the cell on elongate funicles; fruit coriaceous, chartaceous, or drupa-
ceous, often 4-5-winged, sometimes crowned by the persistent calyx limb, inde-
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 269
hiscent or rarely dehiscent for its whole length, the stone osseous or crustaceous;
seed pendulous, usually elongate and sulcate, the testa coriaceous or membrana-
ceous; endosperm none; cotyledons convolute or plicate, usually carnose and oily,
the radicle small, superior.
About 15 genera, widely dispersed in tropical regions. No other
genus is represented in Central America. Buchenavia is in Panama.
Flowers in dense globose cone-like heads; leaves alternate Conocarpus.
Flowers spicate or racemose.
Leaves alternate; petals none; trees.
Calyx limb deciduous; branches unarmed; fruit often winged .... Terminalia.
Calyx limb persistent; branches often armed with spines; fruit not winged.
Leaves opposite; petals present.
Calyx limb persistent; fruit obscurely triangulate; erect trees or shrubs; leaves
thick and fleshy Laguncularia.
Calyx limb deciduous; fruit with 4-6 wings or acute angles; plants scandent;
leaves not fleshy.
Calyx tube short, constricted above the ovary; native plants. .Combretum,
Calyx tube very slender, elongate, tubular, not constricted above the ovary;
cultivated plants Quisqualis.
Trees, the branches or branchlets sometimes armed with spines; leaves alter-
nate, usually crowded at the somewhat swollen apices of the branchlets; flowers
perfect or rarely staminate; inflorescence spicate, axillary; calyx campanulate,
shallowly 5-dentate, persistent; petals none; stamens 10, biseriate, the filaments
exserted, anthers versatile; fruit a fleshy-leathery drupe drawn up into a neck
and crowned with the persistent calyx.
A small genus with 3 or 4 more species, one in Mexico, the others
in the West Indies and South America.
Leaves mostly 3-6 cm. broad, coriaceous, glabrous or essentially so, the lateral
nerves not very conspicuous below; flowers and fruit sericeous. . . .B. buceras.
Leaves mostly 6-11 cm. broad, membranaceous to subcoriaceous, copiously pubes-
cent beneath, at least when young, the lateral nerves conspicuous below;
flowers and fruits loosely tomentose B. macrostachya.
Bucida buceras L. Syst. Nat. ed. 10. 1025. 1759. Cacho de tow;
pucte, pocte (Pete"n, Maya).
Wet forest or thickets, 1,000 meters or usually lower; Pet&i; Alta
Verapaz; Izabal; Huehuetenango. Southern Florida; West Indies;
Mexico; British Honduras; Honduras; Panama.
A tree, usually 9-15 meters high, the trunk often 30 cm. or more in diameter,
sometimes as much as a meter, the bark gray, scaly, the young branchlets sericeous,
FIELDIANA: BOTANY, VOLUME 24
soon glabrate, the branches often armed with stout brown spines 2-3 cm. long;
leaves clustered at the ends of the branchlets, crowded, slender-petiolate, obovate
to oval, 3-9 cm. long, very obtuse to rounded and emarginate at the apex, narrowed
to the base, glabrous in age, when very young somewhat sericeous; flowers whitish;
FIG. 46. Bucida macrostachya. A, Branch with leaves and inflorescences;
H- B, Flower; X 3. C, Fruit; X 3. D, Base of leaf showing glands; X 1H-
spikes pedunculate, slender, sericeous, 3-10 cm. long, usually interrupted; calyx
lobes triangular, acute; fruit ovoid, 8 mm. long, sericeous.
Called "bullet tree" and "bully tree" in British Honduras. The
wood is hard, close-grained, yellowish brown, with a specific gravity
of about 1.04. In some parts of the range it is utilized for poles,
posts, cross-ties, piling, and in general carpentry and construction
where strength and durability are important. The bark is some-
times used for tanning.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 271
Bucida macrostachya Standl. Field Mus. Bot. 4: 240. 1929.
B. megaphylla Exell, Journ. Bot. 68: 244. 1930 (type from Mexico).
Almendro de cerro; roble.
Dry brushy rocky hillsides, 200-700 meters; El Progreso (type
from El Rancho, W. A. Kellerman 7744); Zacapa; Chiquimula.
Southern Mexico; Honduras. Figure 46.
A tree of 5-9 meters, or perhaps sometimes larger, with a short trunk and dense
broad crown, the branchlets usually much thickened at the apex and densely leafy
there; petioles rather slender, mostly 2-4.5 cm. long, densely sericeous or tomen-
tose; leaf blades elliptic or obovate-elliptic, mostly 12-25 cm. long, obtuse to
broadly rounded at the apex, sometimes apiculate, cuneately narrowed to the base,
glabrous above or sparsely pilose on the nerves, densely whitish-puncticulate, be-
neath rather densely pilose with subappressed or spreading hairs, the lateral nerves
about 16 pairs, very prominent, ascending at a narrow angle; spikes numerous,
pedunculate, with the peduncle 19 cm. long or less, very densely flowered, the
flowers green or whitish, the whole inflorescence densely tomentose with lax, mostly
spreading hairs; calyx more or less persistent after anthesis, broadly campanulate,
thin, 3 mm. broad; fruit ovoid, somewhat costate, fulvous-tomentose, 5-6 mm.
long; stamens exserted.
This is a very common small tree on the dry rocky hills between
the cities of Zacapa and Chiquimula, where it often is conspicuous.
The leaves are shed during the dry season. It is noteworthy for the
swollen leaf-bearing tips of the branchlets, which are about 1.5 cm.
thick, much thicker than the leafless portion immediately below.
As remarked by Exell in describing B. megaphylla, these swollen
tips look as if they might be inhabited by ants, but apparently they
Usually woody vines, the pubescence of lepidote scales or of simple hairs;
leaves opposite, rarely verticillate, petiolate, commonly membranaceous; flowers
perfect or some of them sometimes staminate only; inflorescence often secund,
simple or compound; calyx campanulate, 4 (5) -lobate, deciduous; petals 4 (5),
small, inserted between calyx lobes and falling with calyx; stamens 8-10, biseriate;
ovary ovoid or oblong, 4-5-angulate, constricted below the calyx; ovules 2-6; fruit
coriaceous, indehiscent, with 4-6 angles or wings, 1-seeded, the wings membra-
naceous or somewhat coriaceous.
Species about 350, in most tropical and subtropical regions of the
world. A few other species grow in southern Central America. Often
Calyx 3 mm. or less long; flowers small; inflorescence paniculate, segments not
Calyx glabrous; stems spiny C. decandrum.
272 FIELDIANA: BOTANY, VOLUME 24
Calyx puberulent or tomentose; stems unarmed C. laxum.
Calyx 4-5 mm. long or longer; flowers large; inflorescence a lateral raceme, usually
Calyx 12-14 mm. long; bracts large, subfoliaceous C. Cacoucia.
Calyx 4-6 mm. long; bracts absent or minute.
Calyx and fruit pilose to tomentose C. argenteum.
Calyx and fruit lepidote, not pilose or tomentose C. fruticosum.
Combretum argenteum Bertol. Fl. Guat. 412. 1840 (type from
Volcan de Agua, Velasquez). C. erianthum Benth. PL Hartw. 73.
1841 (type from Retalhuleu, Hartweg 526) . Peine de mico; chupamiel.
Moist or dry thickets, 600 meters or less; Zacapa; Jutiapa; Santa
Rosa; Escuintla; Guatemala; Suchitepe"quez; Retalhuleu. Mexico;
El Salvador; Honduras; Nicaragua.
A large vine with brownish or grayish stems; leaves short petiolate, oblong-
elliptic to oblong-ovate, mostly 10-15 cm. long, acute or acuminate, rounded or
obtuse at the base, glabrous above or nearly so, yellowish-lepidote beneath and
more or less puberulent or short pilose; inflorescence simple or branched, often
forming large panicles, densely pilose with short spreading yellowish hairs; flowers
usually yellowish green to bright yellow; calyx limb 5 mm. long; petals glabrous,
about equaling the calyx lobes; fruits 2 cm. long, usually deep red at maturity,
pilose or tomentose, broadly winged.
Called "chupamiel" in El Salvador. This species is very similar
to Combretum fruticosum but even at a distance is of easy separation,
because in C. argenteum the inflorescences are yellow, in C. fruticosum
various shades of red to almost gray.
Combretum Cacoucia Exell in Sandwith, Kew Bull. 1931: 469.
1931. Cacoucia coccinea Aubl. PI. Guian. 450, t. 179. 1775. Termi-
nalia Cacoucia Baill. Hist. PL 6: 275. 1877, nom. illegit. C. coccineum
Engl. & Diels in Engler, Monogr. Afr. Pflanzenfam. 3: 110, 112.
1899, not Lam. 1785.
Wet thickets, usually in swamps, often in Manicaria swamps, at
or near sea level; Izabal; possibly also Baja Verapaz and Chiquimula.
British Honduras; Honduras; Nicaragua; Panama; southward to
A large coarse vine, sometimes suberect; leaves on very short petioles, broadly
elliptic to oblong-elliptic, sometimes 19 cm. long and 10 cm. broad but usually
smaller, acute or acuminate, cordate at the base, almost glabrous; flowers large for
the genus, flame-red, in stout, dense, terminal and axillary spikes 50 cm. long or
shorter, the bracts linear-lanceolate, large, green; calyx densely sericeous, 5-lobate;
petals acute, exceeding the calyx lobes; stamens long-exserted; fruit 5-angulate,
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 273
The vine is an exceptionally showy one because of its large, spire-
like, gorgeously colored flower spikes. It is plentiful about Puerto
Barrios, chiefly in swamps, and probably occurs along the Atlantic
coast throughout Central America. Exell states (Flora of Surinam)
that the seeds are "poisonous, used for killing bats," but he does not
mention the mode of administering them.
Combretum decandrum Jacq. Enum. 19. 1760. C. Palmeri
Rose, Contr. U. S. Nat. Herb. 5: 136. 1897. C. nicoyanum Pittier,
Contr. U. S. Nat. Herb. 17: 247. 1917 (type from Costa Rica, Ton-
Dry or wet thickets, 300 meters or less; Chiquimula; Santa Rosa;
Escuintla; Retalhuleu. Mexico; El Salvador; Costa Rica; Panama;
northern South America.
A large vine with brown branches, these short-pilose or puberulent, usually
armed with stiff hard spines; leaves short-petiolate, membranaceous, elliptic or
oblong-obovate, mostly 5-13 cm. long, obtuse to short-acuminate, rounded at the
base, glabrate above, pilose beneath along the costa and in the nerve axils; spikes
lax, the rachis puberulent, forming small or large and much-branched panicles,
the flowers 4-5-parted, white, sweet-scented; petals 2 mm. long, exceeding the
calyx; fruit suborbicular, 1.5 cm. long, glabrous, green, the wings very thin.
Combretum fruticosum (Loefl.) Stuntz, U. S. Dept. Agr. Bur.
PI. Ind. Seed & PI. Imp. No. 31: 86. 1914; L. Wms. Fieldiana, Bot.
29: 370. 1961. Gaura fruticosa Loefl. Inter Hispan. 248. 1758. Com-
bretum secundum Jacq. Enum. 19. 1760. C. farinosum HBK. Nov.
Gen. & Sp. 6: 110. 1823. C. Warszewiczianum Eichler in Mart. Fl.
Bras. 12, pt. 2: 110. 1867. C. Benthamianum Van Heurck & Muell.-
Arg. in Van Huerck, Obs. Bot. 220. 1871 (type from Bay of Fonseca,
Honduras). C. farinosum var. phaenopetalum Donn.-Sm. Bot. Gaz.
23: 7. 1897 (type from Nenton, Huehuetenango, E. W. Nelson 3534).
C. superbum Pittier, Contr. U. S. Nat. Herb. 18: 242. 1917. C. poly-
stachyum Pittier, I.e. 243. C. phaenopetalum Pittier, I.e. 243. C. lepi-
dopetalum Pittier, I.e. 245. Bejuco de cepillo (Pete"n); chupamiel;
Dry or wet thickets or forest, 1,200 meters or less; Pete*n; Alta
Verapaz; Izabal; El Progreso; Baja Verapaz; Zacapa; Chiquimula;
Jutiapa; Jalapa; Santa Rosa; Escuintla; Guatemala; Sacatep^quez;
Suchitepe"quez; Retalhuleu; Huehuetenango. Western and southern
Mexico; British Honduras to El Salvador and Panama; probably
northwestern South America. Figure 47.
FIELDIANA: BOTANY, VOLUME 24
FIG. 47. Combretum fruticosum. A, Portion of stem and inflorescence; X 1.
B, Individual flower; X 3. C, Hypanthium, dissected; X 4. D, Fruit; X 1^.
E, Petal; X 5.
A small or often large vine, climbing over trees, unarmed; leaves short-petio-
late, broadly oval to elliptic-oblong, 5-15 cm. long, obtuse or short-acuminate,
acute or obtuse at the base, lustrous above and glabrous or nearly so, densely lepi-
dote beneath; flower spikes very thick and dense, secund, usually paniculate;
flowers sweet-scented, usually blood-red to orange-red; petals 1.5-2 mm. long,
obtuse or acute; stamens very long and exserted, red; fruit 2 cm. long, densely
lepidote, broadly winged, usually dark red.
Known in El Salvador by the names "chupamiel," "peineta,"
"chupamiel de peineta," and "chupachupa;" called "tietie" and
"curassow comb" in British Honduras. The plant is a common
and characteristic one of the forest and thickets of the Pacific low-
lands. The showy flowers, full of nectar, are much visited by insects
and hummingbirds. It is said that the cut stem yields a considerable
amount of sap that may be drunk when water is lacking. In Mexico
the branches are used for weaving coarse baskets, and generally they
are employed as a substitute for rope, for tying firewood and other
temporary uses. In the dry lower Motagua Valley the vine is in
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 275
flower in late March. Material referred to C. fruticosum is rather
variable in size of flowers and other characters, but not remarkably
so. We are quite unable to separate most of the species of this group
maintained by Pittier (Contr. U. S. Nat. Herb. 18: 241. 1917).
Combretum laxutn Jacq. Enum. PL Carib. 19. 1760. C. mexi-
canum Humb. & Bonpl. PI. Aequin. 2: 159, t. 132. 1809 (type from
Acapulco, Mexico). C. epiphyticum Pittier, Contr. U. S. Nat. Herb.
17: 247. 1917 (type from Panama, Pittier 6819).
Wet forests or thickets at or little above sea level, sometimes in
mangrove swamps; Pete"n; Alta Verapaz; Izabal; Escuintla; San
Marcos. Mexico; British Honduras to Panama; West Indies; south-
ward to Argentina.
A large vine with tough, brown or blackish stems; leaves short-petiolate, ob-
long or lance-oblong, mostly 11-20 cm. long, glabrous or nearly so, scarcely at all
lepidote, subcoriaceous, acute or acuminate, obtuse or rounded at the base; flowers
creamy white, fragrant, 4-parted, usually in large panicles, the spikes mostly dense;
calyx finely pubescent, rarely glabrate; petals slightly exceeding the calyx lobes,
glabrous; stamens exserted; fruit oblong to suborbicular, 2 cm. long, reddish green
or dark red, broadly winged or sometimes only angulate, glabrate, usually sparsely
sericeous at first, not lepidote.
Called "tamborillo" in Chiapas, Mexico.
Shrubs or trees, glabrous or sericeous; leaves alternate, coriaceous, short-petio-
late, biglandular at the base; flowers minute, in small dense cone-like heads, these
paniculate at the ends of the branches; calyx tube compressed, truncate, not pro-
duced above the ovary, the limb urceolate, 5-fid, deciduous; petals none; stamens 5,
the filaments filiform, exserted, the anthers small, cordate; ovary 1-celled, the style
short, subulate, villous, the stigma simple; ovules 2, pendulous from the apex of
the cell; fruit small, obcordate, angulate, 1-seeded, the fruits densely imbricate,
with corky pericarp.
Two species in mangrove swamps of the tropics of America and
West Africa; only the following in North America.
Conocarpus erecta L. Sp. PI. 176. 1753. Mangle bianco.
Abundant in mangrove swamps or beach thickets along both
coasts; Izabal; Escuintla; Retalhuleu; San Marcos; doubtless in all
the coast departments. Mexico; British Honduras to Panama. Gen-
erally distributed on seacoasts of tropical America and in West Africa.
Variable in size, sometimes a prostrate shrub but usually erect and becoming
a tree of 20 meters with a trunk 80 cm. in diameter, the bark dark brown, fissured
276 FIELDIANA: BOTANY, VOLUME 24
into irregular ridges and thin scales; leaves obovate to elliptic or oval, 2-10 cm.
long, obtuse or acute at each end, glabrous or sericeous, the petiole with 2 glands
on the upper surface at the base of the blade; flowers greenish, the heads 1 cm. or
less in diameter; fruit purplish green, cone-like.
Called "buttonwood" and "button-bush" in British Honduras;
"botoncillo" (El Salvador) ; "canche" (snake tree), "taabche," "tabche"
(Yucatan, Maya); "mangle," "mangle prieto" (Yucatan). This is
one of the usual elements of the mangrove swamps of all tropical
America, growing in association with Rhizophora, Laguncularia and
Avicennia. In the mangrove swamps of San Jose", Escuintla, this is
the most abundant tree. The wood is fine-textured, hard, heavy,
and strong, its specific gravity near 1.00. Locally it is utilized for
fuel and charcoal and sometimes for construction. The bark is used
for tanning skins. The usual form of the species has glabrous or
nearly glabrous leaves. Var. sericea DC. (Prodr. 3: 16. 1828) is a
form with densely sericeous leaves. We have seen no Guatemalan
specimens, but it appears to be common in Yucatan and occurs in
LAGUNCULARIA Gaertner f.
Trees; leaves opposite, petiolate, thick-coriaceous, succulent when fresh, the
venation obsolete, biglandular at the base; flowers polygamous, in elongate axillary
spikes, sericeous, sessile; calyx tube turbinate, not produced beyond the ovary,
bibracteolate, the limb urceolate, 5-fid, persistent; petals 5, minute, caducous;
stamens 10, biseriate, the filaments subulate, included, the anthers cordate; ovary
1-celled, the style filiform, glabrous, the stigma bilobate; ovules 2, collaterally
pendulous from the apex of the cell; fruit coriaceous, crowned by the calyx limb,
elongate-obovoid, subtrigonous, 1-seeded, the angles marginate.
Two species, in tropical America and Africa. Only the following
occurs in North America.
Laguncularia racemosa (L.) Gaertn. in Gaertn. f. Fruct. 3:
209, t. 217. 1805. Conocarpus racemosa L. Syst. Nat. ed. 10. 930.
1759. Mangle Colorado; mangle chaparro; mangle bianco.
Common in mangrove swamps of both coasts, growing with Rhi-
zophora, Conocarpus, and Avicennia; Izabal; Escuintla; Retalhuleu;
San Marcos; doubtless in all the coastal departments. Southern
Mexico; British Honduras to El Salvador and Panama; southern
Florida; West Indies; South America; Africa.
A shrub or tree, sometimes 20 meters tall with a trunk 80 cm. in diameter, the
bark thin, reddish brown, fissured into long scales; leaves petiolate, oblong to oval,
mostly 3-7 cm. long, rounded or very obtuse at each end, glabrous, somewhat
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 277
tuberculate-roughened beneath when dry, the blade with 2 glands at its base;
flower spikes mostly lax and interrupted, often curved; calyx sericeous, 2-3 mm.
long; fruit drupaceous, 1.5 cm. long, 10-costate, reddish.
Called "white mangrove" in British Honduras; "cincahuite" (El
Salvador); "zacolcom" (Yucatan, Maya); "mangle bobo" (Yucatan).
The wood is hard, heavy, strong, dense, yellowish brown, its specific
gravity about 0.86. It is little used except for fuel. The bark is
stated to contain about 14 per cent tannin and is often used for tan-
Woody vines with slender branches; leaves opposite or subopposite, membra-
naceous, petiolate; flowers showy, in short, axillary and terminal spikes or racemes,
usually changing their color with age; calyx tube ovoid below, terete, produced
above the ovary into a very long, slender tube, deciduous, the limb 5-parted, the
small lobes spreading or recurved; petals 5, obtuse; stamens 10, exserted; style fili-
form; ovules 3-4; fruit dry, oblong, coriaceous, acutely 5-angulate or 5- winged,
About 17 species, in tropical Asia and Africa, one of them widely
cultivated for ornament.
Quisqualis indica L. Sp. PI. ed. 2. 556. 1762.
Cultivated in the Parque Central of Guatemala City, and doubt-
less planted elsewhere in the country. Native of tropical Asia.
Vine woody, often large; leaves short-petiolate, oblong or oblong-elliptic, about
14 cm. long and 5 cm. broad or smaller, acuminate, rounded at the base, sparsely
pilose or almost glabrous, spikes with conspicuous linear green bracts; calyx tube
sometimes 8 cm. long, very slender, finely pilose; petals white, turning pink and
red, obovate-oblong, 12 mm. long, much exceeding the calyx lobes; fruit ellipsoid,
5-costate or narrowly 5-winged, sometimes dehiscent along the angles.
Called "Santa Cecilia" and "barbudo" in El Salvador. The vine
is not a common one in Central America.
Trees, unarmed; leaves alternate or subopposite, usually crowded at the ends
of the branches, sometimes pellucid-punctate, mostly petiolate and often glandular
at the base; flowers small, sessile, perfect or polygamo-dioecious, 4-5-parted, green
or whitish, in elongate spikes; calyx tube ovoid or cylindric, constricted above the
ovary, the limb campanulate, generally deciduous; petals none; stamens 10, biseri-
ate; filaments exserted, the anthers small, didymous; style subulate, villous at the
base, the stigma simple; ovules usually 2; fruit dry or drupaceous, often winged,
the putamen coriaceous or osseous.
278 FIELDIANA: BOTANY, VOLUME 24
Species about 200, widely dispersed in tropical regions. One other
species is known from Guatemala by a sterile specimen.
Fruits angled but not winged; leaves broadly obovate, mostly 10-15 cm. broad.
Fruits prominently winged; leaves mostly 6 cm. broad or less.
Fruits 3-3.5 cm. high, with hard thick wings; lower surface of the leaves con-
spicuously reticulate T. nyssaefolia.
Fruits 2 cm. high or less.
Fruits 2-winged, the wings each about 1.5 cm. or more broad T. oblonga.
Fruits usually 5-winged, 2 wings much larger than the others, each not more
than 1 cm. broad T. amazonia.
Terminalia amazonia (J. F. Gmel.) Exell in Pulle, Fl. Surinam
3: 173. 1935. Chuncoa amazonia J. F. Gmel. in L. Syst. Nat. ed. 13,
2: 702. 1791. Gimbernatia obovata Ruiz & Pavon, Fl. Peruv. Prodr.
138. 1794. T. obovata Steud. Norn. Bot. 2: 668. 1841, not Cambess.
1829. T. excelsa Liebm. ex Hemsl. Biol. Centr. Amer. Bot. 1: 402.
1880, nomen nudum. T. Hayesii Pittier, Contr. U. S. Nat. Herb. 18:
239. 1917. Naranjo; canxun (Maya); canxan (Pete"n, Maya); que-
Wet forest or open savannas, 300 meters or less; Peten; Alta
Verapaz; Izabal; Huehuetenango(?). Southern Mexico; British Hon-
duras to Panama; southward to Brazil and Peru.
A tall tree, sometimes 40 meters high, with a tall and rather slender trunk
sometimes a meter in diameter above the often high buttresses, the bark smooth,
pale brown or brownish, often mottled, peeling off in sheets, the slender young
branchlets usually densely pilose with subappressed, yellowish or brownish hairs;
leaves mostly clustered at the tips of the branches and appearing verticillate, mem-
branaceous or subcoriaceous, obovate to oblong-oblanceolate, mostly 7-11 cm.
long, on very short petioles, rounded to acuminate at the apex, cuneate-attenuate
to the base, more or less pilose along the nerves, elsewhere glabrous or nearly so,
often with 2 marginal glands above the base; flowers yellow-green or whitish, in
elongate axillary spikes, these slender, often numerous; calyx limb 3-4 mm. broad;
disk villous; fruit only 4-5 mm. high, the 5 thin wings usually broader than high,
with two larger than the others.
Sometimes called "nargusta" in British Honduras; in Honduras
"membriUo" and "almendro." One of the most common large trees
of the whole Atlantic coast of Central America, and reported as one
of the largest and finest trees of the Pete"n savannas. It grows in
swamps, but also upon hillsides, although it does not extend high on
the slopes. The wood is lustrous light gray or yellowish, becoming
decidedly yellowish upon exposure, the injured portions red or brown;
taste slightly astringent; moderately hard and heavy, the specific
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 279
gravity 0.65-0.75; grain variable, from straight to roey or curly,
medium-textured, easy to cut, takes a good polish, strong, appar-
ently not very durable. The wood is used in the North Coast for
general construction and railroad ties, though not highly esteemed
for the latter purpose. Although not exported, Record states that
it is suitable as a medium-priced wood for furniture, interior finish,
and general carpentry. The name "guayabo" given the tree in Cen-
tral America refers to the facts that the bark is much like that of the
common guava (Psidium) . The bark on the lower part of the trunk
curls up in thin sheets that can easily be pulled loose; it often falls
off and litters the ground.
Terminalia Catappa L. Mant. PI. 128. 1767. Almendro.
Native of tropical Asia. Planted commonly as a shade tree
throughout the tierra caliente of Guatemala, and naturalized in
many or most parts of the lower regions, especially near the sea-
coast; occasionally seen above 300 meters; Pete*n; Izabal; Zacapa;
Santa Rosa; Escuintla; Suchitepe"quez; Retalhuleu; San Marcos.
A large tree, sometimes 25 meters high with a trunk a meter in diameter, but
commonly smaller, the branches rather few, conspicuously whorled and spreading;
leaves mostly clustered at the ends of the branches, obovate, 10-30 cm. long,
rounded and abruptly pointed at the apex, cuneately narrowed to the obtuse or
subcordate base, almost glabrous; flower spikes 5-15 cm. long, the pistillate flowers
on the lower part of the spike, green; fruit a woody drupe, ellipsoid, 2-edged,
4-7 cm. long, the seed 3-4 cm. long.
Called "almond" in British Honduras. The usual English name
is "Indian almond."
The wood is hard, close-grained, and red-brown; it supplies use-
ful lumber when available in quantity, but probably is not used in
Guatemala. The tree is much planted in parks, especially near the
coast, as at Puerto Barrios, San Jose", and Champerico. It tolerates
saline soil better than most trees, and endures neglect. It often is
described as un drbol agradecido, since it repays by rapid and vigorous
growth any care given it. The large thick leaves fall during the dry
months and often form a dense rustling cover over the ground. The
young leaves as well as those about to fall often are vividly tinted
with red and purple, making the foliage very conspicuous. The bark
and fruit are rich in tannin. They also yield a black dye that in India
is used for staining the teeth black. It is said that in Asia silkworms
are fed upon the leaves. The seeds, said to contain 50 per cent oil,
are edible, with a flavor suggestive of almonds (hence the name "al-
FIELDIANA: BOTANY, VOLUME 24
mendro") or filberts, and they often are eaten by Guatemalan chil-
dren, perhaps also by adults. In El Salvador they are used to give a
black dye to textiles. The tree is well known throughout the low-
lands of Central America.
FIG. 48. Terminalia oblonga. A, Branch; X %. B, Flower; X 3. C, Fruit;
XI. D, Fruit in cross section; X 1.
Terminalia nyssaefolia Britton, Bull. Torr. Bot. Club 48: 333.
At sea level; Izabal (Punta Palma, Steyermark 39814). Trinidad.
A tree as much as 20 meters high, the branches slender or stout, brownish-
pilose or glabrate; leaves short-petiolate, obovate or oblong-obovate, 7-12 cm. long,
4-6 cm. broad, abruptly acute or short-acuminate, cuneate-acute at the base, green
above, lustrous, glabrous or nearly so, paler beneath, somewhat brown-pilose when
young but soon glabrate, the veins conspicuously, elevated and closely reticulate
on both surfaces; flowers spicate, densely brown-pilose, the spikes slender-peduncu-
late, short, rather few-flowered, much interrupted; fruit glabrous or nearly so,
usually 3-3.5 cm. long, strongly compressed, the broad wings very thick and hard.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 281
The occurrence of this tree at this one isolated spot in Central
America is curious and not a natural extension of range for a Trinidad
tree, although well within the bounds of possibility. It may be that
the tree is more widely distributed along the Guatemalan coast or in
other parts of Central America, but there is also the possibility that
it may have been planted and introduced at Punta Palma.
Terminalia oblonga (R. & P.) Steud. Norn. Bot. 2: 668. 1841;
L. Wms. Fieldiana, Bot. 29: 385. 1962. Gimbernatia oblonga Ruiz
& Pavon, Syst. Veg. 274. 1798. Chuncoa oblonga Pers. Syn. PI. 1:
486. 1805. T. chiriquensis Pittier, Contr. U. S. Nat. Herb. 18: 238.
1917. Volador; guayabo.
Common in forests of the Pacific lowlands, mostly at 600 meters
or less, sometimes ascending to 1,200 meters; Santa Rosa; Escuintla;
Suchitepe"quez ; Retalhuleu; Quezaltenango; San Marcos. El Salva-
dor and Honduras to Panama; southward to Brazil. Figure 48.
A large tree, similar to T. amazonia in most details, sometimes 45 meters tall,
with a trunk 75 cm. in diameter, the trunk tall and slender, the crown relatively
small and spreading, the bark mottled like that of Platanus, the newly exposed
portions almost white; buttresses usually present but rather small; leaves usually
on longer petioles than in T. amazonia, mostly acute or acuminate, almost glabrous,
usually without glands, pellucid-punctate; flowers green, the spikes mostly longer
than the leaves, slender, interrupted, the stamens long-exserted; fruit with only
2 wings, these subcoriaceous, 2 cm. high or shorter, with each wing about 1.5-2 cm.
broad, the wings finely nerved.
This is one of the most common trees of the Pacific plains, often
forming extensive and dense stands, usually in association with other
trees. It is quite possible that T. amazonia may extend to the Pa-
cific slope of Guatemala, but we have seen no specimens that could
be referred certainly to it.
UNIVERSITY OF ILLINOIS URBANA