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Full text of "Flora of Guatemala"

L I B RAR.Y 

OF THE 

UNIVERSITY 
OF ILLINOIS 

FSo.S 



v. 24- 
pt.? 



BIOLOGY 



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p 

FLORA OF GUATEMALA 



PAUL C. STANDLEY 



AND 



LOUIS 0. WILLIAMS 



FIELDIANA: BOTANY 
VOLUME 24, PART VII, NUMBER 2 

Published by 

CHICAGO NATURAL HISTORY MUSEUM 
JANUARY 26, 1962 



FLORA OF GUATEMALA 

PART VII 



FLORA OF GUATEMALA 



PAUL C. STANDLEY 

Curator Emeritus of the Herbarium 

AND 

LOUIS 0. WILLIAMS 

Curator, Central American Botany 



FIELDIANA: BOTANY 
VOLUME 24, PART VII, NUMBER 2 

Published by 

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 



.. 

re 

v. 



. * 



CONTENTS 
Families Included in Part VII, Number 2 

PAGE PAGE 

Cactaceae. ..................... 187 Punicaceae. . 

Thymelaeaceae ................. 234 Lecythidaceae . . 

Lythraceae ..................... 239 Rhizophoraceae ....... - 263 

Combretaceae .................. 268 



LIST OF ILLUSTRATIONS 
TEXT FIGURES 

PAGE 

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 



vn 



Flora of Guatemala 1 

OPUNTIALES 

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 
it contains. 

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. 

187 



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- 
uous). 
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- 
pressed Rhipsalis. 

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 

epiphytic. 

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. 

Cephalocereus. 

Branches without a covering of white hair-like bristles at the ends. 
Flowers 2-several at each areole; flowers about 3 cm. long. 

Myrtillocactus. 
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. 

Selenicereus. 

Spines, at least the largest ones, 3-4 cm. long; flowers 4-7 cm. long. 

Nyctocereus. 
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 
America. 

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 
(Oaxaca). 

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. 

APOROCACTUS Lemaire 

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." 

CEPHALOCEREUS Pfeiffer 

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; 
Quiche". Honduras. 

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. 

194 



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. 



ECHINOCACTUS 

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. 



ECHINOPSIS 

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 
called tuna. 

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. 

E. Nelsonii. 
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. 

E. strictum. 

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- 
tral America. 

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- 




4 



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. 



199 



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. 



201 



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- 
ure 33. 

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. 



203 



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- 
ama. 

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 
pitted, black. 

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). 
Pitahaya. 

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); 
El Salvador. 

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. 
Figure 34. 

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- 



207 



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, 
black. 

About twenty species, ranging from Arizona to the northern coast 
of South America. There are other species in Mexico and Central 
America. 

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. 

Cereus Yunckeri. 

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 
(Sacapulas). 

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 
Honduras. 

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 
the markets. 

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 
segregates. 



210 FIELDIANA: BOTANY, VOLUME 24 

MAMMILLARIA Haworth 

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 
Mexican. 

Tubercles not emitting milk when cut, the milk tubes, if any, only in the stem. 

M. Ruestii. 
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 
by peddlers. 

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 
Central America. 

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 
3:227.1922. Chile. 

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 
and edible. 

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. 

MYRTILLOCACTUS Console 

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 
Valley. 

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 
small scales. 

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. 



NOPALEA Salm-Dyck 

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. 

N. cochenillifera. 
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- 
duras. 

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. 

Joints glabrous. 

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. 

0. Deamii. 

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 
edible. 

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. 
1908. 

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. 
1837. Aceituno. 

In thickets, about 1,350 meters; Baja Verapaz; Huehuetenango. 
Mexico. 

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 
and local. 

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; 
Quezaltenango; Huehuetenango. 

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 
insects. 



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: 
177. 1909. 

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. 



PERESKIA Miller 

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. 



".'A-' 




223 



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. 

P. aculeata. 
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; 
matial. 

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. 
1907. 

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. 



RHIPSALIS Gaertner 

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 
and furrowed. 

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; 
tropical Africa. 

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- 
bradura. 

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 
36. 

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. 



229 



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. 
1904. Pitajaya. 

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 
axils. 

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 
lobes white. 



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 
in Guatemala. 

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. 
Figure 37. 

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. 
85. 1819. 

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. 



233 



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. 
long. 

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. 



MYRTIFLORAE 

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. 

THYMELAEACEAE 

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 
short, superior. 

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 
small drupe. 

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. 

D. monocephala. 

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- 
chdc. 

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- 
tenango. 

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 
strigose, acute. 

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- 
mark 31758). 

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. 

238 



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; 
Zacapa. 

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 
apex. 

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- 
onymous name. 



ELAEAGNACEAE 

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. 

ADENARIA HBK. 

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. 

AMMANNIA L. 

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 
irregularly. 

Species about 20, mostly in tropical regions. Only the following 
are known in Central America. Ammannia latifolia L. is known in 
Panama. 

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 
and slender. 

CUPHEA Adanson 

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 
scription. 

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. 

C. hyssopifolia. 

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- 



neous. 



U6UUD. 

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. 

C. calophylla. 
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- 
uous. 

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; 
stamens exserted. 

This has been reported from British Honduras as C. axilliflora 
Koehne. 

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 
Honduras. 

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,' 

l.C . 



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 
in Guatemala. 

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- 
ern Mexico. 

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). 
Southern Mexico. 

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. 



248 



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: 
236. 1877. 

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, 
cuneate-oblong. 

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. 
1897. 

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; 
stamens exserted. 

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; 
ovules 13-17. 

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 
petal. 

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 
Tacand) . 

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. 
Seler 2841. 

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 

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. 

LAFOENSIA Vandelli 

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 



254 



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. 
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; 
astronomica. 

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- 
known there. 



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, 
in syn. 

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 
(fide Aguilar). 

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. 



258 



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 
monographer, Koehne. 



ROTALA L. 

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 



259 



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. 




S.H.G. 



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 
short. 

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); 
granad (Quecchi). 

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. 

GRIAS L. 

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. 

G. Gentlei. 

Grias Gentlei Lundell, Wrightia 2: 122. 1961. Bombowood; wild 
mammy. 

Known only from the type collection from British Honduras, 
Gentle 5194. 

"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 
acuminate Cassipourea. 

CASSIPOUREA Aublet 

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 



265 



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 
Colorado. 

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. 
Figure 45. 

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. 
long. 

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." 

COMBRETACEAE 

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. 

Bucida. 
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. 



BUCIDA L. 

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, 



270 



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; 




X 



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 
are not. 

COMBRETUM L. 

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 
ornamental. 

Calyx 3 mm. or less long; flowers small; inflorescence paniculate, segments not 

secund. 
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 

secund. 

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 
Brazil. 

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, 
not winged. 



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- 
duz 13503). 

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; 
peineta. 

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. 



274 



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. 
CONOCARPUS L. 

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 
Honduras. 

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- 
ning skins. 



QUISQUALIS L. 

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, 
1-seeded. 

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. 



TERMINALIA L. 

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. 

T. Catappa. 
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- 
bracho; sisin. 

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- 



280 



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. 
1922. 

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. 



Publication 946 



UNIVERSITY OF ILLINOIS URBANA