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FLORA OF GUATEMALA

PAUL C. STANDLEY

AND

JULIAN A. STEYERMARK

FIELDIANA: BOTANY

VOLUME 24, PART VI

Published by

CHICAGO NATURAL HISTORY MUSEUM

DECEMBER 27, 1949

FLORA OF GUATEMALA

PART VI

FLORA OF GUATEMALA

PAUL C. STANDLEY

Curator of the Herbarium

AND

JULIAN A. STEYERMARK

Associate Curator of the Herbarium

FIELDIANA: BOTANY

VOLUME 24, PART VI

Published by

CHICAGO NATURAL HISTORY MUSEUM

DECEMBER 27, 1949

THL LIBRARY OF THE

JAN 12 1950

Y OF i'i'NCIS

PRINTED IN THE UNITED STATES OF AMERICA

BY CHICAGO NATURAL HISTORY MUSEUM PRESS

CONTENTS

Families Included in Part VI

PAGE

Trigoniaceae 1

Vochysiaceae 2

Polygalaceae 5

Dichapetalaceae 22

Euphorbiaceae 25

Callitrichaceae 171

Buxaceae 172

Coriariaceae 174

Julianiaceae 175

Anacardiaceae 177

Cyrillaceae 195

Aquifoliaceae 196

Celastraceae 201

Hippocrateaceae 218

Staphyleaceae 223

Icacinaceae 225

Aceraceae 229

Hippocastanaceae 233

Sapindaceae 234

Sabiaceae 273

Impatientaceae 275

Rhamnaceae 277

Vitaceae 293

Tiliaceae 302

Malvaceae 324

Bombacaceae 386

Sterculiaceae 403

Saurauiaceae . . . 428

TRIGONIACEAE

Reference: Paul C. Standley, Trigoniaceae, N. Amer. Fl. 25: 297-

298. 1924.

Trees or shrubs, sometimes scandent; leaves opposite or alternate, simple,

entire, penninerved; stipules usually present, large, interpetiolar, sometimes con-

nate; flowers perfect, small, irregular, 2-bracteolate, racemose or paniculate;

sepals 5, free or connate at the base, subequal, imbricate, deciduous; petals 3 or

5, white or pink, free, subperigynous, alternate with the sepals, unequal and

somewhat papilionaceous, contorted in bud; stamens 5-12, with 2-6 of the fertile

ones united, the tube cleft on one side; filaments filiform, the anthers 4-celled,

oval, opening by longitudinal introrse slits; disk sometimes present; ovary free,

3-celled; style terminal, simple, the stigma capitate or obliquely truncate; ovules

2 or more in each cell, 2-seriate, attached to a central placenta, anatropous; fruit

capsular, 3-celled, septicidally 3-valvate, the valves separating from the central

column and themselves often separating into endocarp and epicarp; seeds 2 or

more in each cell, the testa thin, covered with long wool; endosperm carnose, the

embryo straight, the cotyledons plane, the radicle short.

Three genera, one in the Malay Peninsula, the others in tropical

America. Only one is represented in North America.

TRIGONIA Aublet

Usually woody vines; leaves opposite, short-petiolate, generally white-

tomentose beneath; stipules simple or bifid at the apex, deciduous; flowers small,

in terminal panicles or compound racemes; sepals connate at the base, unequal,

the 2 inner ones larger; petals very unequal, the posterior one largest, calcarate

or saccate, pilose in the throat of the spur, the blade reflexed, the 2 anterior ones

ascending, narrow, barbate above the base, the other 2 petals smaller, approximate,

keel-like; stamens 10 but usually only 6 of them fertile; 2-4 hypogynous glands

or a crenate crest present opposite the posterior petal; ovary attenuate to the

style, hirsute, the stigma obliquely truncate; ovules several or numerous; capsule

trigonous, usually pubescent outside and often also within; seeds several in each

cell, compressed-globose.

About 30 species, in tropical America. Two other Central Ameri-

can ones have been described from Nicaragua and Panama.

Leaves mostly 4.5-8.5 cm. long, in age glabrous beneath or nearly so, with about

5 pairs of lateral nerves; capsule 2 cm. long or shorter; petioles 5 mm. long

or shorter T. rasa.

Leaves mostly 9-14 cm. long, usually abundantly pilose beneath even in age,

with about 7 pairs of lateral nerves; capsule usually 2.5 cm. long; petioles

mostly 7-13 mm. long T. floribunda.

2 FIELDIANA: BOTANY, VOLUME 24

Trigonia floribunda Oerst. Vid. Medd. 38. 1856.

Moist or dry thickets on plains and foothills, chiefly on the

Pacific slope, 850 meters or less; Zacapa; Chiquimula; Jutiapa; Santa

Rosa; Suchitepe"quez; Retalhuleu. Chiapas; Honduras and Salvador

to Panama.

A small or large, woody vine, the slender branches densely floccose-tomentose

at first, soon glabrate; petioles 5-20 mm. long, mostly about 10 mm.; leaf blades

elliptic-oblong to lance-oblong or obovate-oblong, 1.5-6 cm. wide, usually acute,

obtuse or acute at the base, floccose-tomentose above when young but soon

glabrate and green, covered beneath at first with a very dense, white tomentum,

in age green but usually pilose, the lateral nerves 7-9 pairs; inflorescence thyrsoid-

paniculate, much exceeding the leaves, 10-25 cm. long, the pedicels 1-2 mm. long;

sepals lance-oblong, acute, 3-4 mm. long, tomentose outside; petals white, 3-4 mm.

long, the spur globose, 2 mm. long; perfect stamens 6, glabrous, shorter than the

petals; capsule obtusely trigonous, 2-3 cm. long, glabrate; seeds 6-9 in each cell,

covered with long, whitish or fulvous, silky hairs.

This is probably the plant reported from Guatemala by Hemsley

as T. guianensis Aubl., on the basis of a collection made by Bernoulli

and Cario. In T. floribunda the young leaves almost always are

covered on the lower surface with a very dense, white tomentum,

but on adult leaves scarcely a trace of the tomentum is to be seen.

Trigonia rasa Standl. & Steyerm. Field Mus. Bot. 23: 59. 1944.

Dry or moist, brushy plains or hillsides, 300 meters or some-

what higher; Santa Rosa (type collected at Rio Panal, lower slopes

of Volcan de Tecuamburro, along road between Cuilapa and Chi-

quimulilla, Standley 78584); Retalhuleu (west of Retalhuleu); en-

demic.

A woody vine with elongate branched stems, the young ones hirtellous or

puberulent; leaves subchartaceous, on petioles 2-5 mm. long, elliptic-oblong to

oblong-ovate or ovate, mostly 4.5-8.5 cm. long and 2-3.5 cm. wide, narrowly

long-acuminate, obtuse at the base, glabrous above or puberulent only on the

costa, almost glabrous beneath in age, with a few scattered straight hairs on the

costa and nerves, the lateral nerves about 5 pairs; inflorescences apparently small

and 6 cm. long or less, the fruiting pedicels 7 mm. long or shorter; capsule oblong-

ovoid, 1.5-2 cm. long, 1 cm. broad, sparsely pilose, densely covered with prominent

pale lenticels.

VOCHYSIACEAE

References: Paul C. Standley, Vochyaceae, N. Amer. Fl. 25: 301-

303. 1924; F. A. Stafleu, A Monograph of the Vochysiaceae, Rec.

Trav. Bot. Neer. 41: 398-540. 1948.

Trees or shrubs, often with resinous sap, the branchlets terete or angulate;

leaves opposite or verticillate, simple, entire, penninerved; stipules small, some-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 3

times reduced to glands or absent; flowers perfect, irregular, large or small, bi-

bracteolate, in racemes, panicles, or thyrses; sepals 5, connate at the base, im-

bricate, subequal or very unequal, deciduous, one of them often calcarate or

saccate; petals yellow, white, bluish, or purplish, free, perigynous or epigynous,

rarely 5, subequal, and imbricate in bud, usually fewer than the sepals (0-3) and

contorted or imbricate in bud; stamens inserted with the petals, typically 10 but

only 1 of them fertile; filaments cylindric or filiform, the anthers linear to oval,

4-celled, dehiscent by 2 longitudinal introrse slits; ovary free or rarely inferior,

1-3-celled; style simple, terminal, the stigmas depressed-capitate, somewhat

3-lobate; ovules 2 or more in each cell, anatropous, attached to a central placenta;

fruit usually capsular, loculicidally 3-valvate; seeds commonly winged, often pilose

or lanate; endosperm none; embryo straight, the cotyledons thin, convolute, the

radicle short, superior.

Five genera, all except the following confined to South America.

VOCHYSIA Jussieu

Trees or shrubs; leaves opposite or verticillate, coriaceous, the stipules small

or none; inflorescence terminal, thyrsoid, composed of racemosely arranged, 2-10-

flowered, scorpioid, pedunculate cymes; sepals unequal, the posterior one large

and usually produced into a spur; petals 3, rarely 1 or none, inserted in the throat

of the calyx, linear or spatulate, the anterior one commonly largest; fertile stamen

1, the filament subulate or filiform, the anther basifixed, elongate, surpassed by

the connective, this cucullate at the apex; staminodia 2 or none; ovary free, 3-

celled, attenuate to a filiform style; capsule coriaceous or ligneous, 3-angulate,

3-celled; seeds 1 in each cell, oblong, compressed, winged.

About 55 species, all except 4 in South America. One species is

known in Central America from Costa Rica and Panama, and an-

other occurs in Tabasco. The genus was first published by Aublet

under the name Vochya. By Jussieu the name was published as

Vochisia, but most authors have used the form Vochysia adopted

here.

Leaves acuminate or long-acuminate, broadest at or below the middle.

V. guatemalensis.

Leaves obtuse or rounded at the apex, usually broadest above the middle.

V. hondurensis.

Vochysia guatemalensis Bonn. Smith, Bot. Gaz. 12: 131. pi. 23.

1887. Vochya guatemalensis Standl. N. Amer. Fl. 25: 302. 1924.

Ruanchap (Quecchi).

Moist or wet forest, 350-1,500 meters; Alta Verapaz (type from

Pansamala, Tuerckheim 943); Huehuetenango. Honduras; Mexico.

A large tree, often 15 meters high or more, with pale bark, the branchlets

glabrous; stipules subulate, 3 mm. long; leaves 3-4- verticillate or the uppermost

opposite, on petioles 2-3 cm. long, oblong-lanceolate, 9-15 cm. long, 2.5-5.5 cm.

wide, rather abruptly acuminate or long-acuminate, acute or acuminate at the

4 FIELD IANA: BOTANY, VOLUME 24

base, coriaceous, glabrous; flowers bright yellow, the thyrses terminal and axillary,

forming large leafy panicles 10-18 cm. long, the rachis sparsely puberulent, the

cymes 3-4-flowered; blade of the posterior sepal 15-20 mm. long, the spur half

as long; petals oblong-obovate, the intermediate one half as long as the calyx and

4 mm. wide, the others slightly shorter and much narrower; anther glabrous, 10

mm. long.

What was presumed to be this species was noted as in flower in

April along river banks near San Pedro Carcha, where the branches

were not accessible.

Vochysia hondurensis Sprague, Kew Bull. 183. 1922. Vochya

hondurensis Standl. N. Amer. Fl. 25: 303. 1924. San Juan; San-

pedrano; Palo bayo (Pete"n); Sayuc (Pete*n, Maya); Robanchab

(Alta Verapaz).

Moist or wet forest, at or little above sea level; Pete"n; Izabal;

Alta Verapaz. Oaxaca to British Honduras; Honduras; Nicaragua;

Costa Rica.

A tree 15-25 meters tall, the crown dense, rounded or depressed, the trunk tall

and slender, much exceeding the crown, the bark smooth and grayish; branchlets

glabrous; leaves 3-4-verticillate, on slender petioles 1-2.5 cm. long, oblong or

oblanceolate-oblong, 8-15 cm. long, 2.5-5 cm. wide, rounded or obtuse at the

apex and usually emarginate, acute or acuminate at the base, coriaceous, glabrous;

flowers bright yellow, the thyrses terminal and axillary, 6-20 cm. long; blade of

the posterior sepal 15 mm. long, the spur half as long; petals obovate, about 5 mm.

long, ciliolate; anther glabrous, 6-10 mm. long; capsule narrowly oblong, deeply

3-sulcate, somewhat verrucose, acutely angulate, about 4.5 cm. long and 1.5 cm.

broad.

Known in British Honduras by the names "white mahogany,"

"yemeri," "emeri," and "emery"; called "corpus" and "corpo" in

Oaxaca; known in Nicaragua as "barba chele." The tree is a con-

spicuous and strikingly handsome one when covered with its brilliant

yellow flowers. It is abundant in many places on the hills of the

Quirigua region, as well as elsewhere in Izabal, where it often stands

high above the surrounding trees. It is common about Puerto

Barrios. The wood is reddish brown or pale brown with a pinkish

hue and a golden sub-luster, although the surface may appear dull

and mealy; light in weight, fairly tough, coarse-textured, inclined

to be gritty and hard on tools when dry, holds its place well when

manufactured; fairly resistant to decay and insects. The wood has

been exported in small amounts from British Honduras to the

United States for use as veneers. In the Lake Izabal region it is

used in making canoes. In Oaxaca the tree is reported to attain a

height of 27 meters or more and a trunk diameter of 60-100 cm.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 5

The wood is used there for railroad ties. It is questionable whether

this species can be maintained as distinct from V. guatemalensis, to

which it is very closely related, and separable by characters that

appear to have slight importance, if any at all.

Stafleu (loc. cit., p. 466) has recently described a var. parvifolia

from British Honduras (El Cayo district, Gentle 2479), characterized

by having smaller 3-verticillate leaves and smaller apiculate flower

buds.

POLYGALACEAE

Reference: S. F. Blake, Polygalaceae, N. Amer. Fl. 25: 305-379.

1924.

Herbs, shrubs, or trees, sometimes woody vines, often with glands in the

tissues of the leaves and also in the flowers and fruits; leaves alternate,, opposite,

or verticillate, simple, entire, short-petiolate, without stipules but sometimes with

small stipular glands; flowers small or medium-sized, perfect, zygomorphic, gen-

erally racemose or spicate, each subtended by a bract and 2 bractlets; sepals 5,

free, or the lower 2 united, one posterior, 2 anterior, 2 lateral and interior, the

last (wings) usually much larger and petaloid; petals 3, rarely 5, hypogynous, the

anterior one (keel) boat-shaped, often with a terminal beak or fimbriate crest;

stamens usually 8, rarely 3-7, the filaments united for most of their length into

a sheath, this split on the upper side, usually united at the base to the keel or

upper petals or both; anthers mostly confluently 1-celled, dehiscent by a sub-

terminal pore; intrastaminal disk present or reduced to a gland at the base of the

ovary, or wanting; gynoecium of 1-2, rarely 3-5, united carpels, the style 1,

the stigma 2-lobate, often penicillate; ovules solitary, rarely 2-6, pendulous; fruit

a capsule, drupe, or samara, loculicidally dehiscent or indehiscent; seeds usually

solitary in each cell, generally pubescent, arillate, and with endosperm; embryo

straight, axial.

Ten genera with numerous species, widely distributed in tropical

and temperate regions. Only the following are represented in North

America.

Ovary and fruit 2-celled; fruit a compressed capsule.

Lower petals united with the keel; capsule usually broader than oblong; herbs or

shrubs; leaves alternate or verticillate Poly gala.

Lower petals free from the keel; capsule narrowly cuneate-oblong; usually

woody vines, sometimes erect shrubs; leaves alternate Bredemeyera.

Ovary and fruit 1-celled; fruit indehiscent, drupaceous or samaroid.

Keel petal with a plicate crest; fruit a samara, with a large broad wing on the

lower side; woody vines Securidaca.

Keel petal not cristate; fruit drupaceous, not or very obscurely winged (in

Central American species) ; erect shrubs or herbs Monnina.

BREDEMEYERA Willdenow

Mostly woody vines, sometimes suberect shrubs; leaves alternate, ovate or

oblong, penninerved; flowers small, in terminal panicles; sepals unequal, the 2

6 FIELDIANA: BOTANY, VOLUME 24

inner ones large and petaloid, wing-like, the 2 lateral ones adnate at the base to

the stamen tube, erect-connivent; keel petal about as long as the lateral ones,

concave-galeate, entire or 3-lobate, ecristate; stamens 8, united below into a sheath;

anthers dehiscent by an oblique introrse pore; ovary 2-celled, the style curved,

stigmatose and emarginate at the apex; capsule compressed, subcarnose, cuneate-

oblong, loculicidally dehiscent on the margins; seeds pendulous, glabrous or pilose,

comose at the hilum with very long hairs; endosperm scant.

About 60 species, in tropical America and Australia. Only one

reaches North America.

Bredemeyera lucida (Benth.) A. Bennett in Mart. Fl. Bras.

13, pt. 3: 51. 1874. Catocoma lucida Benth. in Hook. Journ. Bot. 4:

101. 1842.

Moist or wet thickets or open forest, sometimes on open forested

slopes or in second growth, 300 meters or less; Pete"n; Izabal. British

Honduras; Honduras; northern South America.

A shrub or a woody vine, when erect usually 2-4 meters high, when scandent

as much as 12 meters long, the branches slender, puberulent; leaves short-petiolate,

coriaceous, lustrous, elliptic to oblong, mostly 5-9 cm. long, acute or obtuse, with

an obtuse tip, obtuse at the base, green and usually glabrous above, somewhat

paler beneath, scaberulous or strigillose with lustrous golden hairs or often glabrate,

the nerves and veins prominent and reticulate on both surfaces; panicles usually

large and many-flowered, often much-branched, leafy, often densely flowered,

the flowers greenish yellow, pedicellate, 3 mm. long; sepals broadly ovate or

suborbicular, ciliate and pubescent; ovary pilose; fruit cuneate-oblong, 10-15

mm. long, shallowly emarginate, attenuate at the base, glabrous; seeds densely

hirsute and comose with very long and slender hairs.

MONNINA Ruiz & Pavon

Herbs or shrubs, rarely somewhat scandent; leaves alternate, entire, estipulate

or with stipular glands; flowers small, usually blue or lilac, in terminal and axillary,

sometimes paniculate racemes; sepals 5, deciduous, the 3 outer ones herbaceous,

free or the 2 lower ones united, the 2 inner ones (wings) much larger, petaloid;

petals 3, the lower one (keel) boat-shaped, not unguiculate, not appendaged, free

or nearly so; 2 upper petals usually oblong or liguliform, united below to the

staminal sheath; stamens 8 or 6, the filaments united nearly to the apex into a

sheath, the anthers confluently 1-celled, opening by a large introrse apical pore;

ovary 1-celled, the style sickle-shaped, the stigma lobes dissimilar; ovules solitary,

pendulous; disk usually reduced to a gland at the base of the ovary; fruit samaroid,

narrowly and almost equally winged, or more often drupaceous, the surface rugose;

seed glabrous, not arillate, the testa thin; endosperm thick.

About 85 species, distributed from southwestern United States

to Chile and Argentina, most numerous in the Andes of South

America. Twelve are known in North America, and 6 besides those

listed here are known from other parts of Central America.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 7

Racemes not at all comose, the bracts ovate, acute, 1.2-2.2 mm. long, not exceed-

ing the buds; leaves usually 1-3 cm. wide and broadest above the apex.

Pubescence of the stems closely appressed M. xalapensis.

Racemes more or less comose toward the apex, the bracts mostly lanceolate and

acuminate or attenuate, 2.5-7 mm. long, often conspicuously exceeding the

buds; leaves mostly 3-6 cm. wide, broadest at or below the middle.

Pubescence of the stems and leaves of short spreading hairs . . M. guatemalensis.

Pubescence of the stems and leaves of closely appressed hairs M. sylvatica.

Monnina guatemalensis Chodat, Bull. Herb. Boiss. 4: 249.

1896. Yaxtam (Huehuetenango).

Moist or wet thickets, 1,000-1,700 meters; Alta Verapaz (type

from Coban, Tuerckheim 8377); Baja Verapaz; Quiche"; Huehue-

tenango. Chiapas.

A slender shrub 1.5-2.5 meters high, with few branches, the stems densely

short-pilose with spreading hairs; leaves short-petiolate, rather thick when dried

and often lustrous, oval or elliptic, 6-11 cm. long, acute or acuminate, cuneate

to rounded at the base, pilosulous with short spreading hairs on both surfaces,

the lateral nerves 6-8 pairs; peduncles several near the ends of the branches,

axillary and extra-axillary, the racemes mostly 5-15 cm. long; bracts lance-ovate,

acuminate, 5-7 mm. long, the pedicels 1 mm. long, the flowers violet; sepals

3-3.8 mm. long; wing petals 4.5 mm. long; fruit ovoid, 7.5-8.5 mm. long, coarsely

rugose-reticulate, glabrous, narrowly and obscurely marginate, at first red, glossy

black at maturity.

Monnina sylvatica Schlecht. & Cham. Linnaea 5: 231. 1830.

Llordn de montana (Chiquimula).

Moist or wet thickets or mixed forest, 900-2,400 meters; Baja

Verapaz; Chiquimula; Suchitepe"quez; Quezaltenango; San Marcos.

Southern Mexico; Costa Rica.

An herb or shrub 1-2.5 meters high with few branches, the stems thinly pubes-

cent with short appressed hairs; leaves thin, short-petiolate, mostly elliptic or

ovate-elliptic, 7-15 cm. long, 3-6.5 cm. wide, acuminate, cuneate at the base,

sparsely strigillose on both surfaces, somewhat paler beneath; peduncles several

or numerous, simple or branched, the racemes dense or interrupted, 18 cm. long

or shorter, the flowers short-pedicellate, violet or orchid-purple; bracts lance-

subulate, 4.5-5 mm. long, soon deciduous; sepals suborbicular-ovate to oval-

ovate, 2-2.5 mm. long; wing petals 4-4.5 mm. long; fruit ovoid, 6-7 mm. long,

coarsely rugose-reticulate, cristate on the upper margin and with a fluted wing

1 mm. wide on the lower margin.

Monnina xalapensis HBK. Nov. Gen. & Sp. 5: 414. 1823. San

Benito; Tinta; Lap-chisquit (fide Aguilar); Tintilla; Zacate de venado;

Tintamora.

Wet to dry thickets and forest, often in oak, pine, or Cupressus

forest, common in second growth, 1,200-3,500 meters, abundant in

8 FIELDIANA: BOTANY, VOLUME 24

many regions; Alta Verapaz; Zacapa; Jutiapa; Guatemala; Sacate-

pe"quez; Chimaltenango; Solola; Quiche"; Totonicapan; Huehuete-

nango; Quezaltenango; San Marcos. Southern Mexico; Costa Rica;

Panama.

A shrub 1-3.5 meters high, often much-branched, the branches erect or strongly

ascending, sparsely or densely strigillose; leaves thin, petiolate, oblanceolate or

narrowly obovate, 4-8.5 cm. long, acuminate or acute, attenuate at the base,

sparsely strigillose throughout, paler beneath; peduncles axillary and terminal,

the racemes dense or interrupted, mostly 10 cm. long or shorter, the flowers violet

or purple, short-pedicellate; bracts triangular-ovate, acute or obtuse, deciduous;

sepals triangular-ovate or oblong-ovate, acute or obtuse, 2-3 mm. long; wings

rounded-oval or oval, 5-6 mm. long; fruit ellipsoid-ovoid, rugose-reticulate, 6-8.5

mm. long, at first red, becoming purple or purplish black when ripe, very juicy.

When loaded with ripe fruit, this is a conspicuous and rather

handsome shrub, but the flowers are not conspicuous. Deer browse

on the plant, and presumably it is eaten also by sheep and goats.

In the Occidente the purple juice of the ripe fruit is sometimes used

in place of ink.

POLYGALA L.

Reference: S. F. Blake, A revision of the genus Polygala in

Mexico, Central America, and the West Indies, Contr. Gray Herb.

47: 1-122, pis. 1, 2. 1916.

Herbs, shrubs, or trees; leaves alternate, or often opposite or verticillate,

simple, entire, short-petiolate, very rarely with stipular glands; flowers mostly

small and white, pink, or purple, in terminal or axillary, rarely extra-axillary

racemes, rarely umbellate; sepals 5, the 3 outer ones herbaceous, free or the lower

2 connate, deciduous or persistent, the 2 inner ones (wings) petaloid or rarely

subherbaceous, usually much larger than the others; petals normally 3, united

at the base, the lowest (keel) boat-shaped, unguiculate, sometimes 3-lobate, un-

appendaged or usually with an apical beak or crest; 2 upper petals ligulate to

ovate, sometimes galeate; stamens 8, rarely 6, the filaments united almost to the

apex, the anthers usually confluently 1-celled, opening by an apical pore; ovary

2-celled, the ovules solitary, pendulous from the apex of the central placenta;

style usually slender, bent, more or less excavate at the apex, the stigma 2-lobate;

capsule equally or unequally 2-celled, often winged or marginate, compressed

contrary to the partition, loculicidally dehiscent; seeds globose to fusiform or

conic, usually pubescent, almost always arillate, with or without endosperm, the

testa crustaceous.

Species 500 or more, widely distributed in temperate and tropical

regions. About 180 are known from North America. A very few

besides those listed here are found in southern Central America.

Keel petal obtuse, without a crest or beak.

Calyx with all its sepals free.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 9

Wing sepals very small, about one-third longer than the outer sepals; capsule

subcoriaceous, thick-walled. Erect shrub with somewhat coriaceous

leaves and yellowish flowers P. jamaicensis.

Wing sepals much larger than the outer sepals; capsule membranaceous-

herbaceous, thin-walled.

Lower sepals petaloid, the upper sepal herbaceous, persistent; capsule about

3 mm. long; aril of the seed glabrous P. Purpusii.

Lower sepals herbaceous, like the others, all deciduous; capsule usually

much larger; aril pubescent, at least at the apex.

Wings rounded-oval or broadly oval, usually less than twice as long as

wide P. consobrina.

Wings oblong, oblong-oval, or oblong-obovate, from almost twice to

4 times as long as wide.

Wings 7-10 mm. long P. costaricensis.

Wings 6.5 mm. long or shorter.

Wings glabrous P. polymorpha.

Wings pubescent dorsally.

Wings sparsely long-pilose dorsally; leaves lance-elliptic to linear.

P. trichoplera.

Wings incurved-puberulent along the costa and toward the apex;

leaves ovate or lance-ovate.

Seed glabrous P. guatemalensis.

Seed pubescent P. Durandi.

Calyx with the 2 lower sepals connate.

Plants annual P. bryzoides.

Plants perennial, often woody.

Flowers green P. hondurana.

Flowers purple or violet.

Branches of the inflorescence subtomentose P. Securidaca.

Branches of the inflorescence puberulent or strigose P. floribunda.

Keel petal with a beak or crest at the apex.

Capsule winged or marginate on the upper cell, marginless on the lower cell.

Capsule strongly transverse-rugose or transverse- veined P. rhysocarpa.

Capsule smooth, not transverse-rugose P. Salviniana.

Capsule marginless, or narrowly and equally marginate on both cells.

Leaves all or most of them verticillate.

Racemes subglobose, as broad as long P. conferta.

Racemes much longer than broad.

Capsule 1.5 mm. long P. asperuloides.

Capsule 2-2.5 mm. long P. aparinoides.

Leaves not verticillate except rarely a few whorls on the lowest part of the

stem.

Racemes slender and much elongate, mostly 5 mm. broad or less, tapering

at the apex.

Stems glabrous.

Plants perennial, usually with several stems P. alba.

Plants annual, the stems solitary.

Seed glabrous; flowers white, densely crowded in the racemes.

P. gracillima.

Seed pubescent; flowers purple or white, the racemes rather lax.

P. leptocaulis.

Stems glandular-puberulent.

10 FIELDIANA: BOTANY, VOLUME 24

Capsule more than 3 mm. long; flowers white P. Berlandieri.

Capsule 1.5-1.7 mm. long; flowers purple or white.

Seed appressed-pubescent, the hairs not glochidiate; aril well de-

veloped P. paniculata.

Seed covered with incurved uncinate-tipped hairs; aril obsolete.

P. glochidiata.

Racemes mostly short and broad, generally about as broad as long, usually

not at all tapering at the apex, rarely more elongate but then 8-10 mm.

thick.

Stems with several whorls of leaves below; racemes very dense and many-

flowered, mostly more than twice as long as broad. .P. hygrophila.

Stems with a single whorl of leaves at the very base, or all the leaves

usually alternate.

Wings conspicuously cuspidate at the apex P. longicaulis.

Wings rounded or merely apiculate at the apex.

Wings 4.7-5.3 mm. long; aril minute P. adenophora.

Wings 2.7-3.6 mm. long; aril 0.5-1.3 mm. long P. variabilis.

Polygala adenophora DC. Prodr. 1: 327. 1824.

Wet open pine forest, at or little above sea level; British Hon-

duras. Trinidad and northern South America (Guianas).

A very slender, erect annual 15-35 cm. high, simple or branched above,

glabrous, sometimes papillose below; leaves alternate, linear, 4-16 mm. long,

erect or ascending; racemes solitary at the ends of the peduncles, obtuse, 7-12

mm. broad, 1-4 cm. long, most often about as long as broad; bracts ovate, decidu-

ous, the pedicels 1 mm. long, the flowers purple or rarely white; sepals oval or

ovate, apiculate or obtuse, 1.3-2 mm. long; wings narrowly elliptic, about 5 mm.

long, obscurely apiculate or obtuse; keel with a lobate crest near the apex; capsule

narrowly elliptic, 3.5 mm. long, 1-1.3 mm. wide; seed obconic, pilose, comose at

the apex, 1.8-2 mm. long, the aril minute.

Polygala alba Nutt. Gen. PI. 2: 87. 1818.

Moist or wet, open places, 300-1,000 meters; Huehuetenango

(Cie"naga de Lagartero; between Nenton and Las Palmas). Western

United States; Mexico.

Stems slender, erect, usually numerous from a perennial root, simple or sparsely

branched, usually 15-35 cm. high, glabrous; leaves scattered except for 1-2 verticels

at the base of the stem, the lowest spatulate-obovate and 4-12 mm. long, the others

linear, cuspidate-acuminate, 8-25 mm. long; racemes dense, conic-cylindric, about

5 mm. thick, 2-8 cm. long; flowers white or sometimes tinged with purple; sepals

ovate to oblong, obtuse, 1.3-1.5 mm. long, the wings elliptic, almost 3 mm. long;

keel 3 mm. long, the crest of 4 lobes on each side; capsule elliptic or oblong-elliptic,

2.5-2.9 mm. long; seed pilose, 2.3-2.5 mm. long, the aril 0.8-1.5 mm. high, its 2

lobes oblong, appressed.

The seeds of the Guatemalan specimens are more spreading-

pilose than is typical of the species.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 11

Polygala aparinoides Hook. & Arn. Bot. Beechey Voy. 227.

1836. P. nemoralis A. W. Benn. Journ. Bot. 17: 172. 1879, in part

(type from Chilasco, Baja Verapaz, Salvin & Godmari). P. Vogtii

Chodat, Me"m. Soc. Phys. Geneve 31(2), pt. 2: 144. 1893 (based in

part on material from Coban, Alta Verapaz, Tuerckheim 136).

Ipecacuana blanca; Peor es nada (Huehuetenango).

Moist or wet forest or thickets, sometimes in pine forest, occa-

sionally in marshes, 2,600 meters or lower; Alta Verapaz; Baja

Verapaz; El Progreso; Izabal; Zacapa; Chiquimula; Santa Rosa;

Quiche". Central and southern Mexico.

Stems slender, solitary or several from a slender perennial base, erect or spread-

ing, often 40 cm. long or more; leaves 5-verticillate throughout or the uppermost

scattered, lanceolate or the lowest broader, 1-2 cm. long, 5-10 mm. wide, acute

or subacute, cuspidate, acute at the base, sessile or nearly so, the margins obscurely

denticulate; racemes cylindric, tapering at the apex, rather dense, 5-6 mm. thick,

16 cm. long or usually much shorter, the pedicels less than 1 mm. long; flowers

purplish or reddish mixed with green; sepals broadly oval or ovate, glandular-

ciliate, 1.5-1.8 mm. long; wings elliptic-obovate, 2-3.5 mm. long, rounded; keel

cristate at the apex; capsule broadly elliptic, 2-2.5 mm. long; seed appressed-

pilose, equaled by the 2 narrowly oblong, appressed lobes of the aril.

This has been reported from Guatemala as P. verticillata L.,

P. Boykinii Nutt., and P. galioides Poir.

Polygala asperuloides HBK. Nov. Gen. & Sp. 5: 403. 1823.

At or little above sea level; Izabal (near Izabal, Sereno Watson

19). British Honduras; Panama; Colombia.

Stems 1-several from an annual root, erect or ascending, 10-30 cm. long;

leaves all 5-verticillate or the uppermost in 2's or 3's, the lowest ones rounded-

obovate, 7-9 mm. long, 4.5-5 mm. wide, the middle and upper ones lanceolate or

lance-elliptic, 1-2 cm. long, cuspidate, with obscurely denticulate margins; flowers

pink, the racemes 3-5 mm. thick, 1-5 cm. long, the pedicels 0.4 mm. long; sepals

rounded-ovate to oblong, 1 mm. long or less; wings broadly elliptic, 1.5 mm.

long; keel cristate at the apex; capsule suborbicular, 1.5 mm. long; seed appressed-

pilose, 1.5 mm. long, the aril 1 mm. long, with 2 oblong appressed lobes.

Polygala Berlandieri Wats. Proc. Amer. Acad. 21: 416. 1886.

Grassy hillsides or a weed in cultivated ground, 1,200-2,300

meters; Santa Rosa; Guatemala. Mexico; Salvador.

A slender erect annual, often much-branched, 5-15 cm. high, densely stipitate-

glandular; lowest leaves 4-5-verticillate, most of the leaves scattered and linear,

5-20 mm. long, acute, cuspidate; racemes cylindric, rather lax, 5 cm. long or shorter,

the pedicels 0.8 mm. long; flowers usually white, the sepals rounded-ovate or ovate-

oblong, subacute or obtuse, 1.2 mm. long; wings spatulate-obovate, 2.3 mm. long,

rounded at the apex; keel cristate at the apex; capsule narrowly elliptic, 3.3 mm.

long or slightly shorter; seed 2.5 mm. long, sericeous, the aril 0.5 mm. long, 2-lobate.

12 FIELDIANA: BOTANY, VOLUME 24

Polygala bryzoides St. Hil. Fl. Bras. Merid. 2: 44. 1829. P. an-

gustifolia HBK. Nov. Gen. & Sp. 5: 405. 1823, not P. angustifolia

Gilib. 1781.

Moist or dry, rocky and open or grassy slopes, sometimes on

sandbars along streams or in pine forest, 1,500 meters or less; Pete"n;

Alta Verapaz; Baja Verapaz; Izabal; Zacapa; Jutiapa; Santa Rosa;

Suchitepe"quez ; Retalhuleu ; Huehuetenango. Mexico ; Honduras and

Salvador to Panama; West Indies; southward to Brazil.

Plants annual, simple or branched, 10-30 cm. high, very slender, pubescent

throughout with short, straight or incurved and more or less appressed hairs;

leaves alternate, linear or lanceolate, 1.5-4.5 cm. long, 2-9 mm. wide, acute or

acuminate at each end, 1-nerved, often almost glabrous; flowers pinkish purple

and greenish, the racemes terminal and axillary, 1.5-5 cm. long, the pedicels mostly

recurved, more than 1 mm. long; sepals oblong-ovate, obtuse, sparsely ciliate at

the apex, with 2-3 pairs of pedicellate glands, 1.5-1.8 mm. long; wings broadly

cuneate-obovate, retuse, 3-4 mm. long; capsule oblong-oval, 3 mm. long; seed

2.5 mm. long, the aril 0.7 mm. long.

Polygala conferta A. W. Benn. ex Hemsl. Diag. PL Nov. 2.

1878.

Open pine forest, 1,000-1,200 meters; Huehuetenango; reported

by Blake as collected in Guatemala, "Barranca de Fuerengo,"

Bernoulli 105; the locality name is incorrectly transcribed, and we

are unable to guess what it may be. Central and southern Mexico.

A slender erect annual, simple or sparsely branched, 7-14 cm. high, glabrous;

lowest leaves 5-verticillate, spatulate-obovate, 3.5-7.5 mm. long, the middle leaves

linear, 7-13 mm. long, acuminate, mucronate; racemes headlike, dense and many-

flowered, 6 mm. broad, the pedicels 1.2-1.5 mm. long; flowers pinkish or greenish

white, the sepals rounded-ovate or ovate, obtuse or subacute, 1 mm. long; wings

oval, 1.8 mm. long; keel cristate; capsule rounded-oval, 1.3 mm. long; seed pubes-

cent, 1 mm. long, the aril 0.7 mm. high, with 2 oblong appressed lobes.

Polygala consobrina Blake, Contr. Gray Herb. 47: 48. 1916.

P. Hayesii Blake, loc. cit. (type collected near Guatemala, Sutton

Hayes in 1860). Hierba del rosario; Ipecacuana; Calzdn de nino.

Moist slopes, fields, or thickets, 150-2,000 meters; Chiquimula;

Santa Rosa; Escuintla (type from Escuintla, J. D. Smith 1980);

Guatemala; Chimaltenango. Salvador.

Perennial from a rather thick, woody root, the stems usually several, erect

or ascending, 10-30 cm. high, densely puberulent with mostly incurved hairs;

lower leaves oval, obtuse, the middle and upper ones alternate, ovate, 2-5 cm.

long, acute, broadly cuneate at the base, reticulate- veined, puberulent; racemes

dense and many-flowered, mostly 5 cm. long or shorter, the flowers violet or

greenish violet; sepals lanceolate, acuminate, 3 mm. long; wings oblong-oval,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 13

4.5-5.5 mm. long, 2.5 mm. wide, rounded at the apex, ciliolate, pilosulous on

the outer surface; capsule oval, densely ciliate and pilosulous, 9 mm. long; seed

6 mm. long, the aril 2.3-3.8 mm. high, pilose.

This has been reported from Guatemala as P. americana Mill,

and P. hebantha Benth. The species of this group, formerly passing

as P. americana, have been multiplied beyond reason, upon characters

that apparently are variable and obscure. It is probable that at

least half, and still better two-thirds, of the recently proposed ones

should be reduced to synonymy.

Polygala costaricensis Chodat, Bull. Soc. Bot. Belg. 30, pt. 1:

298. 1891. ?P. guatemalensis Gandog. Bull. Soc. Bot. France 60:

454. 1913, not P. guatemalensis A. W. Benn. 1895. P. platycarpa

var. stricta Chodat, Me"m. Soc. Phys. Geneve 31(2), pt. 2: 27. 1893

(type collected between Rabinal and Santa Ana, Baja Verapaz,

Bernoulli 1092). P, isotricha Blake, Contr. Gray Herb. 47: 53. 1916.

Ipecacuana amarilla (fide Aguilar).

Moist or wet thickets or forest, often in pine-oak forest, or in

rather dry, exposed, rocky places, sometimes a weed in coffee planta-

tions, 150-2,100 meters; Alta Verapaz; Zacapa; Chiquimula; Jalapa;

Jutiapa; Santa Rosa; Guatemala; Sacatepe"quez; Chimaltenango;

Quich^ ; Suchitepe"quez ; Quezaltenango. Chiapas; Costa Rica.

Perennial from a woody root, sometimes suffrutescent at the base, the stems

solitary or several, simple or branched, erect to procumbent, sometimes 75 cm.

long but usually much shorter, puberulent and sometimes short-pilose; leaves

short-petiolate, thin, ovate or ovate-lanceolate, 2.5-6.5 cm. long, acute or acumi-

nate, obtuse or cuneate at the base, reticulate-veined, rather sparsely puberulent

and sometimes short-pilose; racemes lax or rather dense, 3-12 cm. long, the

flowers purple or violet, short-pedicellate; sepals lanceolate, acuminate, 2.5-3 mm.

long; wings oblong to oblong-oval, 8-10 mm. long, sparsely puberulent along the

costa and at the apex; capsule oval, ciliate, puberulent, about 10 mm. long; seed

short-pilose, the aril 2.7 mm. high.

This has been reported from Guatemala as P. americana Mill.

Polygala Durandi Chodat, Bull. Soc. Bot. Belg. 30, pt. 1: 300.

1891. Sopladorcito; Hoja de aire.

Open or shaded banks, moist thickets, or open oak forest, 200-

2,100 meters; El Progreso; Zacapa; Chiquimula; Jalapa; Jutiapa;

Santa Rosa; Escuintla; Guatemala; Chimaltenango; Solola; Hue-

huetenango. Costa Rica.

Perennial from a woody root, the stems solitary or several, erect or ascending,

50 cm. long or less, simple or branched, densely pubescent with short hairs; leaves

short-petiolate, alternate, ovate or ovate-lanceolate, 2-5 cm. long, acute or acumi-

14 FIELDIANA: BOTANY, VOLUME 24

nate, or the lowest oval and obtuse, cuneate or obtuse at the base, thin, reticulate-

veined, sparsely or often densely pubescent; racemes mostly 4-5 cm. long, dense

or lax, many-flowered, the flowers violet or purple, short-pedicellate; sepals lanceo-

late, acute, 2-2.8 mm. long; wings oblong, 6 mm. long, rounded at the apex,

sparsely ciliate, puberulent along the costa and near the apex; capsule oval, pubes-

cent, 1 cm. long; seed 5 mm. long, pilose, the aril 3 mm. high.

This and related species are used in medicine in Guatemala, at

least in household remedies. The roots of some American species

of Poly gala are used like ipecac (Cephaelis Ipecacuanha}.

Polygala floribunda Benth. PI. Hartweg. 58. 1840. P. sphae-

rospora Chodat, Bot. Jahrb. 52, Beibl. 115: 75. 1914 (type collected

above San Jeronimo, Baja Verapaz, Seler 3393). Chupac, Chopac,

Raxjuc (Alta Verapaz, Quecchi).

Moist or wet thickets or forest, often in pine forest, sometimes

in second growth, 1,000-1,800 meters; Alta Verapaz; Baja Verapaz;

Zacapa; Jalapa; Santa Rosa; Guatemala; Quiche"; Huehuetenango.

Chiapas and perhaps elsewhere in southern Mexico.

An erect shrub 1-3 meters high, or often a large woody vine, sparsely pubescent

or puberulent or usually almost glabrous; leaves short-petiolate, rather thick and

firm, rather conspicuously reticulate-veined, ovate or lanceolate, 4-10 cm. long,

acute or acuminate, mucronate, obtuse or cuneate at the base, paler beneath;

racemes lax, many-flowered, mostly 10-20 cm. long, axillary and terminal, often

very numerous, the pedicels 8-13 mm. long; flowers bright purple; sepals sub-

equal, obtuse, oval, ciliate, 4 mm. long; wings suborbicular, 8-11 mm. long or in

fruit somewhat larger, venose, ciliate; capsule transversely broad-oblong and

obcordate, stipitate, ciliate and pubescent, about 8 mm. long and 10 mm. wide;

seed globose, tomentose, 3 mm. thick, the aril 2 mm. high.

A common and showy plant of the Coban region, often occurring

in great abundance, especially in thickets and rather open pine

forest. It is an exceptionally beautiful plant because of its brilliantly

colored flowers, and it is much planted for ornament, not only in

Coban but in other parts of Guatemala. The shrub is usually erect

in cultivation, but in the woods it usually is a small or medium-sized

vine. In North American Flora it is stated that the species of Polygala

are never scandent, but this one is decidedly so. In Alta Verapaz

the roots are used as a substitute for soap, giving abundant suds

when macerated in water. They are used particularly for removing

dandruff, and for treating eczema and other cutaneous diseases.

The roots also are chewed to cleanse the teeth and harden the gums.

Polygala glochidiata HBK. Nov. Gen. & Sp. 5: 400. 1823.

Moist fields, brushy slopes, oak-pine forest, 800-1,900 meters,

or sometimes at even lower elevations; Jutiapa; Santa Rosa; Saca-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 15

tepe"quez; Huehuetenango. Mexico; British Honduras; Honduras;

Costa Rica; Cuba; South America.

A very slender annual 30 cm. high or less, often much-branched, finely stipitate-

glandular; lower leaves 5-verticillate, the lowest lance-obovate, 3-4.5 mm. long,

the middle and upper leaves linear, 5-12 mm. long, cuspidate; racemes cylindric,

rather lax, 5-6 mm. thick, 1-8 cm. long; flowers short-pedicellate, rose-purple,

rarely white; sepals elliptic or oblong, obtuse, 1 mm. long; wings obovate-oval,

2.5 mm. long, rounded at the apex; keel cristate at the apex; capsule elliptic, 1.5

mm. long; seed 1 mm. long, covered with incurved uncinate-tipped hairs, these

spreading when wet; aril obsolete.

Polygala gracillima Wats. Proc. Amer. Acad. 22: 398. 1887.

Open places in pine-oak forest, 1,200-2,400 meters; Baja Verapaz;

Chiquimula; Quiche"; Huehuetenango. Mexico.

A very slender, erect, glabrous annual, 10-20 cm. high, usually much-branched

above; lowest leaves ternate, obovate, 3-3.5 mm. long, the others scattered, linear,

2-7 mm. long; racemes dense, many-flowered, tapering at the apex, 6-25 mm.

long, 2.5 mm. thick; flowers short-pedicellate, white, the bracts persistent, lance-

subulate; sepals ovate or oval, 0.6 mm. long; wings oblong-oval, 1.3 mm. long;

keel cristate at the apex; capsule suborbicular, 1 mm. long; seed black, glabrous,

ellipsoid-fusiform, striate, 0.5 mm. long, the aril minute.

Polygala guatemalensis A. W. Benn. Journ. Bot. 33: 108.

1895.

Known only from the type, Coban, Alta Verapaz, 1,340 meters,

Tuerckheim 298 (in part).

Perennial, suffrutescent, 50 cm. high, with few erect branches, rather densely

and finely pubescent; leaves alternate, short-petiolate, ovate, 2.5-4 cm. long,

acuminate, reticulate-veined, sparsely pubescent on both surfaces; racemes rather

dense, 4.5-9.5 cm. long; sepals lanceolate, acute, 3 mm. long; wings oblong, 5-6

mm. long, rounded at the apex, puberulent along the costa and near the apex;

immature capsule oval, pilosulous, 8 mm. long; seed glabrous, 2.2 mm. long, the

aril 1 mm. high.

Polygala hondurana Chodat, Bot. Jahrb. 52, Beibl. 115: 75.

1914. P. tonsa Blake, Contr. Gray Herb. 47: 63. 1916 (type from La

Vega, Santa Rosa, Heyde & Lux 3067).

Moist or usually wet thickets or forest, 600-1,800 meters; Alta

Verapaz; Chiquimula; Jalapa; Jutiapa; Santa Rosa; Huehuetenango.

Honduras; Salvador; Nicaragua.

Perennial from a rather thick, nodose root, erect, a meter high or less, usually

frutescent, often with numerous, erect or ascending, slender, green branches,

sparsely puberulent or glabrate; leaves short-petiolate, alternate, thin, ovate to

lanceolate, 5-10 cm. long, long-acuminate, usually rounded or obtuse at the base,

16 FIELDIANA: BOTANY, VOLUME 24

deep green above, paler beneath; racemes lax, few-flowered, mostly shorter than

the leaves, the bracts caducous, the pedicels slender, 4-8 mm. long; flowers pale

green; sepals ciliate, puberulent, 3 mm. long; wings ovate-orbicular, 8-9 mm. long,

venose, glabrous; capsule quadrate-orbicular, deeply emarginate, 6-7 mm. long

and equally wide; seed tomentose, 4.8 mm. long, the aril 2 mm. high.

This species is easily recognized by its shrubby habit and large

pale green flowers. The name "hierba del rosario" is applied to

this species in Salvador. It has been reported from Guatemala as

P. monninoides HBK.

Polygala hygrophila HBK. Nov. Gen. & Sp. 5: 395. pi. 508.

1823.

Savannas or grassy pine forest, at or little above sea level;

British Honduras. Chiapas; Panama; northern South America.

A slender erect annual, 35 cm. high or less, simple or with a few erect branches,

glabrous; lowest leaves verti ciliate, the upper ones scattered, linear or lance-

linear, 13-18 mm. long; racemes cylindric-ovoid, slightly comose at the apex,

about 2 cm. long and 8 mm. thick; sepals oval, 1.8 mm. long; wings oval, 4.5 mm.

long, rounded at the apex, green tinged with pale red or deep pink; keel cristate;

capsule subglobose, turgid; seed ellipsoid, 1.7 mm. long, pubescent, the aril 1 mm.

long.

This has been reported from British Honduras as P. Timoutou

Aubl., a South American species, with much broader leaves, extend-

ing northward into Panama.

Polygala jamaicensis Chodat, Me"m. Soc. Phys. Geneve 31(2),

pt. 2: 11. 1893. P. petenensis Lundell, Lloydia 4: 51. 1941 (type

from Sabana Zis, northwestern end of Lago de Pete"n, C. L. Lundell

3187). Limonaria cimarrona.

In forest, 800 meters or less; Pete"n (Camp 36, Guatemalan

boundary). British Honduras; Jamaica.

A shrub or small tree, in Guatemala 1-3 meters high, much-branched, woody

throughout, the branches densely puberulent or finally glabrate; leaves short-

petiolate, ovate to ovate-lanceolate, 3.5-8.5 cm. long, obtuse to acuminate, cuneate

at the base, sparsely and minutely strigillose; peduncles 1-1.5 mm. long, the axis

of the raceme 3-6 mm. long, the pedicels 2-5 mm. long, puberulent, the flowers

few, yellow; sepals rounded-ovate or deltoid-ovate, 1.5 mm. long; wings deltoid-

ovate, 2 mm. long; keel 4 mm. long; capsule subquadrate, lobate one-fifth its

length, with rounded lobes, ciliate and pubescent, 7-11 mm. wide, stipitate;

seed 6 mm. long, the aril 3 mm. high.

After comparing the two known collections of P. petenensis with

numerous Jamaican collections of P. jamaicensis, we are unable to

find any characters by which they can be distinguished.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 17

Polygala leptocaulis Torr. & Gray, Fl. N. Amer. 1: 130. 1838.

P. Pringlei Wats. Proc. Amer. Acad. 25: 142. 1890. Tamiz.

Open, grassy, moist or wet plains, often in marshes or in oak

forest, 2,000 meters or less; Izabal; Jalapa; Jutiapa; Quiche"; Huehue-

tenango. Southern United States; Mexico; British Honduras; Hon-

duras; Salvador; Cuba; South America.

A very slender, erect annual, 50 cm. high or less, glabrous, simple or sparsely

branched; leaves alternate, linear, 8-25 mm. long; racemes cylindric, lax or dense,

13 cm. long or less, about 5 mm. thick; flowers lilac or greenish pink, short-pedicel-

late; sepals ovate, obtuse, 1 mm. long; wings obovate, 2 mm. long, rounded at

the apex; keel cristate; capsule oblong, 1.6-1.8 mm. long; seed subcylindric, ap-

pressed-pubescent, 1.2 mm. long, the aril minute, 2-lobate.

This has sometimes been confused with P. paludosa St. Hil.

Polygala longicaulis HBK. Nov. Gen. & Sp. 5: 396. 1823.

Cambray (Huehuetenango).

Moist savannas or pine forest, sometimes in dry rocky open places,

1,400 meters or less; Pete"n; Izabal; Zacapa; Chiquimula; Jalapa;

Jutiapa; Santa Rosa; Quiche"; Huehuetenango. Central and southern

Mexico; British Honduras to Panama; West Indies; South America.

A slender erect annual, 45 cm. high or less, glabrous, simple or branched above;

leaves alternate, linear, 4-16 mm. long, erect or ascending; racemes very obtuse,

dense, 7-12 mm. broad, usually little longer than broad but sometimes almost

5 cm. long; bracts ovate, deciduous, the pedicels 1 mm. long; flowers rose-purple;

sepals oval or ovate, apiculate or obtuse, 1.5-2 mm. long; wings narrowly elliptic,

about 5 mm. long, obscurely apiculate or obtuse; keel cristate; capsule narrowly

elliptic, 3.5 mm. long; seed obconic, pilose, comose at the apex, 2 mm. long, the

aril minute.

This has been reported from Guatemala as P. trichosperma Torr.

Polygala paniculata L. PI. Jam. Pugill. 18. 1759. P. paniculata

f. leucoptera Blake, ( Contr. Gray Herb. 47: 101. 1916. Lanillo;

Ipecacuana; Rax cukichoj (Coban, Quecchi); Mentol; Menta.

Moist fields, banks, or thickets, on sandbars along streams, often

a weed in cultivated or waste ground, sometimes in open, oak or

pine forest, 2,000 meters or less, most frequent at low elevations;

Alta Verapaz; Zacapa; Chiquimula; Jalapa; Jutiapa; Santa Rosa;

Escuintla; Guatemala; Sacatepe"quez ; Suchitepe"quez ; Retalhuleu;

Quiche" ; Huehuetenango; Quezaltenango; San Marcos. Texas;

southern Mexico; British Honduras to Salvador and Panama;

West Indies; South America.

18 FIELDIANA: BOTANY, VOLUME 24

A slender erect annual 10-30 cm. high, often much-branched, densely stipitate-

glandular; lowest leaves verticillate, the others alternate, linear, 8-18 mm. long;

racemes lax, cylindric, 5-6 mm. thick, 9 cm. long or less, the flowers short-pedicel-

late, purplish, pink, or white; sepals ovate or oblong-ovate, obtuse, 1.3 mm.

long; wings obovate or spatulate-obovate, 2-2.5 mm. long, rounded at the apex;

keel cristate; capsule elliptic, 1.7 mm. long; seeds appressed-pubescent, 1.5 mm.

long, the aril 0.4-0.8 mm. long, the 2 lobes appressed.

Called "hierba del colico" in Salvador, where the plant is used

in domestic medicine. The Guatemalan names "menta" and

"mentol" allude to the fact that the roots, when chewed, have a

mint-like taste. The same flavor is found in the roots of Polygala

species of the southern United States, as is well known to children

living where the plants are found. This is by far the commonest

Polygala of Central America, being an unattractive weedy plant,

often common about dwellings and in waste ground generally.

Polygala polymorpha Chodat, Bot. Jahrb. 52, Beibl. 115: 74.

1914.

Usually on dry limestone hillsides, sometimes a weed in cornfields,

900-2,600 meters; endemic; Huehuetenango (type from Chacula,

Seler 3130).

Perennial from a somewhat ligneous root, the stems few or numerous, erect

or decumbent, 10-20 cm. long, strigillose; leaves short-petiolate, alternate, ovate-

elliptic to narrowly lance-oblong, 2-3.5 cm. long, obtuse or acute, thick, mucronate,

sparsely and minutely strigillose; racemes lax, terminal, few-flowered; flowers

purple, 6-7 mm. long; sepals lance-linear, acute; wings oblong-elliptic, glabrous;

capsule oblong, 9 mm. long, cuneate at the base, pilosulous; seed hirsute, the aril

3-lobate.

Polygala Purpusii Brandeg. Univ. Calif. Publ. Bot. 4: 88. 1910.

Dry, open, often rocky slopes, 1,200-1,600 meters; Huehue-

tenango (region of Cuilco). Puebla, Mexico.

An erect perennial as much as 50 cm. high, herbaceous or suffrutescent below,

the perpendicular root rather thick and lignescent, the stems few or numerous,

simple or sparsely branched, incurved-pilosulous, terete; leaves on short slender

petioles, oval to oblong-oval, 9-18 mm. long, 4-11 mm. wide, rounded or very

obtuse at the apex, acute at the base, incurved-puberulent, usually rather densely

so, especially beneath; racemes terminal, sessile or short-pedunculate, 12 cm. long

or shorter, laxly many-flowered, the flowers lavender, slender-pedicellate, the

pedicels recurved in age; bracts rather conspicuous, narrowly lanceolate; upper

sepal herbaceous, persistent and conspicuous in fruit, 3 mm. long or less; lower

sepals petaloid, oblong-obovate, deciduous, not unguiculate, 3.5 mm. long; wings

obovate-oval, 4-4.7 mm. long; keel whitish, with a yellowish tip; capsule sub-

orbicular, incurved-puberulent toward the apex, 3 mm. long; seed obovoid, pilose,

2.5 mm. long; aril glabrous, 2-lobate.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 19

Polygala rhysocarpa Blake, N. Amer. Fl. 25: 366. 1924.

Dry rocky mountain slopes, 2,800-3,500 meters; Huehuetenango

(below Calaveras, Sierra de los Cuchumatanes, Steyermark 50347).

Veracruz, Mexico.

Stems few or several from a very slender, annual root, 25 cm. high or less,

erect or nearly so, simple or sparsely branched, puberulent with spreading gland-

like hairs; lowest leaves obovate, 3-5 mm. long, puberulent, the other leaves alter-

nate, oblanceolate or linear, 4-15 mm. long, acuminate at each end, puberulent;

racemes cylindric, acute or acuminate, dense above, lax below, 3-5 mm. thick,

1.5-7 cm. long; bracts lance-subulate, deciduous; flowers greenish white or purplish;

sepals ovate to lance-elliptic, obtuse to acuminate, 1-1.6 mm. long; wings obovate,

2-2.2 mm. long, rounded at the apex, cuneate at the base; keel about 2 mm.

long, the crest on each side consisting of a cuneate lamella and 2 entire or 2-parted

lobes; capsule oblique-oval, emarginate, rounded at the base, glabrous, with 6-7

conspicuous, usually greenish veins on each side of the septum and also evidently

transverse-rugose, winged on the upper side at the apex, 3 mm. long; seed cylindric,

often curved, pubescent, 1.8 mm. long, the aril 1 mm. long, its 2 linear lobes ap-

pressed.

Polygala Salviniana A. W. Benn. Journ. Bot. 17: 203. 1879.

P. macroloncha Chodat, Bot. Jahrb. 52, Beibl. 115: 84. 1914 (type

from Zaragoza, Chimaltenango, Seler 2925). P. microloncha Chodat,

loc. cit. (type collected near Chacula, Huehuetenango, Seler 3138).

P. oxysepala Blake, Contr. Gray Herb. 47: 109. 1916 (type from

Santa Rosa, Baja Verapaz, Tuerckheim 1202). Peor es nada (Hue-

huetenango).

Open hillsides, fields, or brushy slopes, often in pine or oak forest,

900-2,700 meters; Baja Verapaz; Jalapa; Santa Rosa; Sacatepe"quez

(type from Volcan de Fuego, above Las Calderas, Salving; Chimal-

tenango; Solola; Quiche"; Huehuetenango; Quezaltenango. Hon-

duras.

Perennial from a somewhat ligneous root, the stems usually several, slender

and wiry, erect or ascending, simple or sparsely branched, 60 cm. long or less,

strigillose or puberulent; leaves alternate, linear-acicular, 5-13 mm. long, cuspidate,

1-nerved, sparsely puberulent or almost glabrous; racemes lax or rather dense,

7 mm. thick, 6 cm. long or shorter, the bracts subulate, deciduous; flowers short-

pedicellate, greenish white; sepals ovate or ovate-lanceolate, obtuse to acuminate,

1-1.8 mm. long; wings obovate, 3-3.5 mm. long, obtuse; capsule elliptic, 2.5 mm.

long; seed oblong, pubescent, 2 mm. long, the aril 1.2-1.5 mm. long, the 2 lobes

oblong.

This has been reported from Guatemala as P. scoparia HBK.

Polygala Securidaca Chodat, Bot. Jahrb. 52, Beibl. 115: 76.

1914. Hierba grande.

20 FIELDIANA: BOTANY, VOLUME 24

Common in mountain forests of Honduras; to be expected in the

mountains of Chiquimula.

A shrub, the branches subtomentose when young; leaves short-petiolate,

alternate, ovate-elliptic, 4-7 cm. long, obtuse, rounded at the base, subcoriaceous,

softly tomentulose; racemes terminal, solitary or subpaniculate, short-cylindric,

3-5 cm. long, 2.5 cm. broad, the rachis tomentulose or hirsute, the pedicels hirsute,

7 mm. long; flowers purple, 10-12 mm. long.

Polygala Seleri Chodat, Bot. Jahrb. 52, Beibl. 115: 73. 1914.

Known only from the type, from Cuesta de la Concepcion,

Jacaltenango, Huehuetenango, Seler 3244.

Perennial from a ligneous root, the stems slender, branched, 20-30 cm. long,

puberulent; leaves alternate, short-petiolate, ovate or ovate-lanceolate, 3-5.5 cm.

long, acute or acuminate, glabrate; racemes rather lax, 5-10 cm. long, the flowers

violet, 9 mm. long, short-pedicellate; upper sepal very acute; wings elliptic-oblong,

glabrous, obtuse or short-acute, twice as long as broad, 9 mm. long; ovary long-

pilose; immature capsule orbicular, ciliate.

We know this plant only by a photograph of the type.

Polygala trichoptera Chodat, Bot. Jahrb. 52, Beibl. 115: 74.

1914.

Known only from the original material, limestone hillsides,

Uaxacanal, Huehuetenango, 1,300-1,400 meters, Seler 2796, 2904.

Perennial, the stems slender, erect, hirsute, 10-20 cm. high, simple or sparsely

branched; leaves lance-elliptic to lanceolate or linear, sparsely hirsute above,

conspicuously venose beneath, 2-4 cm. long, 7-16 mm. wide; racemes 7-12-

flowered, 2-4 cm. long, the pedicels 2.5 mm. long; flowers violaceous, 6 mm. long;

sepals lance-linear, hirsute, acute; wings obovate-oblong, very obtuse, sparsely

long-pilose dorsally; immature capsule elliptic-obovate, very hirsute.

We have seen nothing to represent this species.

Polygala variabilis HBK. Nov. Gen. & Sp. 5: 397. pi. 509.

1823. P. variabilis f. leucanthema Blake, Contr. Gray Herb. 47: 96.

1916. Cola de mico.

Open rocky slopes, sometimes in savannas, 1,500 meters or less;

Zacapa; Chiquimula; Jalapa; Jutiapa. Southern Mexico; British

Honduras; South America.

A slender erect annual, simple or sparsely branched, the stems obscurely

stipitate-glandular, especially below; leaves alternate, linear, 4-9 mm. long;

racemes short and thick, rather dense, 7-10 mm. broad, usually about as long as

broad, the axis sometimes elongating in age; flowers short-pedicellate, rose-purple

or sometimes white; sepals ovate, rounded or obtuse at the apex, 1-2 mm. long;

wings elliptic, 2.7-3.5 mm. long, rounded at the apex; keel cristate; capsule ovate-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 21

oblong, 2.5-3 mm. long; seed obconic, sericeous-comose, 2.7 mm. long, the aril

2-lobate, 0.5 mm. high, appressed.

In the typical form the flowers are rose-purple; in f. leucanthema

they are white. Both forms have been collected in Guatemala.

SECURIDACA Jacquin

Mostly woody vines, rarely erect shrubs; leaves alternate, entire, broad, short-

petiolate, generally with stipular glands; flowers rather large for the family, mostly

rose-colored, in terminal and axillary, often paniculate racemes; sepals 5, free,

deciduous, the 3 outer ones herbaceous, the 2 inner ones (wings) much larger,

petaloid; petals 3, deciduous, united at the base, the lowest (keel) boat-shaped,

unguiculate, with a subapical fimbriate crest, the 2 upper petals united with the

base of the stamen tube; stamens 8, the filaments united almost to the apex to

form a sheath; anthers confluently 1-celled, opening by a large introrse-apical

pore; disk a low fleshy ring at the base of the ovary; ovary 1-celled, the style

sickle-shaped, excavate at the apex, the stigma lobes 2, approximate; ovule

solitary, pendulous; fruit a 1-celled 1-seeded samara, with a large wing on the

lower side, sometimes marginate on the upper side or rarely almost equally winged;

seed glabrous, not arillate, the testa thin; endosperm none; cotyledons thick-

fleshy, oily.

About 30 species in tropical America, Africa, and Asia. Two

other Central American species have been found in Costa Rica and

Panama.

Pubescence of the lower leaf surface of closely appressed hairs S. diversifolia.

Pubescence of the lower leaf surface of spreading hairs S. sylvestris.

Securidaca diversifolia (L.) Blake, Contr. U. S. Nat. Herb. 23:

594. 1923. Polygala diversifolia L. Sp. PL 703. 1753. S. erecta Jacq.

Enum. PL Carib. 27. 1760. Bejuco anisillo (Pete"n, fide Lundell).

Moist or dry thickets, often in second growth, sometimes in

forest, 1,800 meters or less, mostly at 300 meters or lower; Pete*n;

Izabal; Zacapa; Chiquimula; Santa Rosa; Escuintla; Retalhuleu.

Mexico; British Honduras to Panama; Lesser Antilles; South

America.

A small or large, woody vine, the branches slender, strigose; leaves on very

short petioles, elliptic-oblong to broadly ovate or oval, 4-10 cm. long, 2-5.5 cm.

wide, obtuse or acute, rounded or cuneate at the base, chartaceous, sparsely or

densely strigillose on both surfaces, paler beneath, lustrous above, the venation

rather prominent and closely reticulate on both surfaces; racemes many-flowered,

simple or branched, 5-10 cm. long, the bracts lanceolate to ovate, acuminate,

deciduous; pedicels 4-7 mm. long, the flowers rose or purple; sepals oval, ciliate,

2.5-3.5 mm. long; wings 8-11 mm. long; body of the samara turgid, elevated-

reticulate, 5-8 mm. long, the wing obovate, 3-5 cm. long, 1-1.5 cm. wide near

the middle, strigose or glabrate.

22 FIELDIANA: BOTANY, VOLUME 24

This and the following are handsome and showy vines, often

producing large masses of beautiful, rose-colored or purple flowers.

Apparently the vines flower for only a short time. The flowers much

resemble those of Leguminosae, and most persons on seeing the plants

for the first time assume that they belong to that family.

Securidaca sylvestris Schlecht. Linnaea 14: 381. 1840.

Moist or dry thickets or mixed forest, often in second growth,

2,000 meters or less; Pete"n; Alta Verapaz; Zacapa; Chiquimula;

Jutiapa; Escuintla; Solola; Quezaltenango; San Marcos. Western

and southern Mexico; British Honduras; Honduras; Costa Rica.

A small or large, woody vine, the branches densely hirtellous or pilosulous;

leaves on very short petioles, ovate to oblong-elliptic, 3-7 cm. long, 1.5-3 cm. wide,

acute or obtuse, rounded or cuneate at the base, sparsely or densely pilosulous on

both surfaces with spreading hairs, lustrous above, paler beneath, the venation

prominent and closely reticulate on both surfaces; racemes lax or dense, 10 cm.

long or less, the bracts ovate or lance-ovate, deciduous, 1.5-4 mm. long, the pedicels

4-6 mm. long, the flowers rose-purple; sepals oval or rounded, 2.5-4 mm. long,

ciliate and pilosulous; wings 8-11 mm. long; body of the samara 7-9 mm. long,

reticulate, the wing obovate, 3-4 cm. long, 12-14 mm. wide, short-pilosulous.

This is perhaps the plant reported by Hemsley from Guatemala

as S. mollis HBK. on the basis of a Friedrichsthal collection. The

locality is not cited, and there is no certainty that the plant came

from Guatemala. S. mollis is a synonym of S. coriacea Bonpl., a

species ranging from South America north into Panama. In Salvador

S. sylvestris is called Coralmeca, and it is used there, with salt, in

treating certain diseases of cattle. The vine is said to be used there

also as a barbasco or fish poison. Seeds of these plants seem to be

spread widely by some means, for small sterile plants are common

in fields where no adult plants are found, and they often invade

cultivated fields.

DICHAPETALACEAE

Reference: H. A. Gleason, N. Amer. Fl. 25: 381-383. 1924.

Trees or shrubs, sometimes scandent; leaves alternate, 2-ranked, entire, mem-

branaceous or coriaceous, short-petiolate, penninerved; stipules small and narrow,

deciduous; inflorescence a loose or dense cyme, sometimes capitate, axillary, the

peduncle often adnate to the petiole; bracts small and narrow, deciduous; flowers

sessile or short-pedicellate, small and inconspicuous, perfect or unisexual, regular

or somewhat zygomorphic; receptacle flat or concave; sepals 5, imbricate, free

or slightly connate at the base, equal or nearly so; petals 5, alternate with the

sepals, free or connate into a short tube, equal or conspicuously unequal, often

bifid or bilobate and more or less involute or cucullate; stamens 5, alternate with

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 23

the petals, either free, hypogynous, and with slender complanate filaments some-

what dilated at the base, or epipetalous with nearly or quite sessile anthers; anthers

usually 5, sometimes 3, 2-celled, introrse, with a broad connective; hypogynous

disk unilateral in zygomorphic flowers, of 5 scales alternate with the stamens in

regular flowers; ovary superior, 2-3-celled; styles 2-3, united below, with recurved

stigmas; ovules 2 in each cell, suspended near the summit of the cell, anatropous;

fruit drupaceous, more or less compressed, the exocarp thin, leathery or fleshy;

seed usually 1, without endosperm.

Three genera, in the tropics of both hemispheres. Only the

following genus is known from continental North America.

DICHAPETALUM Thouars

Trees or shrubs, sometimes woody vines; leaves short-petiolate, usually mem-

branaceous; stipules linear or narrowly lanceolate; inflorescence usually laxly

cymose, with few or many flowers, arising from the axils of the upper leaves, the

peduncle adnate for part of its length to the petiole; flowers small, whitish, short-

pedicellate, perfect or unisexual, regular or nearly so; receptacle flat or somewhat

concave; sepals free or barely connate at the base, equal or subequal, spreading

or ascending; petals free, equal or nearly so, ascending or erect, short-unguiculate,

bifid or bilobate, the apex cucullate; stamens free and distinct, the filaments slender,

compressed, somewhat dilated below; anthers 5; hypogynous disk of 5 minute

scales.

About 80 species in the tropics of both hemispheres, most abun-

dant in Africa. Two other species are known in Central America,

from Costa Rica and Panama.

Branches of the inflorescence and the sepals hirsute or hispid with long stiff spread-

ing hairs; leaves hirsute beneath, the veins conspicuously impressed on the

upper surface D. bullatum.

Branches of the inflorescence and sepals short-pilose or puberulent; leaves velu-

tinous-pilose to glabrate beneath, the veins not or only slightly impressed on

the upper surface.

Leaf blades mostly oblong or lance-oblong, somewhat narrowed to a rounded

base, mostly 3-5 cm. wide, puberulent or sparsely pilose beneath, the hairs

of the costa often appressed D. chiapense.

Leaf blades mostly broadly obovate to oblong-obovate, usually acute, acutish,

or merely obtuse at the base, generally 7-13 cm. wide, densely velutinous-

pilose beneath with spreading hairs D. Donnell-Smithii.

Dichapetalum bullatum Standl. & Steyerm. Field Mus. Bot.

23: 169. 1944.

Moist or wet thickets or mixed forest, sometimes in Manicaria

swamps, 500 meters or usually at or near sea level; endemic; Izabal

(type collected along road between Puerto Barrios and Santo

Tomas, Steyermark 39874).

An erect or subscandent shrub, the branchlets thick, ochraceous, usually

lustrous, when young densely hispid with stiff, sordid or brownish hairs; leaves

24 FIELD IANA: BOTANY, VOLUME 24

very large, short-petiolate, thick-membranaceous, strongly bullate, the thick

petiole 5-8 mm. long, densely hispid; leaf blades elliptic or broadly elliptic, some-

times oblong-elliptic, 17-28 cm. long, 11-18 cm. wide, abruptly acute or acuminate,

rounded or subcordate at the base, sparsely hispid above or in age glabrate, the

nerves and veins conspicuously impressed, densely hirsute or hispid beneath

with rather long, stiff hairs, the veins elevated and laxly reticulate; inflorescence

small, terminal or pseudoterminal, branched from the base or composed of several

simple inflorescences, the primary branches very slender, 1-1.5 cm. long, densely

hispid, the flowers umbellate at the end of the peduncle, the long slender pedicels

almost filiform, hispidulous; sepals narrowly oblong, obtuse, 3-3.5 mm. long,

densely whitish-tomentulose outside and hispid with appressed stiff white hairs;

petals about as long as the sepals, bilobate at the apex, glabrous, white turning

blackish purple in drying; filaments very slender, glabrous, longer than the petals;

ovary densely white-tomentose.

Dichapetalum chiapense Standl. Field Mus. Bot. 17: 196.

1937.

Wet forest or thickets, 1,500 meters or less; Pete*n; Alta Verapaz;

Suchitepe"quez; Huehuetenango. Chiapas, the type from Mt. Ovando,

E. Matuda 679; British Honduras; Atlantic coast of Honduras.

Shrub or small tree as much as 8 meters high, sometimes scandent, the branch-

lets closely fulvous-tomentulose or glabrate; petioles stout, 5-8 mm. long, the

blades mostly oblong or lance-oblong, subcoriaceous, 8-13 cm. long and 2-3.5 cm.

wide or larger, usually gradually acuminate or long-acuminate, generally somewhat

narrowed to the rounded or very obtuse, often somewhat unequal base, dark

green above when dried, glabrous or somewhat tomentose along the costa, more

or less pilose beneath at first with chiefly appressed hairs, later glabrate, with

about 9 pairs of lateral nerves, the ultimate veins prominent and laxly reticulate;

inflorescence small, with few to many flowers, the branches densely fulvous-

tomentose; fruit 1-2-celled, when 1-celled broadly oval or subglobose, about 2 cm.

long and 1.5 cm. wide, densely fulvous-tomentose.

Dichapetalum Donnell-Smithii Engler, Bot. Jahrb. 23: 144.

1896. Symphyllanthus Donnell-Smithii Gleason, N. Amer. Fl. 25:

381. 1924.

Moist or dry thickets and forest, 1,800 meters or less, chiefly in

the Pacific bocacosta; endemic; Alta Verapaz; Santa Rosa; Es-

cuintla (type from Escuintla, J. D. Smith 2067); Sacatepe"quez ;

Chimaltenango; Retalhuleu; Quezaltenango; San Marcos.

Usually a shrub but sometimes a tree of 10 meters, the young branchlets

densely fulvous-tomentose or short-pilose; leaves on very short petioles, usually

very thin and bright green, oblong-obovate to broadly obovate, mostly 10-25 cm.

long and 7-13 cm. wide, sometimes larger, commonly obtuse or rounded at the

apex and abruptly acuminate, mostly acute or subacute at the base but rarely

somewhat rounded, sparsely or densely pilose above, beneath usually densely

velutinous-pilose, the lateral nerves mostly 5-7 pairs; cymes small and usually

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 25

few-flowered, the branches densely fulvous-pilose; sepals spreading, rounded-

ovate, obtuse, densely tomentose outside; petals shorter than the sepals, 2 mm.

long, bifid almost to the middle; fruit brownish, compressed-ellipsoid, 1.5-2 cm.

long, densely velutinous-pilose.

The species has been 'reported from Guatemala under the name

D. pedunculatum Baillon. The shrub is a common one at many

localities in the Pacific bocacosta or even far down upon the plains,

but it is inconspicuous, even when in flower, when it reminds one

somewhat of some of the shrubby Lauraceae. The plants of this

genus are best marked by the fact that the peduncles are united

with the petioles, a character not found in other Guatemalan plants.

EUPHORBIACEAE. Spurge Family

Trees, shrubs or herbs, sometimes scandent and twining, mostly with milky

sap; leaves chiefly alternate, sometimes opposite or verticillate, simple or rarely

digitately compound, sometimes palmate-lobate, dentate or entire; stipules often

present; inflorescence highly variable in form, the flowers usually small but some-

times large, unisexual, monoecious or dioecious, generally regular; perianth some-

times none, usually small, often dissimilar in flowers of the 2 sexes, either a calyx

or a calyx and a corolla, the segments free or united, imbricate or valvate in bud;

staminate flowers with an intrastaminal or extrastaminal disk, or this of separate

glands or lobes; stamens sometimes indefinite, often as many as the sepals or fewer,

sometimes only 1, the filaments free or united; rudimentary ovary present or none;

disk of the pistillate flower annual or cupular, or of separate glands, or absent;

ovary usually 3-celled, sometimes 1-4-celled or the cells rarely more numerous;

styles as many as the carpels, free or connate, entire, cleft, or laciniate; ovules

1 in each cell, or 2 and collateral, pendulous, anatropous, attached at the inner

angle of the cell, the raphe ventral; micropyle often covered with a caruncle, this

persistent and conspicuous on the seed; fruit generally capsular and separating

into 2-valvate cocci, these separating from the persistent axis or columella, or the

fruit drupaceous and indehiscent; seeds commonly as many as the ovules; endo-

sperm usually abundant and carnose, the cotyledons broad and flat, rarely thick

and carnose.

One of the largest families of plants, with more than 200 genera

and 7,000 species. Other genera besides those listed here are repre-

sented in southern Central America, mostly groups consisting of a

single species. The majority of the plants of the family are dis-

tinguished by the combination of milky sap and dry 3-celled fruit,

but there are numerous exceptions. The family includes many

plants of great economic importance, the most valuable being

Hevea, from which practically all commercial rubber is obtained.

In many species the sap is poisonous or at least highly irritant, and

the seeds often possess purgative properties, or in large amounts

are poisonous. The family has been monographed by Pax in the

26 FIELDIANA: BOTANY, VOLUME 24

Pflanzenreich, as noted on the following pages. The work has been

exceptionally well done, and all the genera have been covered except

a few of the very large ones, notably Croton, Phyllanthus, and

Euphorbia. The nomenclature in the groups that have not been

treated is, naturally, not altogether satisfactory, and it is likely

that numerous changes in names will have to be made when these

large genera are treated critically.

Flowers surrounded by a calyx-like or slipper-shaped involucre containing both

staminate and pistillate flowers; perianth none or minute.

Involucre calyx-like or cupular, regular Euphorbia.

Involucre slipper-shaped or shoe-shaped, very asymmetric Pedilanthus.

Flowers not involucrate or, if so, the involucre containing only staminate or

pistillate flowers, never shoe-shaped; perianth usually present and well

developed.

Ovules 2 in each cell; flowers apetalous or the petals, when present, usually

small and scale-like; flowers fasciculate or solitary in the leaf axils, rarely

spicate.

Fruit capsular; pubescence not lepidote.

Flowers partly in stiff spikes or racemes; leaves coriaceous Amanoa.

Flowers axillary and solitary or fasciculate, or in slender racemes or panicles;

leaves usually membranaceous.

Petals well developed, relatively large Astrocasia.

Petals minute or none Phyllanthus.

Fruit drupaceous; pubescence sometimes lepidote.

Pubescence lepidote Hieronyma.

Pubescence not lepidote Drypetes.

Ovules 1 in each cell; petals often well developed; flowers often in racemes,

spikes, or panicles.

Stamens in bud bent inward, the apex of the anther turned downward.

Flowers usually with petals, mostly in terminal racemes.

Sepals equal or nearly so, not appendaged Croton.

Sepals unequal, the outer ones of the pistillate flowers with conspicuous

stalked glands or appendages Julocroton.

Stamens straight in bud, the tips of the anthers turned upward.

Flowers in dichotomous cymes. Plants often armed with stinging hairs,

herbaceous or woody, never scandent.

Plants bearing few or usually very numerous stinging hairs . Cnidoscolus.

Plants without stinging hairs Jatropha.

Flowers variously arranged but never in dichotomous cymes.

Inflorescence subtended by 2 large, opposite, green, white, or reddish,

foliaceous bracts, these about as broad as long; plants twining herbs

or shrubs, or rarely low erect shrubs, the leaves often digitately

compound or deeply lobate Dalechampia.

Inflorescence not as described above.

Leaves deeply lobate.

Leaves peltate; calyx lobes valvate; stamens numerous. . . .Ricinus.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 27

Leaves usually not peltate; calyx lobes imbricate; stamens 10.

Manihot.

Leaves entire or dentate, rarely with one or two very shallow lobes.

Leaves mottled or spotted with red, yellow or white, entire or ob-

scurely lobate; cultivated shrubs, rarely escaping to thickets.

Codiaeum.

Leaves not mottled with other colors than green; chiefly native

plants.

Segments of the staminate calyx valvate in bud.

Stipules indurate and spinescent Ophellantha.

Stipules not spinescent, sometimes none.

Staminate flowers with well-developed petals.

Plants sericeous; capsule not tuberculate.

. Petals 8-12; stamens numerous Garcia.

Petals 4-5; stamens 5-15 Ditaxis.

Plants hirsute or hispidulous; capsule tuberculate.

Caper onia.

Staminate and pistillate flowers without petals.

Plants twining and scandent, rarely erect but then with

stinging hairs.

Capsule 4-celled; plants woody, without stinging hairs.

Plukenetia.

Capsule 3-celled; plants herbaceous throughout or nearly

so, usually with stinging hairs Tragia.

Plants not twining or scandent, without stinging hairs.

Anther cells elongate and narrow, often flexuous; flowers

spicate or racemose, sometimes subcapitate; herbs,

shrubs, or small trees Acalypha.

Anther cells short, globose or oblong; shrubs or trees.

Flowers solitary or fasciculate in the leaf axils . . . Adelia.

Flowers, at least the staminate ones, racemose, spicate,

or paniculate.

Staminate flowers paniculate Alchornea.

Staminate flowers racemose or spicate.

Pubescence of stellate hairs Bernardia.

Pubescence of simple hairs, or none Cleidion.

Segments of the staminate flowers imbricate or open in bud.

Leaves with scattered brown scales on the lower surface; flowers

enclosed in a globose involucre Pera.

Leaves without scales; flowers not enclosed in an involucre.

Petioles bearing conspicuous glands below the base of the

blade.

Leaves dentate Sapium.

Leaves entire or essentially so Tetrorchidium.

Petioles without glands, or the glands borne at the very apex.

Flowers paniculate.

Bracts foliaceous; panicles broad; leaves very large,

cordate Omphalea.

28 FIELDIANA: BOTANY, VOLUME 24

Bracts not foliaceous; panicles raceme-like; leaves small,

not cordate Mabea.

Flowers spicate or racemose.

Ovary 5-20-celled, the fruit very large; leaves glabrous.

Leaves cordate at the base; fruit dry Hura.

Leaves not cordate at the base; fruit fleshy . Hippomane.

Ovary normally 3-celled.

Leaves tomentose beneath; staminate flowers densely

crowded on the rachis Dalembertia.

Leaves glabrous; staminate flowers not densely

crowded.

Staminate calyx none or rudimentary . . Gymnanthes.

Staminate calyx well developed.

Calyx deeply 3-parted Sebastiania.

Calyx shallowly lobate Stillingia.

ACALYPHA L.

Reference: F. Pax & K. Hoffmann, Acalypha, Pflanzenreich IV.

147, xvi. 1924.

Annual or perennial herbs or more frequently shrubs or small trees; leaves

alternate, petiolate, bistipulate, mostly ovate, dentate, 3-5-nerved or penninerved,

often puncticulate; flowers monoecious or rarely dioecious, apetalous, small or

minute; staminate flowers glomerate within small bracts, short-pedicellate; pistil-

late flowers 1-5 within a conspicuous, often accrescent bract, or pedicellate in the

axis of a small, scarcely foliaceous bract; inflorescences unisexual or bisexual, the

staminate usually ament-like, slender, the pistillate inflorescence paniculate,

racemose, or usually spicate; androgynous spikes usually with pistillate flowers

below and staminate above; androgynous and pistillate inflorescences axillary or

terminal, the staminate ones always axillary; disk none; staminate calyx globose

in bud, in anthesis valvately 4-parted; stamens generally 8, the filaments free;

anther cells distinct, divaricate or pendulous, oblong or linear, in the open flower

flexuous- vermiform; pistillate sepals 3-5, connate at the very base or rarely

higher, small, imbricate; ovary usually 3-celled, often muricate; styles free or

short-connate, generally lacinulate; ovules solitary in the cells; capsule generally

small, tridymous, the cocci bivalvate; seeds small, subglobose, distinctly or

obsoletely carunculate, the testa crustaceous; endosperm carnose, the cotyledons

broad, flat.

Species almost or fully 400, in both hemispheres, chiefly in

tropical regions, very few extending into temperate areas. A few

other species are found in southern Central America.

Pistillate flowers pedicellate; pistillate bracts minute.

Leaves very glutinous on the upper surface, glabrous on both surfaces.

A. gummifera.

Leaves not glutinous-viscid.

Leaves broadly ovate, broadest at or near the base, palmate-nerved.

A. villosa.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 29

Leaves oblong to lance-oblong or elliptic-oblong, penninerved or essentially so.

A. costaricensis.

Pistillate flowers sessile; pistillate bracts mostly large and foliaceous. Shrubs,

trees, or herbs.

Plants herbaceous, mostly annual, sometimes perennial.

Lobes or teeth of the pistillate bracts short, oblong or ovate, obtuse, little if

at all exceeding the united portion of the bract.

Inflorescences all or mostly axillary, very lax and interrupted. A low

annual A. indica.

Inflorescences partly terminal, very dense.

Spikes mostly twice as long as broad or shorter; plants annual.

Stems bearing numerous gland-tipped hairs. . . . A. pseudoalopecuroides.

Stems without gland-tipped hairs A. Poiretii.

Spikes several times as long as broad; plants usually perennial.

Stems stipitate-glandular A. subviscida.

Stems without glandular pubescence.

Leaves small, mostly 3 cm. long or shorter, obtuse or acute.

A. phleoides.

Leaves all or mostly much larger, usually acuminate.

Leaves hirsute with long stiff spreading hairs A. triloba.

Leaves sparsely pilose or hispidulous with short hairs.

A. guatemalensis.

Lobes of the pistillate bracts linear or filiform, much longer than the united

portion of the bract.

Spikes 3 times as long as broad or usually shorter, mostly 3 cm. long or

less.

Spikes all axillary; leaves mostly obtuse A. arvensis.

Spikes partly terminal; leaves abruptly acuminate A. alopecuroides.

Spikes several to many times as long as broad, most of them much more

than 3 cm. long, at least at maturity.

Ovary and capsule pubescent.

Fruiting bracts cleft almost to the base into 7-13 slender, almost

subulate lobes A. setosa.

Fruiting bracts incisely 19-25-dentate to about the middle or somewhat

more deeply A. persimilis,

Ovary and capsule glabrous A. polystachya.

Plants shrubs or trees, woody throughout or nearly so, when shrubs usually tall

and more than a meter high.

Inflorescences terminal and axillary, the terminal spikes wholly pistillate, all

or nearly all the pistillate spikes terminal only.

Leaves penninerved, all or most of them broadest at or above the middle.

Branches hirsute with spreading fulvous hairs; leaves usually densely

pilose beneath A. lancetillae.

Branches glabrous or pubescent with very short, appressed hairs; leaves

glabrous beneath or nearly so.

Bracts pilose with gland-tipped hairs, equaling or longer than the

capsule A. Ferdinandi.

Bracts without gland-tipped hairs, shorter than the capsule . A . Skutchii.

30 FIELDIANA: BOTANY, VOLUME 24

Leaves palmate-nerved, ovate or usually very broadly ovate, broadest at

or near the base.

Lowest pistillate bracts leaf-like, much larger than the upper ones, some-

times 5 cm. long A. chlorocardia.

Lowest pistillate bracts not leaf-like, of about the same size as the

upper ones.

Leaves 7-12 cm. wide, coarsely crenate, densely and finely velutinous-

pilose beneath A. Mortoniana.

Leaves mostly 2-6 cm. wide, finely crenate, variously pubescent be-

neath or almost glabrous.

Pistillate spikes slender, much interrupted A. firmula.

Pistillate spikes dense, continuous.

Pistillate bracts 7-10-dentate A. Schiedeana.

Pistillate bracts mostly 11-15-dentate A. mollis.

Inflorescences all axillary.

Spikes bisexual, a few pistillate bracts present at the base of the spike which

consists mostly of staminate flowers A. diver sifolia.

Spikes unisexual, the staminate and pistillate flowers in separate spikes.

Pistillate spikes short and globose or oblong, or bearing only 1-3 or

rarely more bracts.

Pistillate spikes globose or subglobose, with very numerous bracts.

A. trachyloba.

Pistillate spikes consisting of only 1-few scattered bracts.

Pistillate spikes with mostly 3-6 remote bracts, or sometimes with

only 1-2 A. euphrasiostachys.

Pistillate spikes with only 1-2 bracts, mostly with only 1.

Bracts 9-11-laciniate A. unibracteata.

Bracts 13-17-laciniate A. leptopoda.

Pistillate spikes slender, elongate, linear or oblong-linear, bearing very

numerous bracts.

Pistillate bracts entire, or sometimes dentate but the leaves then

colored; cultivated plants.

Pistillate bracts entire; leaves green A. hispida.

Pistillate bracts dentate; leaves colored A. Wilkesiana.

Pistillate bracts dentate or laciniate; leaves green; native plants.

Branches densely glandular-pubescent A. Langiana.

Branches without glandular pubescence.

Leaves penninerved, broadest at or above the middle, glabrous

or nearly so A. Ferdinandi.

Leaves palmate-nerved, broadest at or near the base, glabrous or

densely pubescent.

Lowest bracts of the pistillate inflorescence leaf-like, much

larger than the upper ones, sometimes 5 cm. long.

A. chlorocardia.

Lowest bracts of the pistillate spikes of about the same size as

the upper ones.

Pistillate bracts 2-lobate at the apex; leaves glabrous or nearly

so A. tenuicauda.

Pistillate bracts not bilobate; leaves usually densely pilose,

sometimes almost glabrous A. macrostachya.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 31

Acalypha alopecuroides Jacq. Icon. PI. Rar. 3: 19. pi. 620.

1786-93.

Rocky mountain slopes with Juniperus, 1,350-1,500 meters;

Huehuetenango. Southern Mexico; British Honduras; Honduras;

Panama; West Indies; northern South America.

An erect annual, mostly 50 cm. high or less, simple or sparsely branched,

the stems pubescent and more or less glandular-pilose; leaves on slender petioles

2-6 cm. long, membranaceous, triangular-ovate or rounded-ovate, 3-7 cm. long,

acuminate or cuspidate-acuminate, rounded or subcordate at the base, crenate,

sparsely hirsute or pilosulous on both surfaces when young, glabrate in age,

palmate-nerved, the petioles glandular-pilose above; spikes terminal and axillary,

the terminal ones pistillate, in fruit sometimes 5 cm. long, 1-1.5 cm. wide, very

dense and many-flowered, the pistillate ones often bearing a few staminate flowers

at the apex, the axillary spikes wholly staminate or with a few pistillate flowers

at the base; pistillate bracts 7-9 mm. long, 3-5-lobate almost to the middle, the

lobes triangular-ovate, setaceous, long-pilose and glandular, 1-flowered; ovary

pilose, the styles 2-fid or entire; capsule 2 mm. long, the lobes dorsally carinate;

seeds 1 mm. long, narrowly ovoid.

The Maya name in Yucatan is recorded as "xmizbil"; names

reported from Salvador are "taba de polio," "gusanillo," and "tarco,"

but these may pertain rather to A. arvensis. The species is rare in

most parts of Central America, the majority of the collections re-

ported from that area as A. alopecuroides being rather A. arvensis.

Acalypha arvensis Poepp. & Endl. Nov. Gen. 3: 21. 1845.

Hierba del cancer; Gusanillo; Gusanito; Mata-gusano; Corrimiento

(Pete"n); Sajoi (Petatan, Huehuetenango); Ccul (Chimaltenango,

fide Tejada); Ztajnoy (Quiche", fide Tejada).

Moist or wet thickets, fields, or banks, often a weed in cultivated

or waste ground, frequent on sandbars along streams, 1,500 meters

or less; Pete"n; Alta Verapaz; Izabal; El Progreso; Zacapa; Santa

Rosa; Escuintla; Guatemala; Suchitepe"quez ; Retalhuleu; San

Marcos; Huehuetenango. Southern Mexico; British Honduras;

Honduras; Costa Rica; Panama; Martinique; tropical South

America.

Plants annual, erect or ascending, usually 50 cm. high or less, simple or

branched, the stems sometimes more elongate, procumbent, and rooting at the

lower nodes, spreading-pilose or glabrate, densely pubescent on the younger parts;

leaves membranaceous, on petioles 2-3.5 cm. long, rhombic-ovate or rhombic-

lanceolate, mostly 3-7 cm. long, acute or obtuse, obtuse at the base, palmate-

nerved, crenate-serrate, pilose on both surfaces with spreading or appressed hairs

or sometimes glabrate; spikes slender-pedunculate, axillary, androgynous, the

upper ones almost wholly pistillate, 1.5-2.5 cm. long, 10-13 mm. broad, bearing

a few staminate flowers at the apex; lower spikes almost wholly staminate, 2 mm.

32 FIELDIANA: BOTANY, VOLUME 24

thick; fruiting bracts 5 mm. wide, 4-7-lobate to the middle, the lobes triangular-

ovate, filiform-acuminate, hirsute, some of the hairs gland-tipped; styles lacinulate;

capsule 2 mm. broad, pilose; seeds broadly ovoid, 1 mm. long.

Called "gatito" in Yucatan; "espinosilla" (Oaxaca); "hierba del

gusano" (Veracruz). The plant is known everywhere in Guatemala

by the name "hierba del cancer," and it is much used in household

medicine. There is a general belief among the country people that

it is a remedy for "cancer" (of which they often have very vague

ideas), and it is used commonly in treating sores, cutaneous and

venereal diseases, and the bites of various poisonous animals.

Acalypha chlorocardia Standl. Field Mus. Bot. 8: 18. 1930.

Known only from the type, Middlesex, British Honduras, on

river bank, 60 meters, W. A. Schipp S-45.

A shrub a meter high, the young branches rather densely hirsute with long

spreading whitish hairs; stipules 1 cm. long, linear-subulate, glandular-denticulate;

leaves membranaceous, on petioles 12-16 cm. long, ovate or broadly ovate, 12-15

cm. long, 7-9 cm. wide, long-acuminate, rounded and shallowly cordate at the

base, closely appressed-serrate, sparsely hirsute above, thinly hispidulous beneath,

palmately 5-7-nerved at the base; terminal spike pistillate, 19 cm. long, rather

dense, the bracts numerous, the lowest ones resembling the leaves, as much as 7 cm.

long, narrowly long-acuminate, the upper ones sessile, cordate-clasping, 1 cm.

long, about 15-serrate, acuminate, appressed-hispidulous, the uppermost bracts

only 5 mm. long, acute, crenate-serrate; pistillate flowers sessile; ovary densely

hispidulous; style branches multilacinulate.

From the single sheet of this species it is difficult to decide whether

the pistillate spikes are really terminal. In general appearance the

plant resembles A. macrostachya.

Acalypha costaricensis (Kuntze) Knobloch in Just, Bot.

Jahresb. 19: 337. 1894. Ricinocarpus costaricensis Kuntze, Rev.

Gen. 615. 1891.

Moist or wet, usually dense, mixed forest or thickets, 2,000

meters or less; Izabal; Escuintla; Sacatepe"quez; Chimaltenango;

Suchitepe"quez; Quezaltenango; San Marcos. Chiapas; British

Honduras; Honduras; Costa Rica.

A slender shrub 1.5-5 meters high, the young branches green, terete, at first

densely pubescent or hirsute, soon glabrate; leaves thin, bright green, on slender

petioles 4-12 cm. long, oblong or elliptic-oblong, mostly 10-20 cm. long and 4-10

cm. wide, long-acuminate, obtuse or acute at the base or often rounded or cuneate

to a narrow, subtruncate or subcordate base, coarsely crenate-dentate, essentially

penninerved, glabrous or nearly so or often hirsute beneath or on both surfaces,

the lateral nerves 6-11 pairs; flowers monoecious or dioecious; pistillate inflores-

cence terminal, paniculate, usually lax and much-branched, often 20 cm. long,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 33

pedunculate, the branches hirsute or almost glabrous; pistillate bracts very small,

1-2-flowered, the pedicels 1.5 mm. long or more; ovary densely muricate; style

elongate, pinnate-lacinulate, usually purple-red.

A very common shrub in the Atlantic lowlands, often in second

growth. Material of this species has been reported from Guatemala

as A. Schlechtendaliana Muell. Arg., a species of southern Mexico

that does not reach Central America, so far as our material indicates.

The key characters used by Pax and Hoffman for separating these

two species are not reliable, but it is believed that both are good

species, separable on other characters.

Acalypha diversifolia Jacq. PI. Hort. Schoenbr. 2: 63. pi. 244.

1797. A. leptostachya HBK. Nov. Gen. & Sp. 2: 96. 1817. A.

diversifolia var. leptostachya Muell. Arg. in DC. Prodr. 15, pt. 2: 854.

1866. A. tabascensis Lundell, Lloydia 4: 51. 1941. Cacucup (Alta

Verapaz); Cliche (Maya); Palo de sangre (Pete"n, fide Lundell).

Moist or wet thickets or forest, often in second growth thickets,

sometimes in open pine forest, 1,000 meters or less; Pete"n; Alta

Verapaz; Izabal. Southern Mexico; British Honduras to Panama;

tropical South America.

Usually a rather slender shrub of 1.5-3 meters, sometimes a tree of 6 meters,

the branches often elongate and recurved, when young villous or appressed-pilose,

in age glabrate and brown or reddish brown; leaves thick-membranaceous, on

petioles 1-2 cm. long or rarely longer, oblong-lanceolate to oblong-ovate, mostly

7-15 cm. long, acuminate or long-acuminate, obtuse at the base, serrate or crenate,

penninerved, with 6-9 pairs of lateral nerves, velutinous-pubescent or glabrate;

stipules 5-6 mm. long, linear-setaceous from a broad base; flowers monoecious;

spikes axillary, staminate or androgynous and then with 1-2 pistillate bracts at

the base, 5-11 cm. long, 2-4 mm. thick, sessile or subsessile, the staminate portion

of the spike often deciduous, or the inflorescence wholly pistillate and consisting

of 1 or few crowded pistillate bracts; pistillate bracts obtuse or acute, shallowly

few-dentate, 1-3-flowered, in fruit 4-6 mm. broad; ovary muricate, hispidulous;

styles pinnately lacinulate; capsule almost 3 mm. broad; seeds 1.5 mm. long,

minutely puncticulate.

Known in Honduras as "costilla de caballo" and "costilla de

danto"; "tapa-camino" (Veracruz). This is a very common shrub

of second growth thickets along almost the whole Atlantic coast of

Central America.

Acalypha euphrasiostachys Bartlett, Proc. Amer. Acad. 43:

55. 1907.

Known in Guatemala only from the type, Zacapa, Dept. Zacapa,

185 meters, C. C. Deam 190. Also in the State of Mexico, Mexico.

34 FIELDIANA: BOTANY, VOLUME 24

A slender shrub 1-2 meters high, the young branches hispidulous and puberu-

lent, in age glabrate and dark reddish brown; leaves membranaceous, on slender

petioles 1-2 cm. long, ovate or elliptic-ovate, 3-8 cm. long, 2-4.5 cm. wide, abruptly

acuminate, obtuse to subcordate at the base, serrate, palmately 3-5-nerved, thinly

or rather densely soft-pilose on both surfaces with rather short, spreading, soft

hairs; spikes all axillary, the staminate about 1 cm. long, lax and interrupted,

sessile; pistillate spikes 2-7 cm. long, the bracts remote, usually 3-7, sometimes

only 1-2, about 8 mm. long and 10 mm. wide, 1-flowered, about 13-dentate, with

unequal subacuminate teeth, hispidulous and glandular-pubescent; ovary densely

pilose; styles multilacinulate.

A little-known plant, of somewhat uncertain status. Although

placed by Pax and Hoffmann far apart from A. leptopoda, it is

actually closely related to that species.

Acalypha Ferdinand! K. Hoffm. Pflanzenreich IV. 147, xvi:

63. 1924.

Moist or wet, mixed forest, often on limestone, 1,300 meters or

less; Alta Verapaz (type from Cubilgiiitz, Tuerckheim 11.187);

Izabal. Atlantic lowlands of Honduras; Costa Rica.

A slender shrub 2 meters high, or sometimes a tree of 7 meters, the branches

slender, usually glabrous; leaves firm-membranaceous, on slender petioles 1.5 cm.

long or shorter, obovate-oblong to oblanceolate or lanceolate, usually broadest

above the middle, 8-18 cm. long, 2.5-7 cm. wide, rather abruptly acuminate or

caudate-acuminate, attenuate to a narrow, truncate or cordate base, serrate,

penninerved, usually glabrous, the lateral nerves 7-10 pairs; stipules 5-10 mm.

long, setaceous-filiform, rigid; flowers monoecious, the spikes axillary, 2.5-7 cm.

long, short-pedunculate, the staminate very dense; pistillate spikes mostly in the

upper leaf axils, sometimes terminal, lax, in fruit as much as 15 cm. long, the

bracts remote, in fruit as much as 1 cm. long and wide, rounded-ovate, acute,

1-2-flowered, puberulent and stipitate-glandular, 13-15-dentate, the teeth short,

acute or acuminate; ovary muricate, hirtellous and often stipitate-glandular;

styles multilacinulate.

Called "costilla de danto" in Honduras. This has been recorded

from Guatemala as A. cuneata Muell. Arg. var. obovata Muell. Arg.,

a quite different South American species.

Acalypha firmula Muell. Arg. Linnaea 34: 21. 1865 (type

from Salvador). A. porphyrantha Standl. Journ. Arnold Arb. 11:

32. 1930 (type from Siguatepeque, Comayagua, Honduras). Hierba

de San Antonio (fide Aguilar).

Usually in moist or dry, pine or oak forest, 1,100-2,000 meters;

Chiquimula; Jalapa; Guatemala. Honduras; Salvador.

A slender shrub 1-3 meters high, the branches pilose with spreading hairs or

almost glabrous, purplish-red or ferruginous in age; leaves firm-membranaceous,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 35

on slender petioles 1-7 cm. long, ovate to broadly ovate or oblong-ovate, 3-11

cm. long, mostly abruptly long-acuminate, rounded and shallowly cordate at the

base, crenate-serrate, glabrous or sometimes rather densely soft-pilose; staminate

spikes axillary, sessile, dense, short; pistillate spikes terminal, subsessile, very lax

and interrupted, the bracts in fruit only 3 mm. long, shallowly about 11-dentate,

stipitate-glandular; styles bright purple-red, showy; ovary muricate and hirtellous;

seeds almost 2 mm. long.

The leaves frequently are deep purple on the lower surface.

Acalypha guatemalensis Pax & Hoffm. Pflanzenreich IV. 147,

xvi: 27. 1924. Hierba del c&ncer.

Moist or dry fields or thickets, sometimes in rather open forest,

especially of oak or Alnus, or on open banks, frequently a weed in

cultivated ground, 750-2,500 meters; Baja Verapaz; Jalapa; Santa

Rosa; Guatemala; Sacatepe"quez; Chimaltenango; Solola; Quiche";

Huehuetenango (type from Jacaltenango, Seler 3261) ; Quezaltenango.

Honduras.

Plants herbaceous, usually perennial but sometimes annual, erect or ascending,

sometimes a meter high but usually lower, simple or branched, mostly erect, some-

times decumbent, when young puberulent or pilosulous with ascending or sub-

appressed hairs; leaves on petioles 3 cm. long or usually shorter, rounded-ovate

or rhombic-ovate, 4-7 cm. long, acuminate or acute, obtuse to usually broadly

rounded at the base, crenate, membranaceous, 5-nerved, thinly pilose along the

nerves and veins or sometimes rather densely and softly pubescent, in age often

glabrate; flowers monoecious, the spikes mostly androgynous, terminal and axillary,

generally numerous, the larger ones 4-5 cm. long or more, very dense, many-

flowered, pedunculate or subsessile; staminate portion of the spike short, dense;

pistillate bracts in fruit 5 mm. broad, 5-7-lobate to the middle, setose and bearing

short gland-tipped hairs, 1-2-flowered, the lobes lanceolate; ovary hirtellous;

styles pinnately 6-10-lacinulate, purple-red; capsule tuber culate, 3 mm. in diam-

eter; seeds ovoid, smooth, 2 mm. long.

This has been reported from Guatemala as A. alopecuroides Jacq.

Acalypha gummifera Lundell, Contr. Univ. Mich. Herb. 4:

10. 1940.

Wet mixed forest, often or usually on limestone, 150-875 meters;

Pete"n (type from Camp 34, British Honduras boundary, W. A.

Schipp 1290); Alta Verapaz; Izabal. British Honduras.

A slender shrub 1-2.5 meters high, glabrous throughout; leaves firm-mem-

branaceous, on slender petioles 1-4 cm. long, lanceolate or narrowly lanceolate,

8-14 cm. long, 1-4 cm. wide, narrowly long-acuminate, narrowed to the narrowly

rounded base, penninerved or somewhat 3-nerved at the base, remotely and incon-

spicuously serrulate, very lustrous and glutinous-viscid on the upper surface,

somewhat paler beneath, sparsely barbellate beneath in the axils of the nerves,

the lateral nerves 5-6 pairs; flowers monoecious, the spikes axillary, the staminate

36 FIELDIANA: BOTANY, VOLUME 24

ones 3-6.5 cm. long, very dense; pistillate flowers paniculate, the panicles as much

as 6 cm. long, with sparse filiform branches; capsule almost 3 mm. long, glabrous;

seeds reddish brown, ovoid, 2 mm. long.

The species is well marked by the very lustrous, glutinous upper

surface of the leaves, a character found in no other local species.

Acalypha hispida Burm. Fl. Ind. 203. pi. 61, f. 1. 1768.

Perhaps native of the islands of the South Pacific, but grown for

ornament in most tropical regions; planted commonly in Guatemala,

mostly in the tierra caliente, but also in such places as Guatemala

and Antigua.

A shrub, the stout branches tomentulose at first; leaves slender-petiolate,

broadly rhombic-ovate, 9-15 cm. long, cuspidate-acuminate, cuneately narrowed

at the base, firm-membranaceous, serrate, glabrate, 3-nerved at the base, penni-

nerved above; flowers dioecious; pistillate spikes axillary, pendent, 30 cm. long

or less, very dense, the style branches red or purple-red; bracts small, ovate-

lanceolate, entire, pubescent.

Sometimes called "chenille plant" in English; "cola de zorro"

(Salvador); "nemiz" (Maya); "cola de gato" (Yucatan). The very

numerous, large, thick, drooping, bright red or purple-red flower

spikes are very showy and ornamental.

Acalypha indica L. Sp. PI. 1030. 1753.

The typical form of this species is widely distributed in the Old

World tropics. In tropical North America it is represented by the

following variety:

Acalypha indica var. mexicana (Muell. Arg.) Pax. & Hoffm.

Pflanzenreich IV. 147, xvi: 35. 1924. A. mexicana Muell. Arg.

Linnaea 34: 41. 1865.

Moist or wet fields or thickets, usually a weed in waste or culti-

vated ground, 1,200-2,400 meters; Alta Verapaz; Guatemala;

Sacatepe"quez; Chimaltenango; Huehuetenango; Quezaltenango.

Southern Mexico; Costa Rica.

A slender annual, erect or decumbent, simple or usually branched, often

branched from the base, mostly 50 cm. high or lower, the stems sparsely pubescent

when young, soon glabrate; leaves on long slender petioles, thin, ovate or rhombic-

ovate, 2-6 cm. long, acute or usually obtuse, cuneate or rounded at the base,

crenate, glabrous or nearly so in age, 5-nerved at the base; spikes axillary, andro-

gynous, solitary or geminate, mostly very short, the staminate portion 1 cm. long

or less; pistillate bracts 1-4, often remote, foliaceous at maturity, suborbicular,

6-12 mm. broad, dentate, 1-2-flowered, sparsely setulose-pilose on the nerves;

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 37

ovary pilose, the style short, sparsely lacinulate; capsule short-pilose, 2 mm.

broad; seeds broadly ovoid, 1.5 mm. long, minutely puncticulate.

The occurrence of this plant in Guatemala is such that it may be

presumed to be of foreign origin. It is rarely found except in culti-

vated ground or in the immediate vicinity of settlements.

Acalypha lancetillae Standl. Field Mus. Bot. 4: 312. 1929.

Wet mixed lowland forest, at or little above sea level; Izabal.

British Honduras; Atlantic coast of Honduras (type from Lancetilla

Valley near Tela).

A shrub or small tree 1-6 meters high, sparsely branched, the branches densely

pilose with soft spreading hairs, the older ones pale brown; stipules setaceous

7-13 mm. long; leaves thin, on petioles 1-4 cm. long, oblong to obovate-oblong

or oblanceolate, mostly 7-17 cm. long and 3-7 cm. wide, short-acuminate or long-

acuminate, gradually narrowed below to the narrow, obtuse to shallowly cordate

base, closely serrate, hirsute or hirtellous above, densely velutinous-pilose beneath,

penninerved, the lateral nerves about 11 pairs; flowers monoecious, the staminate

spikes dense, slender, axillary, subsessile, mostly 5-8 cm. long, 2 mm. thick;

terminal spike about 7 cm. long and 1 cm. thick, the terminal portion caudiform,

staminate, dense, the pistillate portion about 4.5 cm. long, short-pedunculate;

pistillate bracts as much as 7 mm. long, cleft into about 11 linear-subulate lobes

extending almost to the base of the bract, densely hispidulous, eglandular, the

axillary pistillate spikes reduced to usually a single sessile bract; styles much

elongate, with very numerous capillary branches; capsule hispidulous.

Acalypha Langiana Muell. Arg. Linnaea 34: 159. 1865.

In canyon, 1,100 meters, Guatemala (Fiscal, C. C. Deam 6108).

Southern Mexico.

A slender shrub about a meter high, the branches densely short-pilose and

bearing numerous short gland-tipped hairs; leaves slender-petiolate, thin, ovate

or lance-ovate, 4-7 cm. long, acuminate, rounded at the base, crenate, 5-nerved

at the base, glabrate above, densely velutinous-pilose beneath with very short

hairs; stipules setaceous; flowers monoecious, the spikes axillary, unisexual, short-

pedunculate; pistillate spikes laxly flowered, the bracts 3 mm. long, 5 mm. broad,

reniform, about 2-flowered, rounded-obtuse, densely glandular-puberulent, shal-

lowly 9-13-dentate, the teeth triangular, acute; ovary somewhat muricate, pubes-

cent; styles pectinately 6-9-lacinulate; seeds minutely foveolate-puncticulate.

The available material of this species is scant, and the proper

position of the single Guatemalan collection is somewhat uncertain.

Acalypha leptopoda Muell. Arg. Linnaea 34: 39. 1865.

A slender shrub 1-3.5 meters high, the indument of the stems and leaves

variable; leaves on petioles 1-6 cm. long, membranaceous or thick-membranaceous

and rather rigid, lance-ovate to broadly ovate, mostly 4-10 cm. long, acuminate

or long-acuminate, rounded or shallowly cordate at the base, serrate, the young

38 FIELDIANA: BOTANY, VOLUME 24

leaves pubescent on both surfaces, 3-nerved at the base, penninerved above the

base; stipules 5-10 mm. long, subulate-filiform; flowers monoecious, the spikes

axillary, the staminate ones 3-6 cm. long, pedunculate, dense; pistillate spikes

consisting of 1 or 2 bracts, borne on a very slender peduncle 2-3.5 cm. long; fruit-

ing bracts about 8 mm. broad, 1-flowered, orbicular, incised-dentate almost to the

base, the 13-17 teeth lanceolate, acute, sometimes sparsely and minutely stipitate-

glandular; ovary pubescent; styles pectinately multilacinulate; capsule slightly

muricate; seeds 1.5 mm. long, puncticulate.

The species as treated by most authors consists of the two

following varieties:

Acalypha leptopoda var. glabrescens Muell. Arg. in DC.

Prodr. 15, pt. 2: 824. 1866. A. Lotsyi Donn. Smith, Bot. Gaz. 20:

544. 1895 (type from Pansamala, Alta Verapaz, Tuerckheim 1242).

Tejedor; Lolosan, Loasdm (Alta Verapaz, Quecchi); Canilla de venado.

Dry to wet thickets or rather thin forest, sometimes in pine

forest, 2,100 meters or less; Alta Verapaz; Baja Verapaz; Chiquimula;

Jalapa; Jutiapa; Santa Rosa; Guatemala; Sacatepe"quez; Chimal-

tenango; Retalhuleu; Quezaltenango; San Marcos; Huehuetenango.

Southern Mexico; Honduras and Salvador to Panama.

Leaves and stems glabrate, or the leaves often quite glabrous at maturity.

Around Coban the plant is used commonly in domestic medicine,

as a lotion for treating burns, infected cuts, and various skin affec-

tions, and as a shampoo for the hair.

Acalypha leptopoda var. mollis Muell. Arg. in DC. Prodr. 15,

pt. 2: 824. 1866. Bisic (Coban, Quecchi).

Dry to wet thickets, often in open or dense, moist or wet forest,

600-2,300 meters; Alta Verapaz; El Progreso; Jutiapa; Santa Rosa;

Guatemala; Quiche"; Huehuetenango. Southern Mexico; Honduras;

Costa Rica.

Leaves densely velutinous-pilose, especially beneath, the pubescence persistent

in age.

The two varieties probably are not systematically important,

although the plants differ visibly in appearance. They do not have

distinctive ranges in Guatemala.

Acalypha macrostachya Jacq. PI. Hort. Schoenbr. 2: 63. pi.

245. 1797.

A stout shrub 1-4.5 meters high, variable in pubescence, the branches usually

thick and with large pith; leaves membranaceous, on petioles 5-25 cm. long, ovate

to broadly ovate or triangular-ovate, 10-25 cm. long, 6-18 cm. wide, usually

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 39

abruptly acuminate or caudate-acuminate, rounded and often cordate at the

base, serrate, palmate-nerved; stipules 1-1.5 cm. long, linear-lanceolate from an

ovate base, glandular-ciliate, persistent; flowers monoecious or perhaps sometimes

dioecious; spikes axillary, as much as 40 cm. long, sessile or short-pedunculate;

staminate spikes dense, up to 5 mm. in diameter; pistillate spikes dense or lax,

sometimes with staminate flowers at the apex; pistillate bracts very numerous,

broadly ovate, shallowly 13-27-dentate, 1-flowered, the teeth triangular, acuminate,

in fruit 5-7 mm. wide; ovary hispid, the styles purple-red, 10-20-lacinulate; capsule

almost 4 mm. broad, pilose; seeds 2 mm. long, minutely puncticulate.

The species, as treated by Pax and Hoffmann, includes the follow-

ing varieties, separated on the basis of pubescence, and not sharply

separable. Still another variety is reported by the same authors

from Peru and Bolivia.

Acalypha macrostachya var. hirsutissima (Willd.) Muell.

Arg. Linnaea 34: 11. 1865. A. hirsutissima Willd. Sp. PI. 4: 528.

1805. A. sidaefolia HBK. Nov. Gen. & Sp. 2: 95. 1817. A. macro-

stachya var. sidaefolia Muell. Arg. Linnaea 34: 11. 1865. Comida

de venado (Quezaltenango) ; Chichicaste de agua (fide Aguilar); Chi-

chicaste (Santa Rosa; probably an erroneous name); Sesic (Que-

cchi).

Wet to dry, brushy hillsides or ravines or moist or wet forest,

2,000 meters or less, most frequent at about 1,000 meters; Alta

Verapaz; Baja Verapaz; Izabal; Santa Rosa; Escuintla; Guatemala;

Huehuetenango; Quezaltenango; San Marcos. Southern Mexico;

British Honduras; Honduras and Salvador to Panama; tropical

South America.

Young branches and petioles usually very densely pilose; leaves densely

velutinous-pilose on the lower surface, the pubescence persistent in age.

Sometimes called "shuampa" in Salvador. This has been re-

ported from Guatemala as A. caucana Muell. Arg., a South American

species. Both these varieties have approximately the same distribu-

tion in Guatemala and are of apparently little or no taxonomic

importance.

Acalypha macrostachya var. macrophylla (HBK.) Muell.

Arg. in Mart. Fl. Bras. 11, pt. 2: 345. 1874. A. macrophylla HBK.

Nov. Gen. & Sp. 2: 96. 1817.

Moist or wet thickets or rather open, mixed forest, often on steep

rocky hillsides, 250-1,500 meters; Alta Verapaz; Santa Rosa;

Escuintla; Sacatepe"quez; Chimaltenango; Suchitepe'quez ; Retal-

huleu; Quezaltenango; San Marcos. Honduras to Panama; tropical

South America.

40 FIELDIANA: BOTANY, VOLUME 24

Branches and petioles sparsely or rather densely hirsute or pubescent; leaves

in age glabrate except on the nerves.

Acalypha mollis HBK. Nov. Gen. & Sp. 2: 94. 1817.

At 1,200-1,500 meters; Baja Verapaz (Panzal); Guatemala.

Southern Mexico.

A shrub, the branches stout, densely velutinous-pilose; leaves membranaceous,

on slender petioles 1-4 cm. long, broadly ovate to lance-ovate, mostly 7-11 cm.

long, narrowly long-acuminate or cuspidate-acuminate, rounded or very obtuse

at the base, serrate-dentate, 5-nerved at the base, usually very densely soft-pilose,

especially beneath; flowers monoecious, the staminate spikes axillary, 3-5 cm.

long, 2-3 mm. thick, pedunculate, very dense; pistillate spikes terminal and in

the axils of the uppermost leaves, pedunculate, very dense, 1 cm. thick, short or

usually much elongate; pistillate bracts reniform-ovate, 11-15-dentate, 2-3-

flowered; ovary villous-pubescent; styles 8-12-lacinulate.

Acalypha Mortoniana Lundell, Bull. Torrey Club 64: 552.

1937.

Limestone thickets or open forest, 300 meters or less; Pete"n

(type from Uaxactun, H. H. Bartlett 12740). British Honduras.

A shrub of 1.5-4 meters, the branchlets thick, pubescent with mostly sub-

appressed, ochraceous hairs; stipules setaceous, 5 mm. long; leaves on slender

petioles 11 cm. long or less, membranaceous, ovate or elliptic-ovate, 10-25 cm.

long, 4-13 cm. wide, acuminate, rounded and usually shallowly and narrowly

cordate at the base, coarsely crenate, palmately 5-nerved, at first finely velutinous-

pubescent on both surfaces, glabrate in age; flowers monoecious, the staminate

spikes axillary, 5-16 cm. long; pistillate spikes terminal, 10-12 cm. long, many-

flowered, the bracts reniform-cordate, 5-6 mm. long, 6-8 mm. wide, very shallowly

dentate, glandular-pilose, in fruit 9 mm. long and 16 mm. wide, with about 10

acute teeth; ovary sparsely hirsute; capsule 5 mm. long, very sparsely pilose or

almost glabrous; seeds smooth, 4 mm. long.

Acalypha persimilis Muell. Arg. Linnaea 34 : 25. 1865. Chum-

pito.

Moist thickets, dry rocky slopes, on sandbars along streams, or

a weed in waste ground, 200-1,375 meters; Jalapa; Jutiapa; Santa

Rosa; Escuintla; Huehuetenango. Southern Mexico; Greater

Antilles.

An erect annual, 50 cm. high or less, usually branched, sometimes simple,

the stems densely pubescent at first; leaves membranaceous, on slender petioles

2-7 cm. long, ovate, 3-7 cm. long, 2.5-4.5 cm. wide, acute or short-acuminate,

rounded and usually more or less cordate at the base, closely and finely serrulate,

5-nerved at the base, thinly pubescent when young, in age glabrate, puncticulate-

scabrous; flowers monoecious, the staminate spikes axillary, 2 cm. long or less,

slender-pedunculate; pistillate spikes terminal and in the upper leaf axils, the

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 41

terminal ones 3-4 cm. long, in fruit about 5 mm. thick, many-flowered, often lax;

pistillate bracts reniform-orbicular, 4-5 mm. broad, rather shallowly 19-21-dentate,

the teeth narrowly triangular, scabrous and sometimes pilose, 1-flowered; styles

short, 3-4-lacinulate; capsule 3 mm. broad, papillose-hirtous; seeds 2 mm. long,

rugose-tuberculate.

Acalypha phleoides Cav. Icon. PL 6: 42. pi. 569, f. 2. 1801.

Hierba del cancer; Hierba del est6mago (fide Aguilar).

Usually in moist or dry, open, grassy, pine-oak forest, often in

rocky places or in open fields or hillsides, 750-2,100 meters; Jalapa;

Santa Rosa; Guatemala; Sacatepe"quez; Quiche"; Huehuetenango.

Mexico.

A perennial herb, usually from a thick woody root, the stems 50 cm. long or

less, often numerous, erect to procumbent, puberulent or hirsute, little branched;

petioles mostly less than 1 cm. long; leaf blades ovate or elliptic, 2-4 cm. long, 1-3

cm. wide, acute or obtuse, usually obtuse at the base, crenate-serrate, generally

very finely so, 3-5-nerved, pilose or glabrate, punctate-scabrous; spikes andro-

gynous, terminal and axillary, staminate above, or the axillary spikes often

wholly staminate; terminal spikes 2-9 cm. long, the pistillate portion usually

dense; pistillate bracts 1-2-flowered, suborbicular, 6-8 mm. long, coarsely 5-7-

dentate, the teeth broadly triangular, acute, hispidulous or pubescent; ovary

hirsute above, the styles purple-red, 6-8-lacinulate; capsule 2 mm. long; seeds

subglobose, 1.5-2 mm. long, fuscous in age.

Acalypha Poiretii Spreng. Syst. 3: 879. 1826. A. yucatanensis

Millsp. Field Mus. Bot. 1: 371. 1898.

Moist thickets, 200-500 meters; reported by Pax and Hoffmann

as collected in Guatemala by Friedrichsthal, the locality not in-

dicated. Southern Mexico; tropical South America.

An erect annual, generally 50 cm. high or less, slender, branched, the stems

hirsute or pilose; leaves thin, on slender petioles 4 cm. long or shorter, rhombic-

ovate to oblong-ovate, 3-6 cm. long, acute or acuminate, rounded at the base,

crenate-serrate, thinly or densely pilose with soft spreading hairs; spikes andro-

gynous, terminal and axillary, the upper ones very dense, oblong, chiefly pistillate,

the staminate portion very small and hidden among the pistillate bracts, the lower

spikes staminate, with 1-2 pistillate bracts at the base; pistillate bracts 7 mm.

broad, 7-9-fid for one-fifth their length, 1-flowered, densely pilose or hirsute;

styles simple; capsule 2 mm. broad, hirsute and tuberculate near the apex; seeds

narrowly ovoid, foveolate-puncticulate.

Acalypha polystachya Jacq. PI. Hort. Schoenbr. 2: 64. pi. 246.

1797. A. Matudai Lundell, Contr. Univ. Mich. Herb. 4: 10. 1940

(type from Chiapas).

Moist thickets or fields, sometimes on sandbars along streams,

or a weed in cultivated ground, 400 meters or less; Zacapa; Suchi-

tepe"quez; San Marcos; Huehuetenango. Mexico; Costa Rica.

42 FIELDIANA: BOTANY, VOLUME 24

An erect annual, the stems somewhat succulent, a meter high or less, stout,

mostly simple, often fistulous, puberulent when young; leaves on very slender

petioles 4-12 cm. long, thin, ovate or broadly ovate, about 10 cm. long and 6-9

cm. wide or often smaller, acute or abruptly short-acuminate, broadly rounded

at the base, finely and closely serrate, thinly pilose on the upper surface or glabrate,

somewhat paler beneath, glabrate, punctate, palmately 3-5-nerved; flowers

monoecious, the staminate spikes axillary, 4 cm. long or less, slender, dense;

pistillate spikes mostly terminal, in fruit as much as 15 cm. long and 1 cm. broad,

dense or often interrupted below; pistillate bracts 9-11-parted nearly to the base,

the segments almost setaceous, sparsely stipitate-glandular, 1-flowered, in fruit

1 cm. long; ovary glabrous, the styles 2-4-fid; capsule 4-5 mm. in diameter;

seeds ovoid, almost 3 mm. long, acute, scrobiculate-roughened.

Called "equilite" in Veracruz. It is reported from Chiapas that

the plant is sometimes eaten, presumably as a pot herb.

Acalypha pseudoalopecuroides Pax & Hoffm. Pflanzenreich

IV. 147, xvi: 86. 1924.

Moist brushy slopes or in quebradas, 200-500 meters; Zacapa.

Southern Mexico; Honduras.

Plants annual, 50 cm. high or less, erect, usually with numerous spreading

branches, the stems densely pilose and glandular-hirsute, often villous at the base;

leaves on slender petioles 1-2 cm. long, thin, ovate or broadly ovate, 2-4.5 cm. long,

1.5-3 cm. wide, acute or acuminate, rounded and often shallowly cordate at the

base, crenate, 5-nerved at the base, sparsely or densely long-pilose on both surfaces,

glabrate in age, puncticulate, usually more or less glandular-pilose; flowers monoe-

cious, the staminate spikes terminal, 1 cm. long, slender, pedunculate; pistillate

spikes axillary, 1.5-2.5 cm. long, 1 cm. broad, pedunculate, very dense, usually

unisexual, many-flowered; pistillate bracts shallowly about 7-dentate, the teeth

acute, densely glandular-pilose; ovary long-pilose, the style short, simple; capsule

pubescent, 2.5 mm. broad; seeds 1.5 mm. long.

This has been reported from Honduras as A. Poiretii Spreng.

Although A. Poiretii and A. pseudoalopecuroides are placed far apart

in their monograph by Pax and Hoffmann, because of the disposition

of the inflorescences, the two plants are almost exactly alike in

general appearance.

Acalypha Schiedeana Schlecht. Linnaea 7: 384. 1832.

Moist or dry thickets on hillsides or along streams, often in

rocky places, 200-1,350 meters; Zacapa; Chiquimula; Jalapa; Baja

Verapaz; Guatemala; Huehuetenango. Southern Mexico.

A slender, much-branched shrub 1-3 meters high, the branchlets densely

pubescent or glabrate; leaves thin, on slender petioles 1-5 cm. long, broadly ovate

to lance-ovate, 5-13 cm. long, acute or acuminate, often abruptly so, usually

rounded at the base and often cordate, crenate-dentate, varying from densely and

softly pubescent to almost glabrous, 3-5-nerved at the base; stipules setaceous,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 43

1 cm. long or less; staminate spikes axillary, 3 cm. long or less, slender, subsessile,

dense; pistillate spikes terminal, pedunculate, 3-10 cm. long, usually dense and

many-flowered; fruiting bracts 5-10 mm. broad, truncate, 7-11-dentate, the teeth

triangular or lanceolate, acute, 1-flowered, pubescent and stipitate-glandular,

sometimes glabrate; ovary hirsute, muricate; styles 5-10-lacinulate; capsule 3 mm.

broad; seeds broadly ovoid, 1.5-2 mm. long.

Acalypha septemloba Muell. Arg. was described from Guatemala

on the basis of Friedrichsthal 1354, and is reported from the same

country by Pax and Hoffmann. The type actually came, according

to the original label, from Cartago, Costa Rica.

Acalypha setosa A. Rich, in Sagra, Hist. Cuba 3: 204. 1850.

Corrimiento (Pete*n).

Moist or wet thickets, often a weed in waste ground or in fields,

900 meters or less; Pete"n; Alta Verapaz; Zacapa; Santa Rosa;

Escuintla; Solola. Southern Mexico; British Honduras; Honduras;

West Indies; northwestern South America.

An erect annual, usually 75 cm. high or less, simple or sparsely branched, the

young stems puberulent or pilose; leaves thin, on slender petioles 2-7 cm. long,

ovate or broadly ovate, 3-10 cm. long, acute or abruptly short-acuminate, obtuse

or rounded at the base and often shallowly cordate, finely and closely serrulate,

3-5-nerved at the base, thinly pilose or hirsute when young but in age glabrate,

rough-puncticulate; flowers monoecious, the staminate spikes axillary, short, about

1 cm. long, pedunculate; pistillate spikes terminal and in the uppermost leaf

axils, the terminal ones 3-6 cm. long, dense or lax and interrupted, in fruit 5 mm.

broad; fruiting bracts 5-6 mm. long, 7-13-parted almost to the base, 1-flowered,

scaberulous, the lobes setaceous-filiform, eglandular; ovary hirtellous, the styles

4-6-lacinulate; capsule 2 mm. broad, pilose; seeds 1 mm. long, smooth.

Called "gusanillo" and "tarco" in Salvador.

Acalypha Skutchii I. M. Johnston, Journ. Arnold Arb. 19:

120. 1938. Oreja de venado.

Moist or wet, mixed forest, mostly in quebradas, 1,200-2,000

meters; Quezaltenango (type from Volcan de Zunil, A. F. Skutch

981); San Marcos. Oaxaca; Chiapas.

A simple or branched shrub or small tree, 1.5-6 meters high, the branches

stout, strigillose or glabrate; leaves mostly on very long, slender petioles, these

often 15 cm. long; leaf blades oblong to lanceolate or broadly ovate, 10-20 cm.

long, 3-12 cm. wide, abruptly acuminate or long-acuminate, narrowly obtuse to

broadly rounded at the base, crenate-serrate, strigillose when young but in age

almost wholly glabrous, slightly paler beneath, penninerved or rather conspicuously

3-nerved at the base, the lateral nerves 7-10 pairs; stipules 10-18 mm. long;

spikes unisexual, the pistillate ones terminal, with a very stout rachis, 10-20 cm.

long, short-pedunculate, the bracts rather distant, strigillose, in fruit 3-5 mm.

44 FIELDIANA: BOTANY, VOLUME 24

long, deeply 7-11-lobate; styles purple-red, lacinulate; ovary strigose; capsule

5-6 mm. broad.

This shrub is a common one in many of the damp quebradas in

the mountains of the Occidente. The leaves exhibit a good deal of

variation in outline and venation, so much so that two species may

be represented, but the inflorescences seem to be uniform in the

several forms. Closely related to this species is A. laxiflora Muell.

Arg., which is reported from Veracruz and Cuba. While very

similar in foliage and other characters to A. Skutchii, it seems essen-

tially different (according to a photograph of the type) in its slender

flexuous staminate spikes.

Acalypha subviscida Wats. Proc. Amer. Acad. 21: 440. 1886.

Mostly in pine-oak forest, 1,900-2,050 meters; Chimaltenango;

Huehuetenango. Mexico.

An erect or suberect perennial, herbaceous throughout or sometimes suffrutes-

cent below, simple or branched, usually much less than a meter high but sometimes

as much as 1.5 meters, the stems densely pubescent and stipitate-glandular;

leaves on slender petioles 1-6 cm. long, broadly ovate to lance-ovate, thin, 3-9 cm.

long, acuminate, rounded and usually somewhat cordate at the base, crenate,

palmate-nerved at the base, thinly or densely and softly pubescent, usually stipi-

tate-glandular beneath on the nerves; flowers monoecious, the terminal spikes

androgynous or wholly pistillate, 15 cm. long or less, dense, sessile; axillary spikes

staminate or pistillate or androgynous, solitary or 2-3-nate, the staminate some-

times 9 cm. long, slender, dense, the pistillate ones 5-7 cm. long; pistillate bracts

rather lax, in fruit 6-8 mm. broad, reniform, 8-15-crenate, densely pubescent and

stipitate-glandular, 2-4-flowered; styles subpalmately 3-7-lacinulate; capsule 2

mm. broad, pubescent, stipitate-glandular, muricate; seeds 1 mm. long, blackish

gray, almost smooth.

Acalypha tenuicauda Pax & Hoffm. Pflanzenreich IV. 147,

xvi: 149. 1924.

Moist or wet thickets or mixed forest, often in second growth,

700-2,000 meters; Escuintla (type from Los Diamantes, Barranco

del Cucunya, Seler 2508); Suchitepe"quez; Solola; Quezaltenango;

San Marcos. Chiapas.

A slender shrub 1-4.5 meters high, the branchlets fulvescent-pilose or tomen-

tulose at first, soon glabrate; leaves on slender petioles 6-16 cm. long, thin, ovate

to rounded-ovate or elliptic-ovate, mostly 10-20 cm. long and 6-15 cm. wide,

long-acuminate or caudate-acuminate, obtuse to rounded and subcordate at the

base, closely and finely serrate, palmate-nerved at the base, thinly pilose or hirsute

or almost glabrous; stipules 8-12 mm. long, triangular-lanceolate, subulate-

acuminate, sparsely stipitate-glandular on the margins; flowers monoecious, the

spikes axillary, the staminate 5-6 cm. long, short-pedunculate, dense, many-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 45

flowered, 2-3 mm. thick; pistillate spikes slender-pedunculate, often 15 cm. long,

dense or usually lax, sparsely pilose; bracts in fruit about 3 mm. long and 6 mm.

broad or somewhat larger, broadly reniform, subemarginate at the apex, shallowly

about 11-dentate, sparsely stipitate-glandular, the teeth triangular-lanceolate,

acuminate; styles pinnately 10-15-lacinulate; capsule 2 mm. broad, verrucose,

sparsely pilose.

A. tacanensis Lundell (Contr. Univ. Mich. Herb. 4: 11. 1940),

described from Volcan de Tacana, Chiapas, is probably a synonym

of this species, although we have seen no material of it. It is described

as having larger pistillate bracts, as much as 6.5 mm. long.

Acalypha trachyloba Muell. Arg. Flora 55: 25. 1872.

Moist or wet, mixed or oak, usually dense forest, or in moist or

wet thickets, 1,800-3,100 meters; Guatemala; Sacatepe"quez; Chimal-

tenango; Solola; Huehuetenango; Quezaltenango; San Marcos.

Southern Mexico.

A slender shrub, commonly 1-3 meters high, erect or often more elongate and

subscandent, the young branches densely fulvescent-pilose; leaves thin, on slender

petioles 1-5 cm. long, broadly ovate or elliptic-ovate, mostly 5-14 cm. long, usually

caudate-acuminate, rounded and often shallowly cordate at the base, coarsely

crenate-dentate, villous-pubescent on both surfaces, often very densely so, espe-

cially beneath, or in age sometimes glabrate, 5-nerved at the base; stipules 5 mm.

long, linear, reflexed; flowers monoecious, the spikes axillary, the staminate slender,

7-11 cm. long, sessile or subsessile, dense, tomentulose; pistillate spikes very short

in anthesis, borne on long slender peduncles, in fruit 1.5-3 cm. long, very dense,

many-flowered, often as broad as long and head-like; fruiting bracts 1-1.5 cm.

long, cleft almost to the base into 7-9 stiff, linear or subulate segments, these

densely stipitate-glandular; ovary hirtellous and muricate; styles pilose, pecti-

nately dissected; capsule 3 mm. in diameter, tuberculate, hispid; seeds smooth,

almost 2 mm. long.

A common shrub in the central and western highlands.

Acalypha triloba Muell. Arg. Linnaea 34: 23. 1865. Hierba del

cancer.

At 2,500-2,900 meters; Huehuetenango (near San Juan A titan,

Sierra de los Cuchumatanes, Steyermark 51959). Mexico(?).

An erect perennial herb almost a meter high, simple or probably sometimes

branched, densely hispid almost throughout with long spreading stiff hairs; leaves

thin, on long slender petioles, ovate, 5-10 cm. long, acuminate, rounded at the

base and often shallowly cordate, crenate-dentate, 3-nerved at the base; flowers

monoecious, the spikes axillary and terminal, the staminate ones axillary, about

5.5 cm. long and borne on a peduncle of the same length, slender, interrupted

below; pistillate spikes terminal and in the upper leaf axils, subsessile, 5-9 cm. long

or shorter, very dense; fruiting bracts 2-3-flowered, 3-lobate to the middle or

more deeply, long-ciliate, eglandular, the terminal lobe longer than the others,

46 FIELDIANA: BOTANY, VOLUME 24

triangular-lanceolate, acute, the lateral lobes subquadrate, truncate; ovary hirsute,

the styles slender, 5-8-lacinulate, purple-red, showy, the central portion long-

pilose; seeds foveolate-puncticulate.

This species was described as coming from Mexico, but the

labels of the type and other specimens do not indicate any definite

locality. The Guatemalan collection cited was obtained in a region

through which either Sesse" or Mocino or possibly both are known

to have passed, and it is quite possible that the type was collected

in Guatemala rather than Mexico. In general appearance this

species is remarkably like some species of Urtica.

Acalypha unibracteata Muell. Arg. Linnaea 34: 160. 1865.

Tornillo (Pete"n).

Moist or wet, mixed forest or in moist or dry thickets, 200-1,650

meters; Pete*n; Alta Verapaz; Baja Verapaz; Santa Rosa; Guate-

mala; Sacatepe"quez. Southern Mexico; Salvador.

A slender shrub 1-2 meters high, erect, the young branches fulvous-villosulous,

in age brown or reddish brown; leaves thin or in age thick and firm, short-petiolate

or the petioles sometimes slender and elongate, lance-ovate to lance-oblong, mostly

2-5 cm. long and 1-3 cm. wide, acuminate or narrowly long-acuminate, obtuse

to cordate at the base, crenate-serrate, pilose when young but in age often glabrate,

palmate-nerved; stipules small, setaceous-subulate; spikes axillary, unisexual, the

staminate 1-1.5 cm. long, pedunculate, grayish-puberulent; pistillate spikes usually

on very long, almost filiform peduncles, generally reduced to a single bract;

pistillate bracts at anthesis 2 mm. wide, accrescent in age, 1-flowered, reniform-

ovate, 9-11-laciniate to about the middle, the segments lanceolate; ovary slightly

muricate, pubescent, the styles pectinately about 9-lacinulate.

This and A. leptopoda are very closely related, and not always

satisfactorily separable. Called "pie de paloma" in Salvador;

"chilibtux" (Yucatan, Maya). In Yucatan the slender stout

branches are utilized for making baskets and bird cages.

Acalypha villosa Jacq. Sel. Stirp. Amer. 254. pi. 183, f. 61.

1763. A. flagellata Millsp. Field Mus. Bot. 2: 417. 1916 (type from

Yucatan).

Wet to rather dry thickets, frequently in second growth, often

on limestone, 1,000 meters or less; Pete*n; Alta Verapaz; Santa

Rosa; Guatemala; Quiche". Southern Mexico; British Honduras;

Honduras; Salvador; Costa Rica; Panama; tropical South America.

A shrub 1.5-4 meters high, sparsely branched, the young branches stout,

sparsely or densely pubescent; leaves thin, on long slender petioles, ovate to

broadly triangular-ovate, mostly 10-20 cm. long and 5-12 cm. wide, usually long-

acuminate, generally rounded at the base, often subcordate, crenate, palmate-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 47

nerved, sparsely or densely pilose or almost wholly glabrous; stipules subulate;

flowers monoecious, but the branches often unisexual; staminate spikes 3-13 cm.

long, dense, short-pedunculate, sometimes with a few pistillate flowers at the base,

pubescent; pistillate spikes 3-11 cm. long or longer, very lax, usually simple, some-

times branched, the rachis filiform, pubescent; pistillate bracts minute, with 2 or

more flowers, these pedicellate; ovary strongly muricate, otherwise glabrous;

styles short, 5-10-lacinulate; capsule 2.5 mm. in diameter, muricate; seeds sub-

globose, smooth, scarcely 1 mm. long.

Called "tapa-camino" in Veracruz.

Acalypha Wilkesiana Muell. Arg. in DC. Prodr. 15, pt. 2: 817.

1866. Capa del rey; Pastor (Pet&i).

Perhaps native of the southern Pacific region, now grown for

ornament in most tropical and subtropical regions; planted very

commonly in Guatemala, at low and middle elevations, most abundant

in the lowlands.

Usually a shrub of 1-3 meters, the young branchlets tomentulose or pilose,

soon glabrate; leaves rather firm, on long slender petioles, broadly ovate, 10-20

cm. long, short-acuminate, broadly cuneate or very obtuse at the base, crenate-

serrate, 5-nerved at the base, often lustrous, deep green with usually pink or pale

red margins, often spotted with pink or dull red or purple; flowers monoecious,

the spikes axillary; pistillate bracts 1-flowered, sparsely puberulent, 9-13-dentate;

ovary puberulent; styles pectinately 11-15-lacinulate.

Called "manto de Jesus" in Salvador. This is one of the most

plentiful ornamental shrubs everywhere in the lowlands, thriving

with little or no attention, and sometimes persisting around the

sites of former dwellings. It is common in most of the cemeteries

of the tierra caliente. It is used abundantly for hedges, especially

those in the Pacific bocacosta enclosing coffee plantations.

ADELIA L.

Reference: F. Pax, Pflanzenreich IV. 147, vii: 64-71. 1914.

Shrubs or small trees, pubescent or usually soon glabrate, the branchlets

often spinescent; leaves alternate, sometimes crowded on the branchlets, mem-

branaceous or chartaceous, entire, short-petiolate, penninerved, usually barbate

beneath in the axils of the nerves, pellucid-pun cticulate; flowers dioecious, apetal-

ous, small, axillary, the staminate short-pedicellate, the pistillate long-pedicellate;

staminate calyx closed in bud, ovoid, in anthesis valvately 4-5-parted; stamens

8-17, free in bud, in age connate into a short or elongate column, the anthers versa-

tile, dorsifixed near the base, the cells parallel, longitudinally dehiscent; extra-

staminal disk usually annular, rarely of 5 glands; pistillate sepals 5-7, narrow,

reflexed in anthesis, the disk annular, pubescent, adnate to the calyx; ovary

generally 3-celled, the styles free or nearly so, laciniate; ovules 1 in each cell;

capsule 3-lobate, pubescent, separating into 2-valvate cocci that separate from

48 FIELDIANA: BOTANY, VOLUME 24

a persistent central column, the endocarp crustaceous; seeds subglobose, smooth,

gray, not carunculate; endosperm carnose, the cotyledons broad, flat.

About 10 species, all American and chiefly in tropical America.

Two others are known from southern Central America.

Adelia barbinervis Schlecht. & Cham. Linnaea 6: 362. 1831.

Moist or rather dry thickets or thin forest, sometimes in second

growth thickets, 350 meters or less; Pete"n; Alta Verapaz; Izabal;

Retalhuleu; Huehuetenango. Mexico; British Honduras; Salvador.

A shrub or tree, sometimes 9 meters high, with a trunk as much as 20 cm. in

diameter, the trunk round, short, the branches spreading, stiff and stout, the small

ones often spinose, villosulous-pubescent or in age glabrous or nearly so; leaves

on petioles 2-6 mm. long, obovate or obovate-lanceolate, mostly 4-9 cm. long and

2-3.5 cm. wide, cuspidate-acuminate to obtusely acute, narrowed below to a

narrow subobtuse base or more often attenuate, chartaceous, deep green and

glabrous above, penninerved, paler beneath, pubescent along the costa or at least

barbate in the axils of the nerves, the lateral nerves 5-7 pairs; staminate flowers

greenish or whitish, few or numerous in each axillary fascicle, the pedicels 3-7 mm.

long, the pistillate pedicels 12 mm. long or in fruit as much as 2 cm. long, pubes-

cent; staminate sepals 5, ovate-lanceolate, almost 2 mm. long, pubescent; stamens

8-11, the filaments pilose at the base; pistillate sepals 6-7, linear-lanceolate, acute,

2-3 mm. long; ovary densely hirsute; capsule pubescent, shallowly 3-lobate, the

cocci somewhat carinate dorsally, 7 mm. long, 11 mm. broad; seeds globose,

lustrous, 4 mm. in diameter.

Known in Salvador by the names "tintorillo," "macaguite(?),"

and "espino bianco"; "chau" (Yucatan, Maya). In the case of the

Salvadorean name "macagiiite" and the statement that the pul-

verized seeds are applied to the hair to make it soft and sleek, there

is probably a confusion with Trichilia, which bears this name in

Guatemala and is used for this purpose. The wood of A. barbinervis

is said to be whitish throughout, slightly fragrant when fresh;

probably no use is made of it unless as firewood.

ALCHORNEA Swartz

Reference: F. Pax, Pflanzenreich IV. 147, vii: 220-253. 1914.

Shrubs or trees, the indument of simple or stellate hairs; leaves alternate,

2-stipulate, on long or short petioles, membranaceous or coriaceous, more or

less dentate or subentire, usually with 2 glandular spots beneath at the base,

penninerved or palmate-nerved; flowers apetalous, monoecious or dioecious, usually

in unisexual spikes, these simple or paniculately branched, the staminate spikes

commonly axillary, the pistillate terminal; staminate bracts several-flowered, the

pistillate 1-3-flowered; staminate calyx globose in bud and closed; in anthesis

valvately 2-5-parted; stamens 8 or rarely fewer, the filaments connate at the very

base; anthers oblong, dorsifixed, the cells parallel, longitudinally dehiscent;

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 49

pistillate sepals 3-6, usually 4; ovary 2-3-celled, the styles free or short-connate,

usually simple; ovules solitary in each cell; capsule 2-3-coccous or subglobose,

the cocci 2-valvate, separating from the persistent central axis; seeds not caruncu-

late, the endosperm carnose, the cotyledons broad, flat.

Species about 45, widely dispersed in the tropics of both hemi-

spheres. Two or three others are known in southern Central America.

Leaves mostly 8-11 cm. long, entire or nearly so, those of sterile branches often

crenate-serrate A. integrifolia.

Leaves mostly 12-20 cm. long, all or most of them conspicuously crenate-dentate.

A. latifolia.

Alchornea integrifolia Pax & Hoffm. Pflanzenreich IV. 147,

vii: 237. 1914.

Wooded swamps, 1,400-1,500 meters; endemic; Alta Verapaz

(type from Coban, Tuerckheim 103).

A shrub or tree, sometimes 7 meters high, the branches with pale ochraceous

bark, the young ones glabrous; leaves coriaceous, on petioles 1.5-4.5 cm. long,

oblong or ovate-oblong, mostly 7-12 cm. long and 3-6 cm. wide, somewhat

narrowed to the obtuse or rounded apex, obtuse or rounded at the base, entire

or those of sterile branches sometimes crenate, glabrous above, almost glabrous

beneath but with a few minute stellate hairs, with 2-4 glandular spots at the base,

or these sometimes very obscure, 3-nerved at the base, the costa emitting 4-6

lateral nerves above the base; pistillate spikes 4-6 cm. long, solitary in the leaf

axils, simple, the rachis minutely stellate-pubescent; bracts 1 mm. long, triangular,

acute, 1-2-flowered, the flowers sessile, 2-bracteolate; sepals 4, ovate, acute, 2 mm.

long, sparsely pilose, ciliate; ovary 2-3-celled, densely stellate-pilose; styles 2-3,

short-connate, 5-9 mm. long.

We have found this tree only in the large swamp east of Tactic,

probably the type locality, where it is common, growing among or

near the great sphagnum mounds. All the trees found were sterile.

Alchornea latifolia Swartz, Prodr. Veg. Ind. Occ. 98. 1788.

Carreton; Cajeton; Tern (Alta Verapaz).

Moist or wet, mixed forest, sometimes on limestone, often abun-

dant along steep slopes of barrancos, 1,400 meters or less; Peten;

Alta Verapaz; Izabal; Jutiapa; Escuintla; Guatemala; Solola;

Suchitepe"quez; Huehuetenango; Quezaltenango. Southern Mexico;

British Honduras to Salvador and Costa Rica; West Indies.

A tree, sometimes 20 meters high with a trunk 45 cm. or more in diameter,

the crown rounded or irregular, the bark deep gray or light brown, the inner bark

dark reddish, the branchlets minutely stellate-puberulent or almost glabrous;

leaves subcoriaceous, on stout petioles 4-10 cm. long, ovate or elliptic, some-

times very broadly ovate, mostly 12-25 cm. long and 6-18 cm. wide, abruptly

short-acuminate to obtuse, generally obtuse or rounded at the base, crenate-

50 FIELDIANA: BOTANY, VOLUME 24

dentate, 3-nerved from the base, the costa emitting several lateral nerves above

the base, glabrous above, 2-4-glandular beneath at the base, when young sparsely

and very minutely stellate-puberulent beneath, often densely pubescent in the

axils of the nerves; staminate spikes paniculate, slender, 7-20 cm. long, laxly

many-flowered, the pistillate spikes simple or branched, 10-20 cm. long, solitary

or geminate, the rachises minutely stellate-puberulent, the bracts triangular, acute,

scarcely 1 mm. long, the staminate bracts 5-8-flowered, the pistillate 1-3-flowered;

staminate flowers subsessile, 2-bracteolate, 4 mm. broad, the 2 sepals concave,

ovate, acute, glabrous; stamens 8, connate at the base; pistillate sepals 4, ovate,

acute, 1 mm. long; ovary usually 2-celled, sometimes 3-celled, puberulent; styles

6-20 mm. long; capsule 7 mm. high, 10 mm. broad, didymous, dark red or brown-

red, glabrate; seeds echinate, 5 mm. in diameter.

Called "canelito" in Honduras; "pochote," "pochoton," "tambor,"

"tepeachote" (Salvador); "carne de caballo" (Veracruz); "fiddle-

wood" (British Honduras). The wood in this genus is pale brown,

light, soft, and perishable. Little or no use is made of it in Guate-

mala. Large trees often are left standing in the coffee plantations

of the lower Pacific slopes. A probable synonym of this species is

A. similis Muell. Arg., described from Oaxaca. We have seen type

material, which seems to differ in no respect from the material we

have referred here.

Aleurites Fordii Hemsley, the Chinese wood-oil or tung-oil tree,

and A. moluccana (L.) Willd., the candlenut or varnish tree, are

sometimes planted in Guatemala, especially around the capital, as

curiosities or for experimental purposes. The former has ovate-

cordate, sometimes 3-lobate leaves, pubescence of simple hairs, and

rather showy, pink or white flowers; the latter has large ovate-

acuminate leaves, pubescence of stellate hairs, and white flowers.

A. Fordii has been planted on a rather large scale along the Gulf

coast of the United States for its abundant seeds, from which is

obtained tung oil of commerce, used in preparation of paints and

other manufactures.

AMANOA Aublet

Trees or shrubs, glabrous; leaves alternate, penninerved, coriaceous, entire,

short-petiolate; stipules intrapetiolar, sometimes connate; flowers monoecious, petal-

iferous, glomerate-fasciculate in the axils of leaves or bracts, minutely bracteate

or the bracts sometimes rather large and foliaceous, sometimes arranged in strobili-

form dichasia, the flowering branches sometimes simulating racemes or panicles;

staminate sepals 5, subequal, firm, imbricate; petals 5, short, scale-like, unguiculate;

disk extrastaminal, deeply lobate, sometimes small; stamens 5, inserted on a thick

receptacle, episepalous, the filaments free, generally short, the anthers ovoid,

introrse; ovary rudiment columnar, 3-lobate at the apex; pistillate sepals usually

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 51

narrower than the staminate ones; ovary subglobose, 3-celled, the 3 stigmas

sessile, carnose, disk-like; ovules geminate in each cell; capsule drupe-like, indurate

in age, often muriculate, separating into 2-valvate cocci, 3-seeded or by abortion

2-1-seeded; seeds smooth, emarginate at the base, not carunculate, the testa

crustaceous; endosperm thin or none; cotyledons carnose, plane on the inner side,

subtrigonous, the radicle short.

Ten species, 3 of them African, the others in tropical America.

Only one is known in continental North America.

Amanoa potamophila Croizat, Amer. Midi. Nat. 29: 475. 1943.

Moist or wet, mixed forest, at or little above sea level, often along

stream banks; Izabal. British Honduras, the type from seacoast,

Cattle Landing, British Honduras, W. A. Schipp 1204.

A glabrous shrub or tree, sometimes 12 meters high with a trunk 25 cm. or

more in diameter, the smaller branches light brownish or grayish, with large leaf

scars; leaves coriaceous, on petioles 1 cm. long or usually shorter, elliptic-oblong

to obovate-oblong or elliptic, 6-19 cm. long, 3-8 cm. wide, narrowed to an obtuse

apex or abruptly obtuse-acuminate, rounded to broadly cuneate at the base, entire,

with about 10 pairs of lateral nerves; inflorescences spike-like, as much as 9 cm.

long, often leafy, the flowers sessile, subtended by rather large, coriaceous, ovate

bracts; pistillate buds broadly ovoid, 4-5 mm. long; pistillate sepals coriaceous,

oblong-ovate, 6 mm. long, obtuse; fruiting pedicels thick and ligneous, about 1 cm.

long; columella stout, angulate, 1.5 cm. long, the valves of the capsule ligneous,

2 cm. long, 2-3 mm. thick; seeds brownish-marmorate, smooth, lustrous, broadly

somewhat obcordate, about 14 mm. long and 11 mm. broad, the large hilum sub-

central.

Called "swamp icaco" in British Honduras. This has been re-

ported from the region as A. grandiflora Muell. Arg., a closely related

species of northern South America, which seems to differ constantly

in its considerably larger seeds. Croizat states that A. potamophila

"amply differs from this (A. grandiflora) and other species of the

genus in the range and several characters, such as the length of the

pedicel, the thickness of the epicarp and the size of the seed."

"Different range" is scarcely a specific character, in spite of the

fact that it is often invoked to bolster species of weak characters;

of the other characters the only one that holds is the size and form

of the seed, but this does seem to be a constant and probably valid

difference. The wood in this genus is reddish to purplish brown,

moderately or very dense, and difficult to work. In the Amazon

region it is sometimes used for heavy and durable construction.

ASTROCASIA Robinson & Millspaugh

Slender shrubs or small trees, glabrous or nearly so; leaves on long slender

petioles, membranaceous, alternate, entire; flowers small, dioecious, petaliferous,

52 FIELDIANA: BOTANY, VOLUME 24

fasciculate in the leaf axils, on long filiform pedicels; staminate sepals 5, imbricate,

widely spreading in anthesis, the petals erect or ascending; disk cupular, 5-crenate;

stamens 10, the filaments connate into a slender column, this expanded at the

apex into a disk, the anthers ellipsoid, sessile, horizontally dehiscent; ovary rudi-

ment none; fruit capsular, 3-coccous, shallowly 3-sulcate, elastically dehiscent;

seeds pale dull brownish, irregular trigonous-globose, smooth, not strophiolate.

Two other species have been described from Mexico.

Astrocasia phyllanthoides Rob. & Millsp. Bot. Jahrb. 36,

Beibl. 80: 20. 1905. Chinchin (Pet<§n).

Moist or dry thickets, often on limestone, 200 meters or less;

Pete"n; Retalhuleu. Yucatan Peninsula of Mexico; British Honduras.

A glabrous shrub or small tree, sometimes 7 meters high but usually 2-3

meters high, the trunk as much as 15 cm. in diameter, the branches very slender

but stiff, grayish; petioles very slender and unequal, 2-6 cm. long, glandular at

the apex; leaf blades mostly broadly ovate or rounded-ovate, 12 cm. long and 8

cm. wide or usually much smaller, obtuse at the apex, obtuse to almost rounded

at the base, often broadly cuneate, green above, very pale and whitish beneath,

penninerved or somewhat 3-nerved or 3-plinerved at the base; staminate pedicels

filiform, mostly 1-1.5 cm. long, the calyx 4 mm. in diameter; sepals herbaceous,

orbicular, the petals oblong; pistillate pedicels often 4 cm. long, slender but rather

stiff; capsule brownish, about 8 mm. long, minutely papillose; seeds smooth, slightly

lustrous, 5 mm. long.

The Maya names in Yucatan are recorded as "cayuc" and "pix-

toncax." It is probable that the genus Astrocasia should be combined

with Phyllanthtis. In general appearance the species of Astrocasia

are similar to certain species of Phyllanthus, and it is questionable

whether the technical flower characters supposed to separate the

two genera actually exist.

BERNARDIA Adanson

Reference: F. Pax, Pflanzenreich IV. 147, vii: 21-45. 1914.

Shrubs or small trees, the pubescence of simple or fasciculate hairs; leaves

alternate, petiolate or subsessile, dentate, penninerved or sometimes 3-nerved at

the base, usually rather thick, with 2 glandular spots at the base of the blade;

stipules small; flowers monoecious or dioecious, apetalous, the staminate in axillary,

short or elongate spikes, small, usually several in each bract; pistillate flowers

mostly aggregate in a terminal few-flowered inflorescence, or sometimes in the axils

of the uppermost leaves, sometimes in racemiform spikes, the bracts coriaceous,

concave; staminate calyx closed in bud, globose, in anthesis 3-4-parted; stamens

4-22, the filaments short, free, the anthers erect, the cells distinct, subglobose;

pistillate sepals 4-6, subtended by sepal-like bracts; disk annular or of separate

glands; ovary 3-celled, the styles short, distinct at the base, 2-parted, the lobes

smooth or lacerate; ovules solitary in each cell; capsule tridymous, the 2-valvate

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 53

cocci separating from the persistent columella, the endocarp crustaceous; seeds

prismatic-trigonous, not strophiolate, more or less carinate dorsally, the testa

crustaceous; endosperm carnose, the cotyledons broad, flat.

About 35 species, all American and chiefly in tropical regions.

Style branches not laciniate; staminate spikes mostly equaling or little shorter than

the leaves. Leaves glabrous beneath or nearly so B. interrupta.

Style branches laciniate; staminate spikes short, much shorter than the leaves and

often shorter than the petioles.

Leaves glabrous beneath or nearly so, mostly oblong-lanceolate or oblanceolate-

oblong but sometimes broader B. oblanceolata.

Leaves densely pubescent beneath, at least when young, the pubescence some-

times sparse in age but always conspicuous.

Leaves very scabrous on the upper surface, minutely stellate-pubescent be-

neath B. yucatanensis.

Leaves not or very slightly scabrous on the upper surface, densely and softly

stellate-pubescent beneath with coarse hairs B. mollis.

Bernardia interrupta (Schlecht.) Muell. Arg. Linnaea 34: 171.

1865. Acalypha interrupta Schlecht. Linnaea 7: 386. 1832.

Thin forest or thickets, on limestone, little above sea level;

Pete"n. Southern Mexico; British Honduras.

A shrub or tree, sometimes 10 meters high with a trunk as much as 20 cm. in

diameter, the branches minutely stellate-puberulent, the bark gray and smooth,

the inner bark orange; leaves membranaceous, on unequal petioles 1.5-10 cm. long,

ovate or obovate to lanceolate, mostly 7-20 cm. long and 4-10 cm. wide, acuminate,

acute to almost rounded at the base, crenate-dentate, penninerved but definitely

3-nerved at the base, green and almost glabrous above, paler beneath, very minutely

and sparsely stellate-puberulent or almost glabrous, with 4-6 plate-like glands

near the base; flowers dioecious, the staminate spikes axillary, 4-18 cm. long,

floriferous almost to the base, the bracts 2 mm. long, ovate, 5-7-flowered, the

pedicels 3-4 mm. long; pistillate spikes terminal, many-flowered, 6-7 cm. long,

the bracts ovate, 4 mm. long, 1-flowered, the flowers sessile; staminate sepals 3,

oblong, 1.5 mm. long, oblong, apiculate, the stamens usually 12; pistillate sepals

4-5, ovate, long-acuminate, tomentose, the outer ones 3-4 mm. long; ovary densely

fulvous-sericeous, the 3 style branches not laciniate; capsule about 8 mm. long and

12 mm. broad, deeply 3-coccous; seeds 8-9 mm. long, 6-7 mm. broad, slightly

verruculose.

Called "waika ribbon" in British Honduras.

Bernardia mollis Lundell, Contr. Univ. Mich. Herb. 4: 12.

1940.

Dry or moist thickets, 2,500 meters or less; Retalhuleu; Quezal-

tenango; San Marcos. Chiapas, the type from Volcan de Tacana,

E. Matuda 2966.

A shrub or tree 1.5-6 meters high, or reported as sometimes a tree of 12-15

meters with a trunk 25 cm. in diameter, the young branches stout, densely fulvous-

54 FIELDIANA: BOTANY, VOLUME 24

tomentose; leaves on petioles 4 cm. long or shorter, chartaceous, ovate or elliptic,

mostly 5-13 cm. long and 3-7 cm. wide, acuminate, rounded or obtuse at the base,

crenate-serrate, sparsely stellate-pilosulous above or almost glabrous in age, the

veins often more or less impressed, softly and usually densely stellate-pilose

beneath, penninerved but also 3-nerved at the base; flowers dioecious, the stami-

nate spikes axillary, 2.5-5 cm. long, stout, dense, the rachis stellate-tomentulose;

bracts broadly ovate, apiculate, 2 mm. long, 3-5-flowered, the pedicels 3 mm.

long or less; sepals 3, ovate or elliptic, 3.5 mm. long; stamens 22-24; capsule very

densely stellate-hirsute when young with spreading fulvous hairs, 12 mm. long or

more; style branches conspicuously laciniate.

Bernardia oblanceolata Lundell, Contr. Univ. Mich. Herb. 4:

13. 1940. B. mollis var. lanceifolia Lundell, loc. cit. (type from Volcan

de Tacana, Chiapas, E. Matuda 2978).

In forest or thickets, 1,500-2,500 meters; Huehuetenango (Cerro

Huitz, Steyermark 48656); doubtless also in San Marcos. Chiapas,

the type from Siltepec.

A shrub or tree 4-6 meters high, the trunk as much as 20 cm. in diameter,

the branches minutely stellate-puberulent or almost glabrous, in age fuscous;

leaves on petioles 2 cm. long or less, chartaceous, mostly oblanceolate to oblong-

obovate or oblong-lanceolate, 4-11 cm. long, 1.5-3.5 cm. wide, acuminate, obtuse

at the base, irregularly glandular-serrate, penninerved and also 3-nerved at the

base, deep green above and in age almost glabrous, much paler beneath, minutely

stellate-puberulent but soon glabrate, bearing 1-4 glands beneath at the base of

the blade; staminate inflorescences axillary, spike-like, 4.5 cm. long or shorter,

few-flowered, short-pedunculate, the bracts very broadly ovate, 3 mm. long, 3-6-

flowered, the pedicels 2.5 mm. long or less; staminate sepals 3, ovate-elliptic, 4 mm.

long, acute, appressed-pubescent; stamens about 15; pistillate inflorescences

terminal, short, only the basal flower fertile; sepals 5, broadly ovate, 2.5-3 mm.

long; ovary fulvous-tomentose, the styles laciniate; capsule fulvous-tomentose,

13-17 mm. long; seeds about 11 mm. long and 8 mm. broad.

Bernardia yucatanensis Lundell, Contr. Univ. Mich. Herb. 4:

14. 1940.

Moist or rather dry, open forest or in thickets, sometimes on

limestone, 1,500 meters or less; Pete"n (type from La Libertad,

Lundell 3355); Chiquimula. Campeche.

A shrub or tree, sometimes 12 meters high but usually lower, the young

branches densely stellate-tomentose with somewhat fulvous hairs, the older ones

fuscous or dirty brown; leaves on petioles 3 cm. long or less, chartaceous, lance-

oblong to ovate or sometimes rounded-ovate, mostly 4-15 cm. long and 2-6 cm.

wide or sometimes wider, somewhat narrowed to the acute or obtuse apex, obtuse

or rounded at the base, grayish when dried, chartaceous, irregularly crenate-

dentate, when young stellate-tomentose on both surfaces, in age scabrous above

and very rough to the touch, the veins and nerves often impressed, paler beneath,

in age densely and finely stellate-puberulent or sometimes glabrate, penninerved

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 55

and also 3-nerved at the base, bearing a few plate-like glands beneath near the

base of the blade; staminate inflorescence spike-like, axillary, 2 cm. long or less,

stout, densely stellate-tomentose, rather few-flowered; bracts broadly ovate, 2.5

mm. long, 1-3-flowered, the pedicels 1.5 mm. long; staminate sepals generally 3,

obovate-elliptic, 3 mm. long, glabrous within; stamens 23-25.

The plant is variable in foliage, if all the collections we have

referred here are conspecific. While extreme variations seem distinct

from B. mollis, some of the material is more or less intermediate

between them, and it is somewhat questionable whether B. yuca-

tanensis and B. mollis are really different species. Further, the three

species listed here, besides B. interrupta, are all closely related to one

another and toB. mexicana Muell. Arg., and it is doubtful whether,

when ample material has accumulated, all can be distinguished.

Breynia disticha Forster (Phyllanthus nivosus W. G. Smith),

sometimes called Snowbush in English, native of the Pacific islands,

is planted occasionally for ornament in Guatemala, at low and middle

elevations, but it is uncommon. It is a shrub of 1-2 meters, in general

appearance like a species of Phyllanthus. Its small broad leaves,

rounded at the apex, are handsomely variegated or spotted with

white.

CAPERONIA St. Hilaire

Reference: F. Pax, Pflanzenreich IV. 147, vi: 27-49. 1912.

Annual or sometimes perennial herbs, usually growing in wet places, often

in shallow water, hispid, often glandular, rarely glabrate; leaves alternate, short-

petiolate, 2-stipulate, narrow, serrate, generally penninerved, the lateral nerves

prominent beneath, slender, straight or nearly so; flowers small, greenish, monoe-

cious or rarely dioecious, petaliferous, racemose or spicate, the inflorescences pedun-

culate, the flowers solitary within the bracts, the staminate flowers in the upper

part of the inflorescence, the few pistillate ones in the lower part; disk none;

staminate sepals 5, valvate, the 5 petals affixed to the androphore within the

calyx, imbricate, usually unequal; stamens 10 and 2-seriate, the anthers ovoid,

longitudinally dehiscent; ovary rudiment cylindric, entire or 3-dentate at the

apex; pistillate sepals 5, often accrescent in fruit, imbricate, equal or unequal;

petals usually narrower than those of the staminate flower, sometimes much

reduced; ovary sessile, 3-celled, the style short, free or nearly so, palmate-laciniate;

ovules solitary in each cell; capsule tridymous, hispid or echinate, separating into

2-valvate cocci; seeds subglobose, not carunculate; endosperm carnose, the

cotyledons broad, flat.

About 35 species, in the tropics of America and Africa. One

other species is known from Panama.

56 FIELDIANA: BOTANY, VOLUME 24

Pubescence of the young stems of closely appressed hairs; stems conspicuously

fistulose-thickened C. castaneifolia.

Pubescence of the young stems wholly or partly of spreading, usually gland-

tipped hairs C. palustris.

Caperonia castaneifolia (L.) St. Hil. Hist. PI. Bre"sil 245.

1824. Croton castaneifolius L. Sp. PI. 1004. 1753.

In shallow water at the margins of lakes or ponds, 450-1,400

meters; Jalapa; Jutiapa. Southern Mexico; British Honduras;

Honduras; Nicaragua; West Indies; South America.

An erect annual, a meter high or less, simple or sparsely branched, the larger

stems fistulose and transverse-septate, as much as 12 mm. or probably even more

in diameter, the younger stems appressed-setulose, the older ones glabrous or

nearly so; leaves on petioles 0.5-4.5 cm. long, the lower ones elliptic or ovate,

5-16 cm. long, 3-8 cm. wide, obtuse at each end or sometimes acute at the apex,

crenate-dentate, the upper leaves lanceolate to linear-lanceolate, sharply serrate,

glabrate above, somewhat paler beneath, strigose on the nerves; stipules broadly

ovate, acuminate, 5 mm. long; racemes spike-like, with the peduncle 3-7.5 cm.

long, bearing below 1-4 pistillate flowers, interrupted; bracts ovate, acuminate,

1.5 mm. long, the pedicels 1-1.5 mm. long; staminate sepals lanceolate, acute,

green, 2 mm. long; petals white, spatulate-obovate, longer than the sepals; stamens

10; pistillate sepals ovate, in fruit 4-5 mm. long, setulose, the petals lance-obovate;

ovary densely covered with fusiform glands; capsule 7-8 mm. broad, 4 mm.

high, 3-sulcate, muricate; seeds foveolate, 2-3 mm. in diameter.

This is much less common in Central America than the following

species.

Caperonia palustris (L.) St. Hil. Hist. PI. Bre"sil 245. 1824.

Croton palustris L. Sp. PI. 1004. 1753. Caperonia pubescens Blake,

Contr. U. S. Nat. Herb. 24: 12. 1922 (type from Cristina, Izabal,

S. F. Blake 7574).

Moist or wet thickets or fields, or more often in ditches or marshes

or at the margins of lakes, 1,000 meters or less; Pete"n; Izabal;

Zacapa; Jutiapa; Escuintla; Retalhuleu; Huehuetenango. Southern

Mexico; Honduras and Salvador to Panama; West Indies; tropical

South America.

Plants annual, generally less than a meter high, erect or decumbent, the stems

rather stout but not or scarcely fistulose, simple or branched, sparsely or densely

glandular-setose with spreading broad-based hairs, often also hirsute with spread-

ing eglandular hairs; leaves on petioles 3-25 mm. long, the lower ones elliptic to

ovate or oblong, 7-12 cm. long, mostly obtuse, rounded or obtuse at the base, the

upper leaves on short petioles, lance-ovate to almost linear, obtuse to long-at-

tenuate at the apex, serrate, densely pilose to almost glabrous; stipules lanceolate,

acuminate, 3-5 mm. long; racemes spike-like, with the peduncle 4-10 cm. long,

interrupted, bearing near the base 1-5 pistillate flowers; bracts ovate, acuminate,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 57

1.5 mm. long, the flowers almost sessile; staminate sepals 5, lanceolate, acute,

1-1.5 mm. long; petals whitish, slightly longer than the sepals, spatulate-lanceolate,

obtuse; stamens 10; pistillate sepals 5, obovate, acuminate, unequal, in fruit 5 mm.

long, glandular-ciliate and glandular-setulose, the petals spatulate-lanceolate,

acute or obtuse, shorter than the sepals; ovary densely covered with fusiform

glands; capsule 6-7 mm. broad, 5 mm. high, muricate; seeds 2-3 mm. long, ovoid,

foveolate.

The author of C. pubescens remarks that it is "well distinguished

by its pubescence," a statement that is only partially correct. Un-

fortunately, there appears to be no distinguishing character except

the quantity of pubescence, which scarcely is a specific character.

The Guatemalan material of C. palustris is conspicuously variable

in pubescence, leaf shape, and length of petioles, but hardly more so

than material from the West Indies and South America.

CLEIDION Blume

Reference: F. Pax, Pflanzenreich IV. 147, vii: 288-298. 1914.

Trees or shrubs, usually almost glabrous, the indument, if any, of simple hairs;

leaves alternate, petiolate, generally dentate, penninerved, bearing 2 or more

glandular spots on the lower surface at or near the base; stipules caducous; flowers

monoecious or usually dioecious, apetalous; staminate flowers glomerate or fascicu-

late in interrupted spikes, these axillary, mostly elongate; staminate flowers small,

pedicellate, the pistillate in simple or branched racemes or solitary and long-

pedicellate in the leaf axils; disk none; staminate calyx globose or ovoid in bud,

in anthesis valvately 3-4-parted; stamens 35-80, densely crowded on a convex

or conic receptacle; anthers peltately attached dorsally, 4-celled, the connective

produced above the cells; pistillate sepals 3-4, imbricate; ovary usually 3-celled,

the styles elongate, filiform, commonly short-connate below, deeply 2-fid; ovules

solitary in each cell; capsule large or small, 2-3-dymous, separating into 2-valvate

cocci, or by abortion 1-coccous; seeds subglobose, not carunculate; endosperm

carnose, the cotyledons broad, flat.

About 20 species, in the tropics of both hemispheres, but mostly

in the Old World. One or two other species are known from southern

Central America.

Petioles 1 cm. long or less; leaf blades membranaceous, broadest above the middle.

C. nicaraguense.

Petioles mostly 2-3.5 cm. long; leaf blades chartaceous, broadest at or near the

middle C. oblongi folium.

Cleidion nicaraguense Hemsl. Biol. Centr. Amer. Bot. 3: 130.

1883.

Wet mixed forest, at or near sea level; Izabal (between Dartmouth

and Morales, Steyermark 39052). Nicaragua; Costa Rica.

58 FIELDIANA: BOTANY, VOLUME 24

A shrub or small tree, usually 5 meters high or less, the branches very slender,

hirtellous when young, pale in age, the bark light gray; petioles 5-11 mm. long,

hirtellous; leaf blades oblong-oblanceolate, mostly 10-18 cm. long and 4-6.5 cm.

wide, narrowly long-attenuate or caudate-acuminate, gradually narrowed to the

narrow base, the base itself obtuse, coarsely crenate-serrate, bright green above,

somewhat paler beneath, puberulent on the nerves and costa, the lateral nerves

about 8 pairs, ex current into the teeth; stipules narrow, acute, 3-5 mm. long;

staminate flowers fasciculate-racemose, the racemes 2.5 cm. long or less, few-

flowered, axillary, the flowers pilose, white or greenish white, 5-6 mm. in diameter,

short-pedicellate; sepals 3; stamens about 35, the filaments filiform, the connective

minutely penicillate at the apex.

This plant is known only from staminate material, and its proper

generic position is uncertain.

Cleidion oblongifolium (Standl.) Croizat, Journ. Arnold Arb.

24: 166. 1943. Alchornea oblongifolia Standl. Carnegie Inst. Wash.

Publ. 461: 66. 1935.

Moist or wet, mixed forest, about 1,350 meters; Quezaltenango

(Finca Pirineos, below Santa Maria de Jesus); Pete"n (type from

Camp 35, British Honduras boundary, 750 meters, W. A. Schipp

S-279). British Honduras, on limestone.

A tree 8-15 meters high, glabrous throughout, the trunk 15-45 cm. in diameter,

the bark rough, not fissured; leaves chartaceous, on petioles 2-3.5 cm. long, narrow-

ly oblong or lance-oblong, 16-24 cm. long, 4.5-7 cm. wide, short-acuminate

with an acute tip, slightly narrowed to the acute base, crenate-serrate almost to

the base, densely and minutely puncticulate, paler beneath, penninerved, the

lateral nerves about 11 pairs; staminate spikes laxly few-flowered, axillary, solitary

or binate, shorter than the petioles, the bracts very broadly ovate, the flowers

sessile or on very short, thick pedicels, globose in bud, glabrous; pistillate inflores-

cence axillary, about equaling the petioles, few-flowered, the pedicels scarcely

1.5 mm. long; sepals 4, linear-oblong, 1.5 mm. long; young capsule 3-coccous,

sparsely pilose with "minute appressed simple hairs, the styles bifid, with long

slender branches.

CNIDOSCOLUS Pohl

References: Rogers McVaugh, The Mexican species of Jatropha

(with special reference to possible sources of "chilte" rubber, pp. 1-23).

illus. Issued by the Rubber Development Corporation, July, 1943;

The genus Cnidoscolus: Generic limits and intrageneric groups, Bull.

Torrey Club 71: 457-474. 1944.

Herbs, shrubs, or trees, usually abundantly armed with long stiff stinging

hairs; leaves alternate, usually long-petiolate and palmately lobate, sometimes

pinnately lobate (not in Central American species); flowers usually small and

white, in dichotomous, generally long-pedunculate cymes, monoecious, apetalous;

sepals of the staminate flower petaloid, united usually for half their length; petals

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 59

none; stamens in 2 or more verticels, more or less monadelphous; ovary usually

3-celled, the styles connate at the base; ovules solitary in the cell; fruit capsular,

the seeds carunculate.

About 50 species, all in tropical America. Probably only the

following ones grow in Central America. By most authors the genus

has been united with Jatropha, but it has excellent claims to separate

status in both flower and foliage characters. The long stiff hairs

that cover almost all parts of the plants sting the flesh much worse

than most nettles, the pain often being excruciating and persistent

for hours, sometimes with the accompaniment of swelling and blister-

ing. They are probably the most painful of all the stinging plants

of Central America, and all animals naturally leave them strictly

alone. The technical characters used by Pax and some other authors

are difficult to understand, and probably were not understood by

the authors themselves, who had copied them from earlier writers.

Those used by McVaugh for separating the species are much better,

but at best the species are closely related and not always easy to

recognize. Some of the Mexican species have been found to contain

a kind of rubber, which is used locally, but in Central America, so

far as we know, no use is made of the plants.

Glands at the apex of the petiole several, slender, elongate, finger-like.

Stamens, at least the outer ones, with filaments distinct to the base; plants

herbaceous; leaves mostly lobate less than halfway to the base, the lobes

broad, not lobate, without gland-tipped hairs on the margins. . . .C. urens.

Stamens all monadelphous; plants usually more or less woody, often large shrubs

or small trees; leaves mostly lobate to below the middle, the lobes narrow,

usually lobate, with conspicuous gland-tipped hairs or setae along the

margins C. Souzae.

Glands at the apex of the petiole small, cushion-like or papilliform.

Pistillate perianth tubular below, falling off as a whole; seeds mostly 12-13

mm. long or larger. Plants densely armed with stinging hairs . . C. tubulosus.

Pistillate perianth divided nearly or quite to the base, the lobes falling off

separately; seeds (so far as they are known) 6-10 mm. long.

Base of the leaf blade (between the basal lobes) acute and decurrent, the lobes

very broad and somewhat overlapping; plants unarmed or nearly so and

almost glabrous C. Chayamansa.

Base of the leaf blade not at all narrowed or decurrent, truncate or broadly

cordate; plants usually armed with stinging hairs but sometimes unarmed.

Filaments of the outer (lower) stamens shorter than the anthers or equaling

them; seeds 6-8 mm. long; pistillate flowers 6-8 mm. long.

C. aconitifolius.

Filaments of the outer stamens about twice as long as the anthers; seeds

9-10 mm. long; pistillate flowers 8-12 mm. long C. multilobus.

Cnidoscolus aconitifolius (Mill.) I. M. Johnston, Contr. Gray

Herb. 68: 86. 1923. Jatropha aconitifolia Mill. Gard. Diet. ed. 8.

60 FIELDIANA: BOTANY, VOLUME 24

no. 6. 1768. J. Papaya Medik. Bot. Beob. 194. 1783. J. aconitifolia

var. Papaya Pax, Pflanzenreich IV. 147: 101. 1910. Chaya; Chayo;

Copapayo; Chichicaste.

Moist or dry thickets or open forest, often in open rocky places,

most often seen in hedges, where planted, 1,300 meters or less;

Pete"n; Alta Verapaz; Chiquimula; Jutiapa; Santa Rosa; Escuintla;

Suchitepe"quez; Retalhuleu; said to be planted in Quezaltenango, and

doubtless to be found in many of the other departments not listed

here. Southern Mexico; British Honduras to Salvador and Costa

Rica (chiefly or wholly in cultivation).

A shrub or small tree, generally 3-5 meters high, with a thick pale trunk;

petioles 10-20 cm. long or sometimes shorter, usually glabrous in Guatemalan

forms except at the apex, there setose-hispid; leaf blades very variable in form,

mostly 10-20 cm. long, shallowly or deeply 3-7-lobate, cordate at the base, rather

thick and fleshy when fresh, glabrous or nearly so and normally without stinging

bristles, the lobes acute to acuminate, often deeply lobate, but the leaves of culti-

vated forms often merely angulate-lobate or very irregularly and often shallowly

3-4-lobate; stipules lanceolate, deciduous; flowers white; staminate sepals glabrate

or minutely puberulent, white or greenish, usually less than 1 cm. long; stamens

10, monadelphous; pistillate sepals free, spatulate, 6-9 mm. long; ovary pubescent;

capsule bearing few or numerous bristles.

In Salvador sometimes called "chaidra," "chaira," "copapayo,"

and "papayillo"; "chay," "tziminchay," "tzah" (Yucatan). The

cultivated plants are almost free of stinging hairs, and it seems likely

that the almost glabrous form may have resulted from long years

of selection, since people wishing plants for cultivation for food

would naturally have preferred those without painful stinging

bristles. In Central America most of the plants are seen in cultiva-

tion in dooryards or in hedges about dwellings, but some of the clearly

wild plants seem to represent the same species. The leaves, when

young and tender, are cooked and eaten like spinach and other pot

herbs. Their use is rather widespread, but they can not be considered

a very common vegetable, and we have never seen the leaves in

market. Certainly the available supply of them is not very great

in Central America. This plant has been reported from British

Honduras and elsewhere as Jatropha tubulosa Muell. Arg., a species

that is not definitely known from British Honduras.

Cnidoscolus Chayamansa McVaugh, Bull. Torrey Club 71:

466. 1944. Chaya.

In hedges, about 250 meters; Retalhuleu (near Retalhuleu);

doubtless also in other parts of the country. Yucatan Peninsula

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 61

of Mexico; British Honduras (type from Honey Camp, C. L. Lundell

494); cultivated in southern Florida and in Cuba.

A stout shrub about 2 meters high, glabrous or nearly so, bearing few stinging

hairs or none; petioles usually much shorter than the leaf blades, the glands 2 or 1,

ovoid, convex; leaf blades truncate-cordate, broader than long, 3-lobate to below

the middle, the lobes flabellate, often 2-lobate, coarsely undulate-dentate, glabrous

except on the margins; inflorescences 2-5 cm. broad, the peduncles 10-25 cm. long,

the flowers numerous, white; pistillate flowers 9-10 mm. long, cleft almost to the

base, puberulent outside; staminate flowers 6-7 mm. long, tubular below; stamens

10.

The leaves of this species are eaten like those of C. aconitifolius.

McVaugh quotes a letter from Prof. Augusto PeYez Toro of Me"rida,

Yucatan, which would lead one to suppose that in Yucatan only

this species is used for food, but that is not true as regards Central

America, where C. Chayamansa is rare, but C. aconitifolius is rather

common and is used frequently as a green vegetable. The present

species has been in cultivation in southern Florida for 20 years,

and in Cuba still longer, having been introduced to that island from

Yucatan. Its leaves have been found to contain considerable

amounts of vitamin C. It is of some interest that C. Chayamansa

was obtained by Sesse" and Mocino, probably in the Yucatan region,

some 150 years ago.

Cnidoscolus multilobus (Pax) I. M. Johnston, Contr. Gray

Herb. 68: 86. 1923. Jatropha multiloba Pax, Pflanzenreich IV. 147:

107. 1910. Chichicaste de caballo.

Moist or dry thickets, 900 meters or lower; Alta Verapaz; El

Progreso; Jutiapa; Suchitepe"quez. Eastern and southern Mexico.

A shrub or small tree of 3-6 meters; leaves long-petiolate, glabrous at maturity

or nearly so, with abundant stinging hairs on the veins and petioles; leaf blades

mostly broader than long, as much as 30 cm. wide and 25 cm. long, rounded-

cordate at the base, generally 5-lobate to the middle or more deeply, the lobes

obovate, coarsely dentate; inflorescence long-pedunculate, densely furnished with

slender brownish stinging hairs; staminate flowers 12-15 mm. long, almost glabrous

to densely tomentose; pistillate perianth 8-12 mm. long, divided nearly or quite

to the base, usually densely pubescent outside; capsule densely furnished with

stinging hairs; seeds brown, 9-10 mm. long, the caruncle flat or somewhat curled,

2 mm. wide, more or less narrowed at the base and not cordate.

Cnidoscolus Souzae McVaugh, Bull. Torrey Club 71: 468. 1944.

Chaya cimarrona; Sac (Maya; both names used in British Honduras).

Moist or wet thickets, at or little above sea level; British Hon-

duras. Yucatan Peninsula of Mexico.

62 FIELDIANA: BOTANY, VOLUME 24

A stout coarse shrub or herb, commonly 2 meters high, the thick branches

densely covered with long stinging hairs, otherwise glabrous; leaves long-petiolate,

the petioles covered with long stinging hairs; leaf blades 10-15 cm. long, 12-20

cm. wide, lobate to the middle or more deeply, truncate or cordate at the base,

somewhat pilose on both surfaces, especially on the veins, with soft white sharp-

pointed hairs, the lobes oblong or ovate, serrate and often lobate; glands 2-4 at

the apex of the petiole, slender and finger-like, 1.5-3 mm. long, enlarged and

glandular at the apex; inflorescences long-pedunculate, armed with numerous

stinging hairs, the peduncles 20-30 cm. long, the cymes dense and many-flowered,

4-5 cm. broad; pistillate flowers 8-9 mm. long, divided almost to the base, the

perianth minutely puberulent on both surfaces; staminate flowers 8-10 mm.

long, densely puberulent outside; capsule oval to globose, broadly rounded at

each end, 8-9 mm. long, armed with stinging hairs; seeds about 7 mm. long, the

caruncle white or pale yellowish, 1.5-2.5 mm. wide, not at all or scarcely cordate.

This species may be recognized by the combination of glabrous

or nearly glabrous leaf blades with gland-tipped setae along their

margins, and the long slender glands at the apex of the petiole.

Cnidoscolus tubulosus (Muell. Arg.) I. M. Johnston, Contr.

Gray Herb. 68: 86. 1923. Jatropha tubulosa Muell. Arg. Linnaea 34:

212. 1865. J. cordifolia Pax, Pflanzenreich IV. 147: 107. 1910 (type

from Santa Rosa, Santa Rosa, Heyde & Lux 3474). C. cordifolius

I. M. Johnston, loc. cit. Chichicaste.

Moist or dry thickets, often on rocky slopes, 200-1,450 meters;

Zacapa; Chiquimula; Santa Rosa; Escuintla; Guatemala; Sacatepe"-

quez. Central and southern Mexico; Salvador; Colombia.

Usually a shrub of 1.5-4.5 meters, the stems and petioles abundantly armed

with long stinging bristles; leaves long-petiolate, commonly abundantly and softly

pubescent even at maturity, the lower surface with few or numerous stinging hairs;

leaf blades as wide as long or wider, often 20-30 cm. wide, openly rounded-cordate

at the base, mostly 5-lobate to the middle or lower, the lobes oblong to obovate,

usually cuspidate-acuminate, sinuate-dentate or subentire; inflorescences long-

pedunculate, dense and many-flowered, the peduncles densely beset with long

stinging bristles, the branches tomentose; staminate flowers 12-15 mm. long,

densely tomentose; pistillate calyx 10-15 mm. long, lobate to about the middle,

densely pubescent outside; capsule sparsely setose; seeds brown, 12-13 mm. long,

the caruncle flat or curled, 1-1.5 mm. wide, not cordate.

This plant is all too plentiful in some parts of Guatemala, Saca-

tepe"quez, and other departments, where it frequently forms dense

clumps or small thickets.

Cnidoscolus urens (L.) Arthur, Torreya 21: 11. 1921. Jatropha

wrens L. Sp. PL 1007. 1753. J. herbacea L. loc. cit. J. urens var.

herbacea Muell. Arg. in DC. Prodr. 15, pt. 2: 1101. 1866. Chichicaste;

Chichicaste de caballo; Chichicaste de burro.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 63

Brushy, moist or dry, often rocky slopes, sometimes in moist

or wet fields or thickets, occasionally in sandy places, 500 meters

or less; Zacapa; Chiquimula. Mexico; Honduras to Panama;

West Indies.

A shrub of 2 meters, or frequently herbaceous throughout and a meter high or

lower; petioles equaling or longer than the blades, usually densely short-pilose,

armed with numerous long spreading stinging bristles; leaf blades mostly 10-15

cm. long, usually cordate at the base, 3-5-lobate to the middle or more deeply,

usually abundantly pubescent on both surfaces and generally armed with numerous

long yellowish stinging bristles, the lobes mostly ovate or obovate or rounded,

acute or cuspidate-acuminate, sometimes obtuse, subentire or variously dentate

or shallowly lobate; stipules small, laciniate-dentate; cymes long-pedunculate,

few-many-flowered, pubescent and setose; staminate calyx white, 1 cm. long or

shorter, densely tomentulose and bearing few or numerous stinging hairs, lobate

to the middle; stamens 8-10; ovary puberulent and setose; capsule 10-12 mm.

long, usually bearing many long stinging white setae; seeds grayish, somewhat

compressed, 8 mm. long.

CODIAEUM Jussieu

Reference: F. Pax, Pflanzenreich IV. 147, iii: 23-30. 1911.

Shrubs or trees, glabrous or nearly so; leaves alternate, petiolate, coriaceous

or thick-membranaceous, penninerved, entire, undivided or rarely lobate; flowers

mostly monoecious, the staminate petaliferous, the pistillate apetalous, in elongate

racemes, the racemes solitary or binate in the upper leaf axils, unisexual or rarely

with 1-2 pistillate flowers in the staminate racemes, the flowers small and incon-

spicuous, the staminate fasciculate within the bracts, the pistillate solitary;

staminate sepals generally 5, imbricate, the petals small or minute; glands of the

disk 5-15, free; stamens 15-30 or more, inserted on a slightly elevated receptacle,

the filaments free, the anthers erect; ovary 3-celled, the styles distinct, recurved,

simple, the ovules solitary in each cell; capsule globose or tridymous, separating

into 2-valvate cocci; seeds carunculate, the testa lustrous, crustaceous; endosperm

carnose, the cotyledons broad, flat.

Six species, in the Malayan region and the Pacific islands. One

species is widely cultivated for ornament in tropical countries.

Codiaeum variegatum (L.) Blume, Bijdr. Fl. Ned. Ind. 606.

1825. Croton variegatus L. Sp. PI. ed. 3. 1424. 1763. Croton; Pon

(Coban).

Native of the Malayan region and the Pacific islands; grown

for ornament in most tropical regions, and often in hothouses in

the north; planted abundantly in the lowlands of Guatemala, and

less frequently at middle or even higher elevations.

A shrub or small tree; leaves lustrous, on petioles 1-4 cm. long, exceedingly

variable in shape and coloring, ovate-oblong or obovate-oblong to elliptic, spatu-

64 FIELDIANA: BOTANY, VOLUME 24

late, or linear, often shallowly lobate or crispate, penninerved, green or variously

colored with white, yellow, pink, red, or purple.

Sometimes called "laurel" in Honduras; "cintillo" (Veracruz).

This well-known ornamental plant of the tropics is abundant every-

where in the Guatemalan lowlands, being much grown in hedges or

as an ornamental shrub. In the Pacific bocacosta there are many

long roadside hedges bordering the fincas and coffee plantations.

The plant grows easily and thrives with little or no attention. It

is sometimes found more or less naturalized, probably about the

sites of former dwellings. The shrub is grown commonly in the

warmer parts of the United States, especially in Florida, where it

is known by the name Croton, although it is not closely related to

plants of that genus.

CROTON L.

Trees, shrubs, or herbs, the indument usually of stellate hairs or of scales;

leaves mostly alternate, often with 2 sessile or stipitate glands at the base or at

the apex of the petiole, petiolate, entire or dentate, rarely lobate, usually 3-several-

nerved from the base, sometimes penninerved; flowers spicate or racemose,

monoecious or rarely dioecious, the staminate in the upper part of the inflorescence,

the pistillate flowers below, or the two sexes sometimes mixed together; pistillate

flowers solitary under each bract or sometimes with 2-3 staminate ones, the bracts

small; petals usually present in the staminate flowers, often absent or rudimentary

in the pistillate ones; staminate sepals usually 5, valvate or subimbricate, the

disk represented by glands opposite the sepals; stamens 5-many, mostly 10-16,

the filaments inflexed in bud, erect in anthesis; receptacle usually pilose; pistillate

sepals often unequal, the disk annular or of glands; ovary 3-celled, the styles

1-many times bifid or parted; ovules solitary in each cell; capsule separating into

3 bivalvate cocci; seeds smooth, with a small caruncle.

About 700 species, in both hemispheres, most numerous in tropical

regions, only a few species reaching the temperate zones. Other

species besides those listed here are found in southern Central

America, especially in Costa Rica. The genus has received little

serious attention in recent years, so far as North American species

are concerned, and the status of many names is still uncertain.

Plants herbaceous throughout, usually annual.

Leaves deeply palmate-lobate C. lobatus.

Leaves entire or dentate, never lobate.

Leaves entire C. punctatus.

Leaves dentate.

Stems densely and coarsely hispid; leaves green beneath C. hirtus.

Stems stellate-pilose with short hairs; leaves pale beneath.

Leaves acute or acuminate, usually more or less cordate at the base.

C. trinitatis.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 65

Leaves mostly very obtuse, at the base obtuse or subacute . C. glandulosus.

Plants shrubs or trees.

Leaves without glands at the base of the blade or at the apex of the petiole;

pubescence all of closely appressed scales or scale-like hairs, sometimes none.

Leaves acute or rarely obtuse at the base, penninerved C. glabellus.

Leaves cordate to very obtuse at the base, palmate-nerved.

Leaves densely covered on the upper surface with minute appressed hairs,

obtuse or rounded at the apex, mostly about 2 cm. wide . C. punctatus.

Leaves glabrate on the upper surface, usually acute or acuminate, much

larger.

Ovary and fruit smooth, densely lepidote C. guatemalensis.

* Ovary and fruit tuberculate, stellate-hispidulous C. reflexifolius.

Leaves with glands at the base of the blade or at the apex of the petiole, the

glands sometimes (rarely) very small and hard to see, but the pubescence

then of relatively coarse, stellate hairs.

Leaves densely ciliate with long gland-tipped hairs C. ciliatoglandulosus.

Leaves not glandular-ciliate.

Leaves rounded or very obtuse at the apex, less than twice as long as broad

and often as broad as long.

Leaves subentire, densely stellate-tomentose beneath. . .C. payaquensis.

Leaves very coarsely dentate, green and glabrate beneath C. repens.

Leaves acute or acuminate, or, if rarely subobtuse, much longer than broad.

Lowest bracts of the inflorescence embracing both pistillate and staminate

flowers.

Stipules ovate or lanceolate C. draco.

Stipules subulate or setaceous.

Leaf blades more or less cordate at the base, usually conspicuously

so C. callistanthus.

Leaf blades rounded at the base, not at all cordate. C. verapazensis.

Lowest bracts of the inflorescence embracing only pistillate flowers.

Leaves wholly or chiefly penninerved, the lowest nerves not more

conspicuous than the upper ones.

Leaves coarsely crenate C. jutiapensis.

Leaves entire or very finely and evenly serrate.

Leaves glabrous on the upper surface C. Cortesianus.

Leaves pubescent on the upper surface.

Leaves pseudoverti ciliate at the base of the inflorescence;

pistillate flowers crowded at the very base of the inflores-

cence C. adspersus.

Leaves not pseudoverticillate; pistillate flowers not crowded or,

if so, forming an elongate spike.

Leaves obtuse at the base, small, mostly 3.5-6 cm. long.

C. fragilis.

Leaves broadly rounded or shallowly cordate at the base,

mostly 9-16 cm. long or larger.

Glands of the leaves very small and inconspicuous or none.

C. axillaris.

Glands at the base of the leaves large and conspicuous.

C. xalapensis.

66 FIELDIANA: BOTANY, VOLUME 24

Leaves palmate-nerved at the base, the lower nerves much more con-

spicuous than the upper ones.

Pistillate sepals densely covered with very long, gland-tipped hairs.

Leaves glabrate C. glandulosepalus.

Pistillate sepals without gland-tipped hairs, or these very small and

inconspicuous.

Pedicels equaling the fruiting calyx, this reflexed in age.

C. pyramidalis.

Pedicels usually shorter than the fruiting calyx, this not reflexed.

Leaves entire or finely and evenly serrate or dentate, often

cordate at the base.

Inflorescences many-flowered, mostly 10-20 cm. long.

C. xalapensis.

Inflorescences relatively few-flowered, mostly 5 cm. long or

shorter.

Leaves mostly oblong to oblong-ovate C. axillaris.

Leaves ovate or rounded-ovate.

Ovary and capsule long-pilose and densely stellate-

tomentulose; leaves rounded or subcordate at the

base C. lasiopetaloides.

Ovary and capsule merely stellate-tomentulose, without

long spreading hairs; leaves conspicuously cordate

at the base C. limnocharis.

Leaves coarsely and unevenly crenate, dentate, or serrate, not

at all or very obscurely cordate at the base.

Glands at the base of the leaf blade sessile . . . C. quercetorum.

Glands of the leaf blades or petioles conspicuously stipitate.

Leaves practically glabrous on the upper surface, the in-

dument of minute appressed scales without appreciable

branches C. Lundellii.

Leaves thinly or rather densely pubescent on the upper

surface, except sometimes in age, the hairs with easily

visible branches.

Petioles finely stellate-pilose, the branches of the hairs

all of about the same length C. pagiveteris.

Petioles finely stellate-tomentulose and also pilose with

longer spreading hairs.

Columella of the capsule 6-7 mm. long . . C. jalapensis.

Columella of the capsule about 3 mm. long . C. lotorius.

Croton adspersus Benth. PI. Hartweg. 51. 1840. C. botryocarpus

Croizat, Field Mus. Bot. 22: 445. 1942 (type from Jalapa, Jalapa,

Standley 77519). Granadito amarillo; Tomatillo de sensontle; Hierba

de chucho.

Moist thickets or dry open rocky slopes, 1,100-2,000 meters;

Jalapa; Guatemala; Chimaltenango; Huehuetenango. Southern

Mexico.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 67

A slender shrub 2-3 meters high, the young branches and petioles densely

stellate-hispidulous or tomentose; leaves membranaceous, on slender petioles 1 cm.

long or less, the petiole bearing at the apex 2 conspicuous cylindric glands; leaves

alternate, but those at the base of the inflorescence numerous, crowded, and

subverticillate, ovate to lance-ovate, mostly 3-6 cm. long and 1.5-2.5 cm. wide,

acute or acuminate, rounded at the base or obscurely cordate, finely serrulate or

subentire, sparsely and minutely stellate-pilosulous above or in age practically

glabrous, grayish or green beneath, at first stellate-tomentose, in age usually

glabrate, penninerved or the basal nerves sometimes more conspicuous; inflores-

cences solitary or clustered at the ends of the branches, mostly 5-6 cm. long, the

pistillate flowers crowded at the base of the spike, the upper portion staminate,

slender, the flowers soon deciduous; pistillate sepals in fruit 1.5-2 mm. long, linear,

entire; staminate flowers subglobose, the sepals glabrous outside or sparsely stellate-

puberulent; capsule about 6 mm. long, glabrate in age, when young densely stellate-

tomentose and usually bearing a few long slender spreading hairs; seeds ellipsoid,

4 mm. long, brown.

Sometimes called "cuahuilotillo" in Mexico. The several Guate-

malan collections that have been referred to C. botryocarpus seem

to be in no way distinguishable from Mexican material determined

at Kew as C. adspersus, and they match well a photograph of the

type specimen of the latter species.

Croton axillaris Muell. Arg. Linnaea 34: 126. 1865.

Dry or moist, brushy hillsides, 400-1,600 meters; Chiquimula;

Sacatepe"quez; Huehuetenango. Nicaragua, the type from Granada.

A branched shrub about 3 meters high, the branches densely stellate-tomentose

with rather close, often blackish hairs; leaves on rather slender petioles 1-3 cm.

long, lance-oblong or ovate-oblong, 5-13 cm. long, 2-6 cm. wide, acute to long-

acuminate, rounded to subcordate at the base, entire or practically so, sparsely

or densely stellate-pilosulous with very short spreading hairs on both surfaces,

sometimes grayish-tomentose beneath, usually green on both surfaces, penninerved

or sometimes rather conspicuously 5-nerved at the base; inflorescences axillary

and terminal, the axillary ones staminate, dense, sessile; bracts setaceous, 1-

flowered; staminate calyx depressed-globose, the petals lance-obovate, dorsally

glabrous, sericeous within; stamens about 11; staminate calyx 3 mm. broad;

pistillate flowers and fruit unknown.

The only authentic representation of this species that we have

seen is a photograph of the type, formerly in the Berlin herbarium.

Croizat has referred to the species Steyermark 51069 from Huehue-

tenango, and there are available a number of sterile collections

apparently conspecific.

Croton callistan thus Croizat, Journ. Arnold Arb. 21 : 84. 1940.

Llora-sangre.

68 FIELDIANA: BOTANY, VOLUME 24

Wet to rather dry thickets, often in second growth, 300-1,800

meters; known only from Guatemala, but to be expected in Chiapas;

Retalhuleu; Solola; Quezaltenango (type from Colomba, A. F.

Skutch 2025) ; Quiche".

A large shrub or a small tree, often 7-13 meters high, the trunk as much as

18 cm. or more in diameter, usually slender, the branches rather few and stout,

the young branches stout, very densely tomentose and somewhat tuberculate

because of the very unequal, many-branched, stellate, fulvous hairs; leaves very

large, the slender petioles mostly 8-16 cm. long, with 2-3 large glands at or near

the apex; leaf blades membranaceous, broadly ovate or triangular-ovate, as much

as 30 cm. long and 18 cm. wide but mostly smaller, acuminate or long-acuminate,

rather deeply cordate at the base, irregularly and finely dentate or subentire,

palmate-nerved at the base, green above, much paler beneath, thinly or rather

densely covered with minute many-branched hairs, in age often glabrate; inflores-

cences at or near the ends of the branches, sometimes 70 cm. long but mostly

shorter, with very numerous, dense or remote flowers, the flowers glomerate, the

lower bracts with both pistillate and staminate flowers, the upper flowers all

staminate; pistillate flowers on pedicels 5-6 mm. long, the sepals lepidote-tomen-

tose, broadly ovate, 2 mm. long in anthesis; ovary 3 mm. broad, densely covered

with a coarse stellate orange-brown tomentum; staminate flowers slender-pedicel-

late; capsule about 4 mm. long, finally glabrate, densely verruculose.

The leaves have a fetid odor resembling that of Lantana foliage.

The sap, as in other closely related species, is blood-red on exposure

to the air. This species has been confused with C. panamensis

Muell. Arg. and C. draco, to both of which it is closely related. The

taxonomy of this group of Croton species is very imperfectly under-

stood at present, for lack of critical attention. The plant is widely

distributed in Guatemala.

Croton ciliatoglandulosus Ortega, Hort. Matr. Dec. 51. 1797.

Comemano; Hoja de sierra; Hierba mala; Copalito (fide Aguilar);

Ciega-vista; Chirca.

Moist or dry thickets, often in dry rocky places, sometimes in

sandy river bottoms, 200-1,150 meters; Baja Verapaz; El Progreso;

Zacapa; Chiquimula; Jalapa; Jutiapa; Escuintla; Guatemala;

Quiche" ; Huehuetenango. Mexico; Honduras Und Salvador; Cuba.

An erect shrub, seldom more than a meter high but sometimes as much as

2.5 meters, often densely branched, the slender branches densely stellate-pilose;

stipules dissected into numerous long, slender, almost filiform, gland-tipped

divisions; leaves thin and soft, on long slender petioles, rounded-ovate, mostly

3-6 cm. long, abruptly acute or acuminate, rounded or subcordate at the base,

subentire, very conspicuously ciliate with long slender spreading gland-tipped

hairs, palmate-nerved, green above and usually glabrate, usually pale beneath

and densely and softly stellate-pilose; inflorescences short, rather few-flowered,

densely stellate-pilose, the flowers solitary, short-pedicellate; pistillate sepals

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 69

spatulate-lanceolate, glandular-ciliate, accrescent in age; capsule 7 mm. long,

glabrate; seeds lead-colored, undulate-costate.

Sometimes called "ciega-ojo" in Honduras. The plant is easily

recognized among all Central American species of Croton by the

very numerous long gland-tipped hairs and the conspicuous gland-

tipped divisions of the stipules. When taken in the hand all parts

of the plant are extremely viscid, clinging to the skin and leaving

a large amount of an unpleasant and sticky substance. This material

is said to be very dangerous if in contact with the eyes, causing severe

irritation and inflammation, and on this account people leave the

bushes strictly alone and warn strangers against handling them.

It is stated that cattle are sometimes permanently blinded after

feeding on the foliage. The plant has a strong and extremely

unpleasant odor.

Croton Cortesianus HBK. Nov. Gen. & Sp. 2: 83. 1817.

At about 800 meters; Huehuetenango (between Santa Ana

Huista and Nenton, Steyermark 51381). Southern Mexico, the type

from Campeche.

A shrub or small tree as much as 6 meters high, the branches stellate-pilose

with often blackish hairs; leaves firm-membranaceous, on slender petioles 3 cm.

long or shorter, these without apical glands, or the glands very small and in-

conspicuous; leaf blades mostly lance-oblong and 4-12 cm. long, acute or acuminate,

obtuse or rounded at the base, very minutely and closely serrulate or almost entire,

green and glabrous on the upper surface, rather thinly or densely stellate-tomentose

beneath with grayish or sometimes fuscous hairs; inflorescences terminal, mostly

5 cm. long or less, usually very dense, the pistillate flowers numerous, densely

crowded in the lower part of the raceme, the staminate portion of the inflorescence

often very short, sometimes elongate; staminate petals fimbriate-ciliate below;

capsule stellate-hirsute, about 5 mm. long; seeds smooth.

Croton draco Schlecht. Linnaea 6: 360. 1831. Sangre de drago;

Sangre de perro; Llora-sangre ; Calelu (Huehuetenango).

Moist or wet thickets or forest, frequently in second growth,

often on very steep sides of barrancos, 600-1,600 meters; Alta

Verapaz(?); Guatemala(?); Retalhuleu; Quezaltenango; San Marcos;

Huehuetenango. Southern Mexico.

A large shrub or a tree, rarely 20 meters high with a trunk 50 cm. in diameter,

the bark smooth, light brownish or grayish, the branches densely tomentose with

ochraceous or brownish, very unequal, tubercle-like stellate hairs; leaves large,

the rather stout petioles mostly 12 cm. long or shorter, with conspicuous glands

at the apex; leaf blades thin, broadly ovate or deltoid-ovate, mostly 8-20 cm. long,

acute to abruptly caudate-acuminate, shallowly cordate at the base or broadly

rounded, minutely serrulate or subentire, palmate-nerved at the base, green above

70 FIELDIANA: BOTANY, VOLUME 24

and thinly stellate-pubescent or glabrate, usually rough to the touch, beneath

grayish or whitish and often very densely stellate-tomentose; racemes mostly at

the ends of the stout branches, sometimes 60 cm. long, with very numerous, dense

or often remote flowers, the lower bracts subtending both staminate and pistillate

flowers; stamens 15-25; capsule about 5 mm. long, very finely and closely tuber-

culate-tomentulose; seeds lustrous, oval, olivaceous brown, 3 mm. long, somewhat

rugulose or almost tuberculate.

This, like the other related species, is not very well understood,

and it quite possibly includes C. callistanthus. The shrubs or trees

of this alliance are common and conspicuous in the mountains of

the Occidente of Guatemala, often occurring in abundance on the

steep slopes of the barrancos in the lower parts of the vast gorge of

the Rio Samala.

Croton fragilis HBK. Nov. Gen. & Sp. 2: 75. 1817. C. sericeus

Schlecht. Linnaea 5: 85. 1830.

Moist or dry, brushy, rocky hillsides; El Progreso; Chiquimula(?) ;

Guatemala(?); Huehuetenango. Southern Mexico; Honduras(?);

northwestern South America.

A shrub 2.5 meters high or less, the young branches densely stellate-pilose

with whitish or grayish hairs; leaves membranaceous, on petioles 1.5 cm. long or

shorter, the petiole usually with 2 glands at the apex, these conspicuous or hidden

under the pubescence; leaf blades ovate or elliptic-ovate, mostly 3-5 cm. long,

acute or acuminate, obtuse at the base, entire or serrulate, penninerved, finely

stellate-pubescent on the upper surface, usually densely stellate-tomentose beneath;

inflorescences mostly terminal, short and rather few-flowered, dense, the pistillate

flowers subsessile, the staminate slender-pedicellate; stipules subulate; pistillate

sepals ovate-lanceolate, not accrescent; stamens about 15; ovary densely stellate-

tomentose.

This is another species that is poorly understood at the present

time, and when ample material can be assembled and studied criti-

cally, it seems probable that the South American plant and that of

Mexico and Central America will be found distinct. The Maya

name in Yucatan is reported as "tanche."

Groton glabellus L. Sp. PI. ed. 2. 1425. 1763. C. Eluteria Swartz,

Prodr. Veg. Ind. Occ. 100. 1788. C. nitens Swartz, loc. cit. C. perob-

tusus Lundell, Phytologia 1: 405. 1940 (type from Tabasco). Fruta

de chacha; Paujil (fide Aguilar); Perescuch (Pete"n, Maya); Canoh

(reported as the Quecchi name); Cache (Alta Verapaz).

Moist or wet, mixed forest or thickets, sometimes in second

growth, growing on plains or hillsides, 1,000 meters or less, mostly

at 500 meters or lower; Pete"n; Alta Verapaz; Izabal; Suchitepe"quez ;

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 71

Retalhuleu ; San Marcos ; Huehuetenango ; Quiche". Southern Mexico ;

British Honduras to Panama; West Indies; northern South America.

A large shrub or a tree, often flowering when only 2-3 meters high but com-

monly a tree of 6-12 meters, the trunk sometimes 30 cm. or more in diameter,

the young branchlets covered with closely appressed, brown scales; leaves on

eglandular petioles 1-3 cm. long, firm-membranaceous, mostly elliptic-oblong or

lance-oblong, sometimes broader, mostly 5-16 cm. long, usually abruptly acumi-

nate or caudate-acuminate, sometimes obtuse or even narrowly rounded at the

apex, usually subacute or obtuse at the base, entire, penninerved, bearing ap-

pressed brown scales on both surfaces, these usually very few on the upper surface

but often dense on the lower surface; racemes simple or branched, axillary, generally

shorter than the leaves, dense or interrupted, the flowers whitish, somewhat

fragrant; pedicels of the staminate flowers 2 mm. long, in the pistillate flowers

6 mm. long; staminate sepals ovate-triangular, stellate-pubescent, petals spatulate,

pellucid-punctate; capsule oblong-globose, lepidote, tuberculate, 7-10 mm. long;

seeds brown, smooth.

Maya names in Yucatan are "cuxub," "cocche," and "chuts";

"copalchi" (Tabasco); "wild cinnamon" (British Honduras); "cas-

carilian," "lian," "barenillo" (Honduras). A very common small

tree in many parts of the Guatemalan lowlands. C. perobtusus is a

leaf form in which the leaves are very obtuse or rounded at the apex,

a character in which there is much variation.

Croton glandulosepalus Millsp. Field Mus. Bot. 2: 419. 1916.

British Honduras (Maskall Pine Ridge, P. H. Gentle 1170).

Yucatan Peninsula of Mexico.

A slender branched shrub, the older branches gray or whitish, the young

branchlets rather densely stellate-pilose; leaves thin, eglandular, on slender petioles

2-3 cm. long, lance-ovate to oblong-lanceolate, mostly 5-11 cm. long, gradually

or abruptly acuminate or long-acuminate, rounded or obtuse at the base, entire,

3-nerved from the base, when young thinly and very finely stellate-pubescent, in

age almost wholly glabrous, paler beneath; stipules setaceous, 5-14 mm. long;

racemes slender, dense or somewhat interrupted, terminal, mostly 4-7 cm. long;

staminate flowers short-pedicellate, the sepals stellate-tomentulose, ovate, the

petals glabrous or nearly so; stamens 10; pistillate flowers on short thick pedicels,

few, the sepals ovate-lanceolate, fleshy, densely covered with reddish glands

terminating in long slender hairs; ovary densely stellate-tomentulose; capsule

6 mm. long; seeds brown, smooth, 4 mm. long.

This species may be recognized at once by the very dense covering

of long-stalked glands on the pistillate sepals.

Croton glandulosus L. Syst. ed. 10. 1275. 1759.

Dry grassy hillsides or on sandbars along streams, 200-1,250

meters; reported, perhaps in error, from Pete"n; Zacapa; El Progreso;

72 FIELDIANA: BOTANY, VOLUME 24

Quich4. Southern United States; Mexico; West Indies; tropical

South America.

An erect annual, mostly 50 cm. high or less, usually branched, the stems

stellate-pilose with appressed hairs; leaves long-petiolate, with 2 saucer-shaped

glands beneath at the base of the blade; leaf blades oblong-ovate to ovate or

elliptic, 2-3.5 cm. long, rounded or very obtuse at the apex, obtuse at the base,

coarsely dentate or crenate, stellate-pilose on both surfaces; stipules linear, 2 mm.

long, deciduous; racemes short and few-flowered, 2 cm. long or less, the flowers

subsessile; staminate sepals 2 mm. long, the petals slightly longer; stamens 10;

pistillate sepals unequal, spatulate, 3 mm. long, accrescent in age; ovary hirsute;

capsule 5-6 mm. long, glabrate or with a few stellate hairs; seeds 4 mm. long,

bearing dorsal rows of minute pits.

Croton guatemalensis Lotsy, Bot. Gaz. 20: 353. pi. 25. 1895.

C. eluterioides Lotsy, op. cit. 352. pi. 25 in part. 1895 (type from

Santa Rosa, Dept. Santa Rosa, Heyde & Lux 3470). Copalchi;

Zicche (Pete"n) ; Quina (fide Tejada).

Wet to dry thickets or rather thin, mixed forest, often on rocky

hillsides, 1,800 meters or less, mostly at rather low elevations, often

planted for hedges or as windbreaks in coffee plantations; Pete'n;

Alta Verapaz; Baja Verapaz; Chiquimula; Santa Rosa (type from

Santa Rosa, Heyde & Lux 3035); Escuintla; Guatemala; Sacate-

p^quez; Chimaltenango; Suchitepe"quez; Retalhuleu; Quiche"; Quezal-

tenango. Southern Mexico; Honduras and Salvador and perhaps

farther southward.

A rather slender shrub or tree, sometimes 8 meters high, usually lower, usually

densely appressed-lepidote throughout with whitish to brownish scales; leaves

firm-membranaceous, mostly on long slender petioles, or the petioles sometimes

short, eglandular; leaf blades ovate to broadly triangular-ovate, mostly 7-15 cm.

long, acuminate to long-acuminate, shallowly cordate or truncate at the base,

entire, palmate-nerved at the base, green on the upper surface and in age often

almost glabrate, sparsely or very densely lepidote beneath, often silvery and

whitish; racemes axillary, often very numerous, usually much shorter than the

leaves, many-flowered, the flowers often remote, almost sessile, densely lepidote;

sepals ovate, acute; petals ovate-lanceolate, ciliate, glabrous; stamens about 15;

ovary densely lepidote; capsule subglobose, about 8 mm. long, very densely

lepidote, sometimes obscurely tuberculate.

Known in Yucatan by the Maya names "chul" and "chulche";

"copalchin" (Oaxaca). This shrub or small tree is very common in

many parts of Guatemala, but especially on the Pacific plains and

foothills, where often it forms thickets of wide extent. It and

C. reflexifolius are much planted for hedges or windbreaks around

coffee plantations that are exposed to the wind. The plant of

Guatemala and southern Mexico has been referred generally to

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 73

C. niveus Jacq., a species of northwestern South America and southern

Central America (Costa Rica; Panama), which may be distinct.

For a discussion of the subject see Croizat, Field Mus. Bot. 22: 447.

1942. That author thinks that C. eluterioides may be a distinct

species, but this seems highly improbable, since both it and C.

guatemalensis were described from the town of Santa Rosa, where

they probably were growing in coffee plantations. The bark of this

and related species of Croton is said to have a bitter taste, and on

that account it inevitably found application in local medicine as a

"remedio" for intermittent fevers. The bark is said to have been

exported formerly to Europe for medicinal purposes.

Croton hirtus L'HeY. Stirp. Nov. 17. 1784. C. glandulosus var.

hirtus Muell. Arg. in DC. Prodr. 15, pt. 2: 684. 1866.

Open fields or hillsides, 900 meters or less; Pete"n; Suchitepe*quez ;

Huehuetenango. Southern Mexico; Honduras to Panama; West

Indies; tropical South America.

An erect, rather stout annual, commonly 60 cm. high or less, simple or usually

branched, the stems and petioles yellow-hispid with long stiff hairs; leaves mem-

branaceous, long-petiolate, rhombic-ovate, often broadly so, 3-7 cm. long, obtuse

or acute, rounded or obtuse at the base, coarsely crenate, 3-5-nerved from the base,

with 2 long-stipitate glands at the base, stellate-hirsute or in age glabrate; stipules

linear, 3-5 mm. long; racemes 1-several at the ends of the branches, 1.5-3 cm.

long, the flowers subsessile; bracts linear, with conspicuous stipitate glands;

staminate sepals elliptic, acute, stellate-hispidulous, the petals slightly longer;

stamens about 10; pistillate sepals unequal, lanceolate or spatulate, 3-4 times as

long as the capsule; ovary hirsute; capsule 3-4 mm. long, globose, hirsute; seeds

2.5-3 mm. long, with a small caruncle.

Croton jalapensis Croizat, Field Mus. Bot. 22: 449. 1942.

Moist or dry thickets, 1,300-2,500 meters; Jalapa (type from

Jalapa, Standley 76414); Guatemala; Huehuetenango; endemic.

A shrub or small tree 2-6 meters high, densely branched, the young branches

densely stellate-hispidulous; leaves on petioles 2-2.5 cm. long, firm-membranaceous,

ovate or broadly elliptic-ovate, 5-11 cm. long, acuminate or long-acuminate,

rounded or obtuse at the base, coarsely and irregularly serrate, green above,

sparsely stellate-pubescent or glabrate, rough to the touch, paler and often ochra-

ceous beneath, densely stellate-pubescent, 3-5-nerved at the base; petiole bearing

at the apex 2 conspicuous stipitate glands; racemes short or usually elongate and

as much as 11 cm. long; pistillate flowers remote, almost sessile; ovary densely

covered with a yellowish or pale orange, stellate indument; capsule 9 mm. long,

stellate-tomentulose, the columella 5 mm. long; seeds 5.5 mm. long, carunculate,

rugose-costate.

It is worthy of mention that C. jalapensis and C. xalapensis

HBK. were named for two towns having the same name, Jalapa

74 FIELDIANA: BOTANY, VOLUME 24

in Guatemala and Jalapa in Veracruz, C. xalapensis when the old

Spanish spelling of certain geographic names was still in vogue.

However, since the two species names differ in their initial letter

and appear very different to northern European ears, if not to

Spanish ones, both may be retained in the genus.

Croton jutiapensis Croizat, Field Mus. Bot. 22: 450. 1942.

Moist or dry, brushy, often rocky plains and hillsides, 300-900

meters; Zacapa; Chiquimula; Jalapa; Jutiapa (type from Jutiapa,

Standley 74971); Huehuetenango. Honduras.

A shrub 1-3 meters high, the branches rather stout, when young stellate-

hispid with ochraceous or ferruginous hairs, in age ferruginous or fuscous; leaves

thick-membranaceous, on thick petioles 1.5 cm. long or less, the petiole with 2

stipitate glands at the apex; leaf blades ovate to ovate-lanceolate, mostly 4-11

cm. long, acute or acuminate, usually somewhat narrowed to the base, the base

itself broadly cuneate to subcordate, coarsely and unequally serrate or crenate,

penninerved, softly stellate-pubescent on the upper surface, paler and densely

velutinous-tomentose beneath, the nerves usually very prominent beneath, the

lateral nerves about 6 pairs; inflorescences mostly terminal and shorter than the

leaves, lax and rather few-flowered; pistillate sepals unequal, in fruit as much as

12 mm. long, spatulate or linear; ovary densely hispid-tomentose.

Croton lasiopetaloides Croizat, Field Mus. Bot. 22: 450. 1942.

Open oak forest, about 2,000 meters; endemic; Huehuetenango

(type from mountains west of Aguacatan, on the road to Huehue-

tenango, Standley 81219).

A shrub 1-1.5 meters high, the branches slender, laxly stellate-tomentose at

first; petioles 1-2 cm. long, with 2 small obscure glands at the apex; leaf blades

membranaceous, broadly ovate or orbicular-ovate, 5-8 cm. long, 3.5-7 cm. wide,

acute or abruptly short-acuminate or sometimes broadly obtuse, rounded at the

base and often subcordate, stellate-pubescent above at first, soon glabrate and

green, densely and softly stellate-tomentose beneath, whitish or grayish, minutely

denticulate or almost entire, about 5-nerved at the base; inflorescences terminal,

solitary 5.5 cm. long or less, the flowers subsessile; staminate calyx densely stellate-

pilose, globose in bud; pistillate sepals 2 mm. long, stellate-tomentose; ovary

whitish-hispidulous; capsule subglobose, densely and closely stellate-tomentulose

and sparsely long-pilose, the columella 6 mm. long; seeds oval, very lustrous,

castaneous, 4 mm. long.

Croton limnocharis Croizat, Field Mus. Bot. 22: 451. 1942.

Moist or dry, brushy, often rocky plains and hillsides, 300-1,400

meters; endemic; Zacapa; Chiquimula; Jutiapa (type from Jutiapa,

Standley 75215).

A shrub or small tree, 2-4 meters high or perhaps larger, the young branchlets

very densely and closely stellate-tomentulose with whitish hairs, in age cinnamom-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 75

eous; petioles slender, 2.5 cm. long or shorter, with 2 sessile saucer-shaped glands

at the apex; leaf blades thin, ovate or broadly ovate, 6-13 cm. long, 3-7.5 cm. wide,

abruptly acuminate or long-acuminate, shallowly but conspicuously cordate at

the base, obscurely undulate-denticulate or almost entire, 5-nerved at the base,

green on the upper surface and almost glabrate but with scattered minute stellate

hairs, pale beneath and closely stellate-tomentose; stipules minute and deciduous;

inflorescences terminal, very short, few-flowered; pistillate flowers few, short-

pedicellate, the sepals triangular, 1-1.5 mm. long; ovary whitish-tomentulose;

capsule subglobose, 7 mm. long, at first densely and minutely stellate-tomentulose

but the tomentum easily removed and the capsule often glabrate; seeds 4.5 mm.

long, smooth.

Croton lobatus L. Sp. PI. 1005. 1753. Malm (Suchitepe'quez).

Open fields or moist thickets, frequently a weed in cultivated

or waste ground, often on sandbars along streams, 400 meters or

less; Pete"n; Alta Verapaz; Izabal; Zacapa; Escuintla; Suchitepe'quez.

Southern Mexico; British Honduras to Salvador and Panama; West

Indies; South America; tropical Africa.

An erect branched annual, usually 75 cm. high or less, the stems green, when

young thinly stellate-hispidulous; stipules subulate; leaves on slender eglandular

petioles 3-10 cm. long, digitately and deeply 3-5-lobate, membranaceous, sparsely

stellate-pubescent or almost glabrous, green, the segments oblanceolate, acuminate

or caudate-acuminate, narrowed below, 3-5.5 cm. long; inflorescences terminal or

axillary, 10 cm. long or less, usually remotely flowered; staminate flowers short-

pedicellate, the sepals elliptic, glabrous, the petals lanceolate or oblanceolate,

glabrous; stamens 10-13; pistillate flowers subsessile, the sepals linear or lanceolate,

acute, with a few gland-tipped hairs on the margins; ovary stellate-pubescent and

pilose; capsule 8 mm. long, stellate-pubescent and setose, becoming glabrate;

seeds 5 mm. long.

A weedy plant, seldom plentiful in Central America.

Croton lotorius Croizat, Journ. Arnold Arb. 26: 185. 1945.

Sanalotodo.

Known only from the type, Huehuetenango, between Santa Ana

Huista and forest of Rancho Lucas, 800-900 meters, Steyermark

51332.

A shrub 1.5 meters high, the branches laxly cinereous-tomentulose; leaves on

slender petioles 1-2 cm. long, ovate, grayish green, acuminate, rounded at the

base, glabrate above, grayish-tomentose beneath, 2.5-5 cm. long, 1.5-3 cm. wide,

3-5-plinerved, obtusely dentate or duplicate-dentate; petiole bearing at the apex

2 stipitate glands, the stipules acute or setaceous; spikes slender, about 10 cm.

long; staminate perianth 1.5-2 mm. broad, the stamens few; pistillate perianth

1.5-2 mm. long, subcampanulate, hispidulous, cleft almost to the base; ovary glo-

bose, hispidulous.

76 FIELDIANA: BOTANY, VOLUME 24

Croton Lundellii Standl. Carnegie Inst. Wash. Publ. 461: 67.

1935. C. petensis Lundell, Phytologia 1: 406. 1940 (type from

Aguada Tigre-Yaxha road, Pete"n, C. L. Lundell 4128).

Open forest or thickets, 1,500 meters or less; Pete"n; Guatemala;

Huehuetenango(?). Campeche, the type from Tuxpena.

A shrub or small tree, the branches rather stout, when young densely ap-

pressed-stellate-lepidote, soon glabrate; petioles stout, 1.5-3 cm. long, bearing at

the apex 2 large crateriform glands; leaf blades chartaceous or firm-membranaceous,

oblong or ovate-oblong, mostly 7-11 cm. long and 3-5 cm. wide, somewhat nar-

rowed to the obtuse apex, rounded or very obtuse at the base, irregularly sinuate-

crenate or doubly serrate, appressed-stellate-lepidote above at first, soon glabrate,

pale beneath or often almost silvery, densely appressed-lepidote, in age often

glabrate, 3-nerved at the base; racemes as much as 20 cm. long, usually long-

pedunculate, lax, many-flowered; pistillate flowers remote, solitary or fasciculate,

sessile; staminate calyx densely stellate-lepidote; capsule 5-6 mm. long, stellate-

lepidote, smooth; seeds blackish brown or almost black, lustrous, almost 5 mm.

long.

The inflorescences are sometimes fantastically distorted by large

insect galls.

Croton pagiveteris Croizat, Journ. Arnold Arb. 21: 85. 1940.

Copalchi.

On limestone hillsides, 800-900 meters; endemic; Huehuetenango

(type from Pueblo Vie jo, Seler 2776; collected also near Santa Ana

Huista); Baja Verapaz(?).

A shrub 3 meters high, the young branches densely ochraceous-tomentose

with stellate hairs, in age reddish brown; petioles slender, 2-5.5 cm. long, bearing

2 conspicuous stipitate glands at the apex; leaf blades firm-membranaceous, ovate

or elliptic-ovate, mostly 6-12 cm. long and 3-7 cm. wide, acute or short-acuminate,

rounded or broadly cuneate at the base, remotely and inconspicuously crenate-

serrate or serrulate, stellate-puberulent above at first but in age green and almost

glabrate, paler beneath and stellate-tomentulose, 5-nerved at the base; racemes

terminal, 13 cm. long or less, pedunculate, interrupted, the flowers glomerate or

solitary; pistillate flowers almost sessile, the sepals 1.5 mm. long, erect, triangular;

ovary ochraceous-tomentulose; staminate flowers globose in bud, short-pedicellate,

tomentulose.

Croton ortholobus Muell. Arg. (Flora 55: 9. 1872) was described

as coming from Guatemala, but the plant actually was collected by

Friedrichsthal at Cartago, Costa Rica.

Croton payaquensis Standl. Journ. Wash. Acad. Sci. 14: 97.

1924. Hierba mala.

Moist or dry, brushy, often rocky plains and hillsides, sometimes

along rocky stream beds, 400-1,500 meters; Zacapa; Chiquimula;

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 77

Jalapa; Jutiapa. Salvador, the type from Cerro de la Olla near

Chalchuapa; Honduras.

A stout shrub, usually only 30-60 cm. high, with few branches, the stems very

densely stellate-tomentose with mostly appressed, fulvous or yellowish hairs;

stipules subulate, entire, caducous; petioles stout, 0.5-7 cm. long, the glands small

and inconspicuous or none; leaf blades oblong-ovate to oval or suborbicular,

2.5-10.5 cm. long, 1.6-8.5 cm. wide, rounded at the apex, sometimes emarginate,

broadly rounded or subcordate at the base, thick and flannel-like, entire or nearly

so, 5-nerved at the base, very densely stellate-tomentose on both surfaces with a

fulvous tomentum, in age sometimes glabrate and green above; inflorescences

mostly terminal, sometimes 6 cm. long and many-flowered but usually very short

and few-flowered, the flowers short-pedicellate; staminate flowers subglobose in

bud and 1-1.5 mm. in diameter, petaliferous, densely stellate-tomentose; stamens

about 8; pistillate calyx stellate-pubescent, the sepals lance-oblong, acute, sub-

equal; capsule sparsely stellate-puberulent, the columella 4 mm. long; seeds almost

black, very lustrous, 4 mm. long, nearly smooth.

Called "friega-plato" in Salvador. This low shrub is very com-

mon in some hilly regions of the Oriente.

Croton punctatus Jacq. Coll. Bot. 1: 166. 1786.

Usually on sandy sea beaches; British Honduras; doubtless

occurring on the coast of Izabal. Southeastern United States;

Mexico; Honduras to Panama; northern South America.

Plants probably perennial, erect or diffuse, a meter high or less, sometimes

suffrutescent below, the stout stems densely appressed-stellate-pubescent, brown-

ish; leaves thick, on long stout eglandular petioles, elliptic to ovate or oblong,

1-5 cm. long, rounded or obtuse at the apex, rounded at the base, entire, very

densely and finely stellate-pubescent with subappressed hairs, these often brown,

at least in part; flowers monoecious or dioecious; staminate racemes rather few-

flowered, interrupted, 1-2 cm. long, the pedicels 2-4 mm. long; petals none or

rudimentary; stamens about 12; pistillate racemes about 1 cm. long and 1-3-

flowered; sepals equal, cuneate or oblong; petals none; capsule depressed-globose,

5-8 mm. high; seeds 6 mm. long, dark or variegated, the caruncle large, sub-

stipitate.

The Maya name in Yucatan is reported as "zac-chunum";

"hierba de jabali" (Yucatan). This is a characteristic strand plant,

but it is not common in Central America.

Croton pyramidalis Bonn. Smith, Bot. Gaz. 35: 7. 1903.

Hediondilla (Huehuetenango).

Moist or wet, mixed forest, 450 meters or less; Alta Verapaz

(type from Rio Dolores near Cubilgliitz, Tuerckheim 7974); Hue-

huetenango. Southern Mexico; British Honduras; Honduras.

A large shrub or a tree, sometimes 12 meters high, the bark thin, rather

smooth, yellowish brown or grayish, the young branches slender, very densely

78 FIELDIANA: BOTANY, VOLUME 24

lepidote with brownish or silvery scales; petioles slender, 3-7 cm. long, 2-glandular

at the apex; leaf blades firm-membranaceous, oblong-ovate to broadly ovate, 10-20

cm. long and 5-13 cm. wide, abruptly long-acuminate or short-acuminate, broadly

rounded or shallowly cordate at the base, entire or nearly so, bright green above

and glabrous or nearly so, very densely appressed-lepidote beneath and usually

silvery, sometimes brownish in age; racemes solitary or often forming large uni-

sexual panicles, these sometimes 20 cm. long, openly branched, the flowers pedicel-

late; staminate calyx 4 mm. long, the sepals ovate, glabrous within; petals equaling

the calyx, villosulous outside; stamens about 15; ovary densely lepidote; capsule

globose, 5.5 mm. in diameter; seeds lustrous, black, rugulose.

Called "sangre de grado" in Honduras; "cascarillo bianco"

(Veracruz). The wood is whitish. This is perhaps the plant reported

by Hemsley from Guatemala as C. Billbergianus Muell. Arg.

Croton quercetorum Croizat, Field Mus. Bot. 22: 452. 1942.

Known only from the type, Jalapa, mountains about Chahuite,

northwest of Jalapa, moist oak forest, 1,650 meters, Standley 77460.

A shrub or tree as much as 8 meters high, the young branches densely covered

with a very close, uneven, fulvous, stellate tomentum; petioles slender, 2.5-3.5

cm. long, bearing about 4 patelliform glands at the apex; leaf blades ovate-elliptic

or ovate, mostly 7-14 cm. long, acuminate or long-acuminate, rounded or obtuse

at the base, coarsely and unevenly dentate, thick-membranaceous, 5-nerved at

the base, green above, glabrate, sparsely stellate-puberulent, rather rough to the

touch, somewhat paler green beneath and similarly pubescent; stipules 15 mm.

long or less, persistent; inflorescences terminal, simple, 20 cm. long or less; pistillate

flowers almost sessile, the sepals triangular, 4 mm. long, stellate-tomentose;

capsule 12-14 mm. long, densely and finely stellate-tomentulose; seeds 10 mm.

long, costate-rugulose.

Croton reflexifolius HBK. Nov. Gen. & Sp. 2: 68. 1817. Copal-

chi; Hoja amarga.

Dry to wet thickets or thin forest, on plains or hillsides, sometimes

on limestone, 1,500 meters or less; Pete"n; El Progreso; Jutiapa;

Suchitepe"quez ; Huehuetenango. Southern Mexico ; British Honduras

to Salvador and Costa Rica.

A shrub or small tree, commonly 2-8 meters high, the branches slender, at

first densely appressed-lepidote, grayish or brownish; leaves on long slender

eglandular petioles, ovate to broadly triangular-ovate, mostly 8-15 cm. long,

usually cuspidate-acuminate, broadly rounded to truncate or shallowly cordate

at the base, entire, thick-membranaceous or chartaceous, 3-5-nerved at the base,

usually green and almost glabrous above, beneath very densely lepidote with

closely appressed scales, somewhat ferruginous or silvery; stipules small, ovate;

racemes many-flowered, dense or somewhat interrupted, most of them wholly

staminate, the flowers sessile or short-pedicellate, the racemes thus spike-like,

commonly 4-8 cm. long; staminate sepals lance-ovate, the petals lance-obovate,

pubescent; stamens about 10; pistillate flowers few, in age long-pedicellate, the

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 79

sepals 3 mm. long, not accrescent; ovary stellate-pubescent and sometimes pilose;

capsule almost or fully 1 cm. long, muricate or almost echinate; seeds smooth.

Sometimes called "sasafras" in Salvador, a name derived from one

of the languages of the Indians of the eastern United States and

applied properly to a quite different tree of the Lauraceae (Sassa-

fras). It would be interesting to learn how this name of remote

origin reached Central America (it is sometimes used also in South

America). Unless mature capsules are present, it is difficult to

separate this species from C. guatemalensis and C. niveus Jacq., and

one wonders whether there is any important or essential difference

between these supposed species. The distribution data given here

for C. reflexifolius and C. guatemalensis, as represented in Guatemala,

are not wholly dependable, since few of the numerous available

specimens bear mature fruit. However, since the two forms are

alike in almost every character, this is scarcely of practical impor-

tance. The dry leaves of these two species are sold commonly in

the markets as remedios, to be used in home treatment of malaria,

inflammation, and other affections, and as a tonic. In Salvador the

bark is used for flavoring some alcoholic beverages. The leaves

have a strong aromatic odor. The Maya name in Yucatan is re-

corded variously as "perexcutz," "pereschuch," and "pelezcutz."

Croton repens Schlecht. Linnaea 19: 237. 1847. Tostoncillo;

Toston; Chacotote (Izabal).

Dry rocky brushy fields and hillsides, often in open pine forest

or in savannas, 1,500 meters or less; Pete*n; Alta Verapaz; Izabal;

Chiquimula; Jalapa; Jutiapa; Santa Rosa; Guatemala (Fiscal).

Southern Mexico; British Honduras; Honduras; Salvador.

Plants a meter high or usually lower, arising from a hard woody root, the

stems slender, tough, often several, simple or branched, woody throughout or nearly

so; petioles slender or stout, 2 cm. long or usually much shorter, with 2 conspicuous

glands at the apex; leaf blades rather thick, broadly ovate to suborbicular, generally

2-5 cm. long, mostly rounded or very obtuse at the apex, broadly rounded or

usually subcordate at the base, very coarsely and irregularly dentate or duplicate-

dentate or sometimes shallowly lobate, 5-nerved at the base, thinly stellate-

puberulent on both surfaces, very rough to the touch; racemes mostly terminal,

usually much shorter than the leaves, few-flowered; staminate calyx densely

stellate-pilosulous, the petals lanceolate, pilose at the apex; pistillate sepals some-

what accrescent, lance-ovate, obtuse or subacute, densely stellate-pubescent,

spreading in age; capsule 5-6 mm. long, finely tuberculate, pilose with long spread-

ing white hairs.

One of the characteristic low plants of open pine or oak forest

in the mountains and foothills of eastern Guatemala.

80 FIELDIANA: BOTANY, VOLUME 24

Croton trinitatis Millsp. Field Mus. Bot. 2: 57. 1900. C.

Miquelensis Ferguson, Kept. Mo. Bot. Gard. 12: 48. 1901. C.

tragioides Blake, Contr. U. S. Nat. Herb. 24: 12. 1922 (type from

Lago de Izabal, Izabal, S. F. Blake 7854). C. Miquelianus Lanjouw

in Pulle Fl. Surinam 2: 38. 1932.

Moist or wet fields or open pine forest, often a weed in cultivated

ground, especially banana plantations, or in waste ground about

settlements, 300 meters or less; Alta Verapaz; Izabal. Southern

Mexico; British Honduras to Panama; West Indies; tropical South

America.

An erect annual a meter high or usually lower, usually sparsely branched,

slender, the branches stellate-pilosulous or glabrate; petioles 2-10 mm. long, with

2 stipitate glands at the apex or on the base of the blade; leaf blades ovate or

deltoid-ovate, 2-4 cm. long, acute, usually shallowly cordate at the base, coarsely

crenate, green above, thinly long-pilose or almost glabrate, paler beneath, stellate-

pilose; stipules subulate; racemes axillary or terminal, 1-2 cm. long, the bracts

triangular, 2-lobate at the base, the flowers short-pedicellate; staminate flowers

4-5-parted, the sepals ovate-triangular, pilose outside; stamens 8-10; pistillate

sepals often unequal, stellate-pubescent, the petals rudimentary, subulate; ovary

hirsute; capsule 3-4 mm. long; seeds dark olive-gray, lustrous, 3 mm. long, finely

and obscurely impressed-puncticulate in longitudinal lines.

Called "quema-nariz" in Honduras; "wild sage" (British Hon-

duras).

Croton Tuerckheimii Donn. Smith is Olmediella Betschleriana

(Goepp.) Loes. of the family Flacourtiaceae.

Croton verapazensis Donn. Smith, Bot. Gaz. 54: 242. 1912.

Brushy dry rocky hillsides, about 1,600 meters; Baja Verapaz

(type from Santa Rosa, Tuerckheim 11.2297); Jalapa. Endemic.

A tree of 6-9 meters, the branchlets densely stellate-tomentulose; petioles

long and slender, with 2 stipitate glands at the apex; leaf blades membranaceous,

broadly ovate or rounded-ovate, 8-16 cm. long, 7-12 cm. wide, acute or acuminate,

broadly rounded at the base, minutely and inconspicuously denticulate or almost

entire, 5-nerved at the base, sparsely and very finely stellate-pubescent on both

surfaces or glabrate above, only slightly paler beneath; racemes terminal, about

12 cm. long, the lowest bracts embracing both staminate and pistillate flowers,

the pedicels 5 mm. long or often much shorter; stamens about 15; pistillate sepals

2.5 mm. long, oblong-ovate, the petals rudimentary; staminate petals oblong-el-

liptic, 2.5 mm. long; capsule stellate-pubescent, 1 cm. long; seeds 5 mm. long,

fuscous, slightly rugose.

Croton xalapensis HBK. Nov. Gen. & Sp. 2: 85. 1817. C.

pseudoxalapensis Croizat, Journ. Arnold Arb. 21: 85. 1940 (type

from Siguatepeque, Dept. Comayagua, Honduras — given in the

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 81

original publication erroneously as "Dept. of Comaguaya, vicinity

of Siguatepec")- C. pseudoxalapensis var. cobanensis Croizat, op.

cit. 86 (type from Coban, Alta Verapaz, Tuerckheim 11.1015).

Drago; Chirca; Llora-sangre.

Moist or wet thickets, 2,000 meters or less, most frequent at low

elevations; Baja Verapaz; El Progreso; Jutiapa; Santa Rosa; Guate-

mala; Huehuetenango. Southern Mexico; British Honduras to

Costa Rica and Panama.

A shrub or small tree 1-6 meters high, sparsely or densely branched, the

branches stout, densely ochraceous-tomentose with stellate hairs or in age glabrate;

petioles 2-3 cm. long, with 2 conspicuous glands at the apex; leaf blades ovate

or oblong-ovate, mostly 9-20 cm. long, long-acuminate or cuspidate-acuminate,

rounded and cordate at the base, usually shallowly so, serrulate or subentire,

membranaceous, essentially penninerved but the basal nerves often conspicuous,

the lateral nerves about 10 pairs; racemes 25 cm. long or less, with very numerous

flowers, dense or often interrupted; pistillate flowers on very short pedicels, the

sepals triangular-acute, 3.5 mm. long; ovary densely yellowish-tomentose; stami-

nate flowers short-petiolate; capsule subglobose, sparsely or rather densely stellate-

puberulent, the columella about 5 mm. long.

Called "pela-nariz" in Honduras. This has been reported from

Guatemala as C. panamensis Muell. Arg. To the synonymy of

C. xalapensis probably belong C. Aguilarii Lundell (Phytologia 1:

401. 1940; type from La Libertad, Pete"n, M. Aguilar 463) and C.

aster -aides Lundell (op. cit. 402; type from Vaca, El Cayo District,

British Honduras, P. H. Gentle 2218).

DALECHAMPIA L.

Reference: F. Pax, Pflanzenreich IV. 147, xii: 3-56. 1919.

Plants woody or almost wholly herbaceous, usually twining and scandent,

rarely erect shrubs, glabrous or with pubescence of simple hairs, the hairs sometimes

stinging; leaves alternate, petiolate, undivided or 3-5-lobate or digitately 3-5-

foliolate, 2-stipulate, generally 2-stipellate at the apex of the petiole; flowers

monoecious, apetalous, terminal or axillary, sometimes fasciculate, surrounded by

2 large, often colored bracts, the inflorescence with 3 pistillate flowers below and

numerous or few staminate flowers above, the flowers short-pedicellate, the in-

florescences sometimes wholly staminate and dichasial; staminate calyx in bud

globose and closed, in anthesis valvately 4-6-parted; disk none; stamens 15-30

or more, inserted on a convex receptacle or on a column, the filaments short;

anthers erect, generally didymous, longitudinally dehiscent; pistillate sepals 5-12,

imbricate, narrow, mostly pinnatifid, usually accrescent and indurate after

anthesis and surrounding the capsule like an involucre; disk none, or rarely

annular; ovary 3-celled, the styles connate into a long column, the ovules solitary

in each cell; capsule usually tridymous, separating into 2-valvate cocci, these

separating from a persistent columella, the endocarp crustaceous or ligneous;

seeds globose, not carunculate; endosperm carnose, the cotyledons broad and flat.

82 FIELDIANA: BOTANY, VOLUME 24

About 90 species, mostly in tropical America, a few in tropical

Africa and southern Asia. At least one other species is found in

southern Central America.

Plants erect shrubs; leaves simple, undivided, obovate-oblong or oblanceolate-

oblong, attenuate to the base D. spathulata.

Plants scandent, the stems mostly or wholly herbaceous; leaves usually cordate

at the base, often deeply lobate or digitately compound.

Leaves simple, not lobate, or compound leaves sometimes mixed with the simple

ones.

Leaves not uniform, some of them simple and undivided, others digitately

3-foliolate D. heteromorpha.

Leaves uniform, all simple and unlobed.

Bracts pink D. Schippii.

Bracts green.

Leaves glabrous beneath or nearly so; stems and petioles puberulent or

glabrate; bracts merely dentate D. laevigata.

Leaves rather densely and finely pubescent beneath; stems and petioles

hirsute; bracts shallowly laciniate-lobate D. Friedrichsthalii.

Leaves deeply lobate or digitately 3-foliolate, in one species the leaves partly

3-foliolate and partly undivided.

Leaves dimorphous, some of them 3-foliolate, others simple and undivided.

D. heteromorpha.

Leaves uniform, all lobate or digitately compound.

Leaves 3-foliolate.

Leaflets glabrous beneath or nearly so except on the veins . . D. panamensis.

Leaflets densely velutinous-pilosulous beneath D. molliuscula.

Leaves simple, deeply lobate.

Involucral bracts green, deeply 3-lobate D. scandens.

Involucral bracts cream-colored, shallowly 3-dentate at the apex.

D. tiliifolia.

Dalechampia Friedrichsthalii Muell. Arg. Flora 55: 45. 1872.

Moist thickets, 240 meters; Retalhuleu (near Retalhuleu, Standley

88554). Nicaragua; Panama.

A small herbaceous vine, the slender stems puberulent and also hirsute with

long spreading fulvous hairs; leaves on petioles 8 cm. long or less, ovate or broadly

ovate, 7-13 cm. long, 5-8.5 cm. wide, abruptly acute or cuspidate, deeply and

narrowly cordate, the basal lobes often overlapping, denticulate, palmate-nerved,

pilose above, more densely pubescent beneath with short hairs; stipules linear-

lanceolate, 8-12 mm. long, reflexed; inflorescences terminal or axillary, the in-

volucral bracts as much as 3 cm. long, broadly ovate, green, cuspidate-acuminate,

5-nerved, incised-dentate near the apex and usually shallowly 3-lobate; stamens

about 30; pistillate sepals 5-6, lance-ovate, acuminate, multidentate, after anthesis

indurate and as much as 1.5 cm. long; ovary tomentulose; style column slender,

1 cm. long, very slightly dilated at the apex; cocci of the capsule 8 mm. long,

minutely puberulent; seeds 4 mm. in diameter, mottled with white and fuscous.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 83

The type locality has been stated as "Guatemala," but the

locality is the San Juan River, and presumably that of Nicaragua,

where Friedrichsthal is known to have collected.

Dalechampia guatemalensis Gandoger, Bull. Soc. Bot. France 66:

286. 1920. This species scarcely is properly published, being men-

tioned only in a brief key separating three American species proposed

by Gandoger, but not formerly described. It is said to be based

upon a Tuerckheim collection from Alta Verapaz. This collection

is probably Tuerckheim 7978, which is D. panamensis Pax & Hoffm.,

published in 1919.

Dalechampia heteromorpha Pax & Hoffm. Pflanzenreich IV.

147, xii: 26. 1919.

Moist or wet thickets, at or little above sea level ; Izabal (Quirigua,

Standley 23711, 24242). British Honduras; southern Mexico

(Veracruz); Costa Rica.

A twining herb, the stems densely pilose; leaves heteromorphous, simple or

3-foliolate, on petioles 3 cm. long or shorter; leaflets of the compound leaves sessile

or short-petiolulate, oblanceolate or lance-oblong to obliquely ovate-oblong, acumi-

nate, denticulate, puberulent on the veins, reticulate-veined beneath, chartaceous

or membranaceous; simple leaves oblong-ovate to broadly ovate, acute or obtuse,

deeply cordate at the base, short-petiolate; stipules triangular-lanceolate, reflexed,

3-4 mm. long; inflorescences short-pedunculate, the involucral bracts green, 2.5 cm.

long and wide, deltoid-ovate, acute, cordate at the base, denticulate, puberulent

on the veins, 5-nerved; pistillate sepals 10, pinnately parted, long-pilose, not

capitate-glandular; ovary pubescent; style column clavate, shallowly lobate at

the apex; capsule puberulent, 6 mm. high.

This has been reported from British Honduras as D. Schottii

Greenm., a species of the Yucatan Peninsula of Mexico that is to

be expected in British Honduras and Pet&i.

Dalechampia laevigata Standl. Field Mus. Bot. 4: 312. 1929.

Moist or wet thickets, 360 meters or less; Alta Verapaz (Pan-

cajche", Standley 70756). British Honduras; Atlantic coast of

Honduras.

A small herbaceous vine, the stems sparsely whitish-pubescent; stipules linear-

lanceolate, 4-6 mm. long; leaves simple, on petioles 2-6.5 cm. long, ovate to broadly

ovate or oblong-ovate, 7-16 cm. long, 3.5-10 cm. wide, obtuse and apiculate or

acute or acuminate, truncate or shallowly cordate at the base, palmately 3-5-

nerved, obsoletely and remotely serrulate or subentire, firm-membranaceous,

minutely puberulent above on the nerves, minutely pilosulous beneath on the

nerves, between them minutely and sparsely strigillose or almost glabrous; in-

volucral bracts green, membranaceous, broadly rounded-ovate, about 2 cm. long

84 FIELDIANA: BOTANY, VOLUME 24

and 3 cm. wide, obtuse, repand-denticulate, almost glabrous; pistillate sepals

about 7, pinnately parted, densely hispid, in fruit 1 cm. long or more; ovary

densely puberulent; style column filiform, slightly dilated at the apex; capsule

depressed-globose, deeply 3-lobate, minutely puberulent, 1 cm. broad; seeds

globose, 4 mm. in diameter, smooth, brownish.

Dalechampia molliuscula Blake, Contr. U. S. Nat. Herb. 24:

12. 1922.

Moist or wet thickets, 300 meters or less; endemic; Izabal (type

from Quebradas, S. F. Blake 7547; collected also at other localities

in the same general region).

Stems and petioles densely whitish-pilosulous or puberulent; leaves 3-f olio-

late, or sometimes 5-foliolate, on petioles 3 cm. long or shorter, the stipules subu-

late, 2.5-4 mm. long; leaflets abruptly contracted into a very short petiolule or

sessile, the lateral ones obliquely ovate or lanceolate, acute or acuminate, broadly

rounded at the base on the outer side, acute on the inner side, crenate-serrulate,

sparsely or densely pilosulous above, in age glabrate, beneath very densely velu-

tinous-pilosulous; terminal leaflet elliptic to oblong-lanceolate, 5-8 cm. long, 2-3.5

cm. wide, acute at the base; peduncles solitary in the leaf axils, about 1 cm. long;

involucral bracts green, orbicular-ovate, in fruit 2 cm. long and 2.5 cm. wide,

3-lobate almost to the middle, 5-nerved, densely pubescent like the leaves; pistil-

late sepals 12, in fruit 8 mm. long, linear, pinnate-laciniate, hispid; capsule 9 mm.

broad, sparsely setose, the style column slightly dilated at the apex, 7 mm. long;

seeds globose, 3 mm. long, dull grayish with 5 lighter-colored lines.

Dalechampia panamensis Pax & Hoffm. Pflanzenreich IV.

147, xii: 19. 1919. D. scandens var. trisecta Bonn. Smith, Bot.

Gaz. 13: 199. 1888 (type from San Juan Mixtan, Escuintla, J. D.

Smith 2079). Guachipi (fide Aguilar).

Moist or wet thickets or forest, sometimes in rather open, rocky

forest, 1,400 meters or less; Pete'n; Alta Verapaz; Izabal; Santa

Rosa; Escuintla; Guatemala; Retalhuleu. Southern Mexico (Oaxaca) ;

Honduras; Costa Rica; Panama.

A small or rather large, herbaceous vine, the stems slender, puberulent or

almost glabrous; leaves on petioles 3-5.5 cm. long, 3-foliolate; stipules linear, 6 mm.

long or less; leaflets short-petiolulate or sessile, 9-12 cm. long, 1.5-3.5 cm. wide,

the middle ones lanceolate or oblong-lanceolate, the lateral ones obliquely ovate-

oblong, all the leaflets acutely acuminate, acute at the base, or the lateral ones

broadly rounded on the outer side, subentire or remotely denticulate, membrana-

ceous, puberulent on the nerves, elsewhere glabrous or nearly so; inflorescences

axillary, short-pedunculate, the peduncles mostly 1.5 cm. long or less; involucral

bracts green, about 1.5 cm. long and slightly wider, in fruit accrescent and as much

as 3 cm. long and 4 cm. wide, 3-lobate, the lobes short, acute, glandular-dentate,

5-nerved; stamens about 24; pistillate sepals about 10, pinnatifid-laciniate, capitate-

glandular, in fruit 1 cm. long, whitish-hispid; ovary short-pubescent; style column

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 85

clavate, funnelform-dilated at the apex; capsule 8 mm. broad, appressed-pubescent;

seeds globose, 3 mm. in diameter, gray, mottled with whitish and blackish.

Dalechampia scandens L. Sp. PL 1054. 1753.

Moist or dry thickets, often in hedges or second growth, 1,800

meters or less, chiefly at low elevations; Zacapa; Jutiapa; Santa

Rosa; Escuintla; Guatemala; Sacatepe"quez; Retalhuleu; probably

in all the Pacific coast departments. Mexico; British Honduras to

Salvador and Panama; West Indies; South America; tropical Africa

and Asia.

A large or small vine, herbaceous or often somewhat woody below, the stems

usually densely fulvous-hirsute; leaves simple, on petioles 2-12 cm. long, 3-14 cm.

long and wide, membranaceous or chartaceous, deeply cordate at the base, 3-lobate

to the middle or more deeply, denticulate or subentire, usually densely short-

pilose on both surfaces, more densely so and paler beneath, the lobes oblong-el-

liptic or oblong-ovate, short-acuminate to obtuse and mucronate, the middle one

narrowed at the base; stipules broadly lanceolate or ovate, reflexed, 2-10 mm.

long; inflorescences axillary, the peduncles short or elongate; involucral bracts

pale green, 5-nerved, 1.5-3 cm. long and wide, cordate at the base, 3-lobate to

the middle or more deeply, glandular-denticulate or fimbriate-ciliate, densely

pilose; stamens 20-27; pistillate sepals 7-10, pinnate-lobate, capitate-glandular,

white-setose, in fruit 5-12 mm. long; ovary pubescent; style column cylindric or

clavate, somewhat dilated at the apex; capsule 7-10 mm. broad, pilose; seeds

globose, 2.5-4 mm. in diameter, mottled with whitish and brown, smooth.

Called "bejuco de pan" in Salvador; the Maya names in Yucatan

are recorded as "xmoolcoh" and "xmolcoh," signifying "puma foot."

The stiff hairs covering the inflorescences in this and other species

penetrate the skin and flesh readily, causing intense irritation.

Dalechampia Schippii Standl. Field Mus. Bot. 11: 133. 1932.

Open pine forest, at or little above sea level; British Honduras

(type from Sarawee, W. A. Schipp S-181).

A small twining vine, herbaceous or somewhat frutescent, the stems pilose

with short, spreading or somewhat reflexed hairs; stipules lanceolate, 3 mm.

long; leaves on petioles 3-6 mm. long, chartaceous, oblong or lance-oblong, 3.5-6

cm. long, 1.2-2.5 cm. wide, obtuse to short-acuminate, subcordate at the base,

remotely and obscurely serrulate or more coarsely serrate near the apex, densely

and softly short-pilose on both surfaces, much paler beneath, 3-nerved, the veins

prominent and closely reticulate beneath; peduncles axillary, solitary, usually

longer than the subtending leaves; involucral bracts pink veined with salmon,

ovate-rounded, 2-2.5 cm. long and wide, shallowly 3-lobate at the apex, sinuate-

denticulate, velutinous-pilosulous; capsule 5 mm. long, puberulent; seeds globose,

3 mm. in diameter.

Dalechampia spathulata (Scheidw.) Baill. Etud. Euphorb. 487.

1858. Cremophyllum spathulatum Scheidw. Bull. Acad. Brux. 9,

86 FIELDIANA: BOTANY, VOLUME 24

pt. 1: 23. 1842. D. Roezliana Muell. Arg. in DC. Prodr. 15, pt. 2:

1233. 1866. D. Roezliana var. viridis Muell. Arg. op. cit. 1234.

Dense wet mixed forest, 350 meters or less; Pete"n; Alta Verapaz;

Izabal. Southern Mexico; British Honduras; Atlantic coast of

Honduras; Costa Rica; Peru.

A simple shrub 30-90 cm. high, bearing few or numerous leaves, the young

branches sparsely pubescent, glabrate in age; leaves on petioles 2-10 mm. long,

obovate-spatulate or oblanceolate, 13-25 cm. long, 4.5-8 cm. wide, acuminate,

usually abruptly so, cuneate-attenuate to the base, the base itself narrow and

very obtuse or subcordate, glabrous or somewhat puberulent beneath along the

costa, penninerved, entire or sometimes dentate toward the apex; stipules 8-10

mm. long, ovate, acute, stiff, striate; inflorescences axillary, on short slender

peduncles, the involucral bracts 2.5-4 cm. long and almost as wide, ovate or

rounded-ovate, pink, dark red, or green, acute or acuminate, denticulate, 3-nerved;

stamens about 15; pistillate sepals 6, linear-lanceolate, 1 mm. long; ovary pubes-

cent; style column slender, 5-10 mm. long; capsule tridymous, 5-6 mm. long,

9 mm. broad, puberulent; seeds globose, muriculate, white.

In habit this is altogether unlike other Central American species.

It has been introduced into cultivation in hothouses of Europe, but

is not sufficiently decorative to merit cultivation.

Dalechampia tiliifolia Lam, Encycl. 2: 257. 1786.

Moist or wet thickets, 800 meters or less; reported from Pete"n;

Alta Verapaz; Izabal. Southern Mexico; British Honduras; Costa

Rica; Panama; South America.

A small or large vine, herbaceous or somewhat frutescent, the stems rather

stout, densely fulvous-pilose or subtomentose; leaves long-petiolate, simple,

membranaceous, 7-17 cm. long and wide, shallowly or rather deeply cordate at

the base, 3-lobate to the middle or lower, denticulate, pubescent above on the

veins or glabrate, usually densely pubescent beneath, the lobes elliptic or ovate-

oblong, acute or obtuse, the middle one narrowed at the base, the outer ones

angulate or rounded at the base; inflorescences terminal or on short few-leafed

branches, the involucral bracts creamy white, 3-6 cm. long and wide, broadly

ovate or rounded, rounded at the base, shallowly 3-dentate at the apex, sub-

entire, 7-9-costate, fulvous-tomentose; pistillate sepals about 12, pinnate-laciniate,

hispid, not capitate-glandular, in fruit 1.5 cm. long, plumose-hispid with stinging

hairs; ovary hispid; style column dilated and 3-lobate at the apex; capsule 11 mm.

broad, verruculose, hirsute; seeds globose, 5 mm. in diameter, greenish gray,

mottled with blackish.

DALEMBERTIA Baillon

Reference: F. Pax, Pflanzenreich IV. 146, v: 268-270. 1912.

Herbs or shrubs, sometimes at least with tuberous roots, glabrous or with

pubescence of simple hairs; leaves alternate, long-petiolate, entire, dentate, or

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 87

lobate, membranaceous, 3-7-nerved at the base; flowers monoecious, apetalous,

the spikes bisexual, pedunculate, the staminate flowers numerous, 3 inside each

bract, pedicellate; pistillate flowers few at the base of the spike or subsolitary on

a distinct branch, solitary within the bracts; disk none; staminate sepal 1, sub-

cochlea te at the apex; stamen 1, enclosed in the sepal, the anther longitudinally

dehiscent; ovary rudiment none; pistillate sepals 3, imbricate, 2-glandular at the

base; ovary 3-celled, the styles connate at the base into a column, recurved above,

simple; ovules solitary in each cell; capsule tridymous, the 2-valvate cocci separat-

ing from a persistent columella, the endocarp hard; seeds globose, not carunculate,

the testa crustaceous.

Four species in Mexico, one of them extending into Guatemala.

Dalembertia triangularis Muell. Arg. Linnaea 34: 218. 1865.

Rocky barranco slopes, 1,300-2,400 meters; Huehuetenango (re-

gions of Cuilco and San Ildefonso Ixtahuacan). Southern Mexico,

the exact locality unknown.

A slender shrub about 3 meters high, the branches terete, sordid-tomentose

at first, the flowering branchlets subumbellate at the apex of the stem; petioles

slender, 3 cm. long or less, villous-pilose; leaf blades triangular-ovate or triangular-

lanceolate, 4-7 cm. long, 2-4.5 cm. wide, narrowly long-acuminate, shallowly

cordate or truncate at the base, obscurely repand-dentate, usually shallowly and

obscurely 3-lobate or somewhat hastate, thinly or rather densely villous-pilose

or in age glabrate, somewhat paler beneath; staminate spikes dark red, conic-

ovoid, 1 cm. long, very densely many-flowered, tapering to the apex, sessile;

pistillate flowers axillary, solitary, the slender pedicels as much as 7 cm. long,

often curved; capsule deeply 3-lobate, 12 mm. broad, sparsely pubescent or

glabrous; seeds globose, grayish brown.

DITAXIS Vahl

Reference: F. Pax, Pflanzenreich IV. 147, vi: 51-77. 1912.

Shrubs or herbs, annual or perennial, the pubescence usually abundant, consist-

ing wholly or chiefly of malpighiaceous hairs (appressed and attached by the

middle); leaves alternate, short-petiolate, entire or dentate, membranaceous, 3-

nerved at the base, the stipules small; flowers monoecious or rarely dioecious,

petaliferous, the racemes mostly bisexual, pistillate below, staminate above, often

greatly abbreviated and congested; bracts small, 1-flowered, the flowers short-

pedicellate or subsessile, the pedicels often reflexed in fruit; staminate calyx ovoid

in bud and closed, valvately 5-parted in anthesis; petals 5, entire, equaling or longer

than the calyx, adnate to the androphore or almost free; disk of 5 glands; fertile

stamens 2-verticillate, usually 10, with or without filiform staminodia, the fila-

ments short, the anthers ovate, longitudinally dehiscent by introrse slits; ovary

rudiment none; pistillate sepals 5, the petals 4 or 5, entire, equaling or shorter than

the sepals; ovary subsessile, 3-celled, the styles free or connate at the base, 2-fid;

ovules solitary in each cell; capsule tridymous, the 2-valvate cocci separating from

a persistent columella; seeds not carunculate, subglobose, often reticulate or

foveolate; endosperm carnose, the cotyledons broad, flat.

88 FIELDIANA: BOTANY, VOLUME 24

Species about 45, in tropical and subtemperate America. At

least one other species is found in southern Central America.

Ditaxis guatemalensis (Muell. Arg.) Pax & Hoffm. Pflan-

zenreich IV. 147, vi: 59. 1912. Argyrothamnia guatemalensis Muell.

Arg. Linnaea 34: 145. 1865 (type collected in "Guatemala" by

Friedrichsthal).

Moist or dry thickets, open rocky places, sometimes in thin

forest, rarely a weed in cafetales, 1,500 meters or less, chiefly at

500 meters or lower; Alta Verapaz; Zacapa; Sacatepe"quez. Southern

and western Mexico; Salvador; Honduras.

Plants annual, perhaps sometimes more enduring and suffrutescent at the

base, erect, branched, 75 cm. high or less, the branches ascending, angulate, whitish-

sericeous when young; leaves grayish green, on very short petioles, ovate to lanceo-

late, 1.5-8.5 cm. long, acute or obtuse, acute to rounded at the base, inconspicu-

ously serrulate or subentire, densely whitish-strigose when young, in age glabrate,

often somewhat purplish, 3-nerved at the base; stipules narrow, 1 mm. long;

racemes very short, the flowers subglomerate, the rachis very short, bearing a

single pistillate flower and 3-8 staminate ones, the flowers sessile; staminate

sepals 5, lanceolate, acuminate, 4 mm. long, pilose on both surfaces; petals rhombic-

ovate to lanceolate, acute, equaling the calyx, pilose outside; stamens 10 and

2-seriate, accompanied by 3 small staminodia; pistillate sepals 5, linear-lanceolate,

6-7 mm. long, acuminate, the petals narrowly lanceolate, acuminate, shorter

than the sepals; ovary densely strigose, the styles connate almost to the middle,

pilose, bifid almost to the middle; capsule 4 mm. broad, depressed, densely pilose;

seeds globose, reticulate, 2-2.5 mm. in diameter.

Called "pericon" in Salvador. D. tinctoria (Millsp.) Pax &

Hoffm., which probably is synonymous, is called "tinta roja" in

Yucatan, where it is used to give a rose-pink dye. Many plants of

this genus impart a purple or reddish stain to the sheets of paper

between which they are dried.

DRYPETES Vahl

Reference: F. Pax, Pflanzenreich IV. 147, xv: 229-279. 1922.

Trees or shrubs; leaves alternate, short-petiolate, coriaceous, penninerved,

entire or dentate, generally unequal at the base, 2-stipulate; flowers dioecious,

apetalous, fasciculate in the leaf axils or at defoliate nodes; staminate sepals 4-5,

strongly imbricate, broad and concave, often unequal, ciliate; stamens 3-12 or

more, the filaments free, the anthers large, mostly introrse, the cells parallel;

disk central, plane or with an elevated margin, the margin often lobate or laciniate;

ovary rudiment small or none; pistillate calyx similar to that of the staminate

flower; hypogynous disk annular or rarely none; ovary 1-3-celled, the styles mostly

very short, the stigmas dilated, reniform-discoid; ovules geminate in each cell;

fruit drupaceous, globose or ovoid, subcarnose, in age coriaceous or subcrustaceous,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 89

the endocarp coriaceous to osseous, the fruit by abortion sometimes 1-seeded;

seeds not carunculate, the endosperm carnose, the cotyledons broad, flat.

Species about 140, in the tropics of both hemispheres. No others

are known from Central America.

Pedicels densely pubescent when young; fruit 2-2.5 cm. long; leaves mostly 14-20

cm. long D. Brownii.

Pedicels glabrous; fruit about 1 cm. long; leaves mostly 7-12 cm. long.

D. lateriflora.

Drypetes Brownii Standl. Trop. Woods 20: 20. 1929. Bulhop

(Pete*n, fide Lundell; Maya?).

Common in climax forest, 550 meters or less; Pete"n. British

Honduras (type from Hillbank, C. S. Brown 38).

A tree as much as 15 meters high with a trunk 15-20 cm. in diameter, glabrous

throughout except in the inflorescence; leaves coriaceous, on short stout petioles

3-6 mm. long, oblong to lance-oblong or elliptic-oblong, mostly 14-20 cm. long

and 5-8 cm. wide, abruptly acute or short-acuminate, with an obtuse or subacute

tip, contracted at the base and cuneate-decurrent, the costa and nerves prominent

on both surfaces, the lateral nerves about 10 pairs, irregular, ascending at an

acute angle, united remote from the margin; flowers fasciculate in the leaf axils,

the pedicels 3-5 mm. long, densely pubescent; sepals 2.5-3.5 mm. long, obtuse,

tomentulose, broadly oval, rounded at the apex; anthers short-exserted; ovary

densely tomentose; fruit obovoid-globose, 2-2.5 cm. long, very densely ochraceous-

tomentose, broadly rounded at the apex; seed oval, almost 1.5 cm. long, ochraceous.

Called "bullhoof" and "bullhoof macho" in British Honduras.

The wood is yellowish brown, often with reddish brown streaks;

hard, heavy, strong, somewhat brittle, medium-textured, fairly

straight-grained, not difficult to work, finishes smoothly, is not

durable. It is suitable for implements and tool handles.

Drypetes lateriflora (Swartz) Krug & Urban, Bot. Jahrb. 15:

357. 1892. Schaefferia lateriflora Swartz, Prodr. Veg. Ind. Occ. 329.

1787. D. crocea Poit. Me"m. Mus. Paris 1: 159. 1815. D. lateri-

flora var. guatemalensis Pax & Hoffm. Pflanzenreich IV. 147, xv:

255. 1922 (type from Teosinte, Santa Rosa, Heyde & Lux 4414).

Moist or wet, mixed forest, usually on limestone, 850 meters or

less; Pete*n; Santa Rosa. Southern Mexico; British Honduras;

Salvador; West Indies.

A tree, 10 meters high or less, with a trunk as much as 25 cm. in diameter, the

branchlets sparsely puberulent or glabrous; leaves coriaceous, on petioles 7-10

mm. long, lanceolate to lance-ovate or ovate, mostly 7-12 cm. long and 2-4 cm.

wide, acute or short-acuminate, acute to very obtuse at the base and usually

asymmetric, entire, glabrous, the lateral nerves about 8 pairs; flowers densely

90 FIELDIANA: BOTANY, VOLUME 24

fasciculate, the pedicels 3-5 mm. long, glabrous; staminate flowers 3 mm. broad,

the usually 4 sepals ovate, puberulent, ciliate; stamens generally 4, exserted; disk

and ovary pubescent, the ovary 2-celled; drupe subglobose, 1 cm. long, densely

tomentulose.

Called "mula" in Salvador.

EUPHORBIA L.

Annual or perennial herbs, shrubs, or small trees, with milky sap; leaves

alternate or opposite, or only the upper leaves opposite, sometimes verticillate,

usually membranaceous, entire or dentate, rarely lobate; inflorescences consisting

of one pistillate flower and several staminate ones enclosed in a calyx-like cup-

shaped involucre or cyathium; involucre campanulate, sometimes oblique, with

4-5 lobes and as many or fewer glands outside the lobes and alternate with them,

the glands often with petal-like, white or colored, spreading appendages; staminate

flower consisting of a single stamen, geniculate with the pedicel and soon deciduous,

usually without a calyx; anther cells generally globose; bractlets within the in-

volucre linear or setaceous, often lanate; pistillate flower with or without a minute

3-lobate calyx; ovary sessile at the apex of the pedicel, 3-celled, 3-ovulate; styles 3,

free or somewhat united, often bifid; capsule of three 2-valvate cocci, these separat-

ing at maturity from the central persistent axis (columella), ventrally dehiscent;

seeds with or without a caruncle, with a thin crustaceous testa; cotyledons broad,

flat.

One of the largest genera of plants, comprising about 1,500

species, widely dispersed in temperate and tropical regions, but most

numerous in the tropics. A very few species besides those listed

here are represented in southern Central America. The tropical

species, aside from a few isolated groups, have not received mono-

graphic attention in recent years, and the nomenclature in the genus

as a whole is in a confused state. Most of the Central American and

Mexican species are represented in the Herbarium of Chicago

Natural History Museum by authentic material, and there is little

or no uncertainty as to the proper application of most of the names

used here. It is probable, however, that some of them may be

antedated by names proposed for the same species in remote parts

of their range, particularly in South America or the West Indies.

A. Leaves all alternate, or the lowest ones, at least, alternate, and the upper

ones opposite or verticillate. (In E. scabrella, most of leaves present may be

opposite.)

Plants armed with spines; plants cultivated or rarely naturalized.

Inflorescences sessile or nearly so, greenish yellow, inconspicuous; spines

stout, scarcely 5 mm. long; branches deeply angulate E. neriifolia.

Inflorescences slender-pedunculate, showy, red ; spines long and slender, mostly

about 1.5 cm. long; branches obtusely if at all angulate. . .E. splendens.

Plants unarmed; native plants.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 91

Leaves very large, most of them 30 cm. long or longer, entire. Shrubs.

E. data.

Leaves much smaller or, if large, coarsely dentate.

Glands of the involucre without petal-like appendages, naked, sometimes

with crescent-like horns.

Plants tall shrubs, commonly 1-3 meters high; leaves of the inflorescence

large, usually bright red, and very conspicuous; leaves entire or

lobate E. pulcherrima.

Plants herbaceous, or rarely low shrubs, generally much less than a

meter high; leaves of the inflorescence green or, if colored, small

and not conspicuous.

Inflorescence usually umbel-like; stipules none; involucres in open

cymes, each involucre with 4 glands and entire or dentate lobes.

Leaves closely and minutely serrulate, about 7 mm. long, crowded.

E. trichotoma.

Leaves entire, much larger, usually spreading.

Plants annual; leaves rounded at the apex E. Peplus.

Plants perennial.

Leaves rounded or very obtuse at the apex . . . . E. Steyermarkii.

Leaves acute or acuminate E. orizabae.

Inflorescence not umbel-like; stipules gland-like; involucres in small

cymes, each involucre with a single gland or rarely 4 glands and

fimbriate lobes.

Glands of the involucre sessile.

Leaves mostly ovate or panduriform, variously dentate or lobate.

E. heterophylla.

Leaves linear or nearly so, almost or quite entire.

E. heterophylla var. graminifolia.

Glands of the involucre short-stipitate E. dentata.

Glands of the involucre with petal-like appendages, these often large,

white, and conspicuous, sometimes much reduced.

Plants annual.

Leaves of the inflorescence linear or lance-linear E. Francoana.

Leaves of the inflorescence orbicular or nearly so.

Stems densely glandular-pilose, at least above E. astroites.

Stems usually glabrous, sometimes sparsely short-pilose with

eglandular hairs E. ocymoidea.

Plants perennial.

Leaves very small, only 4-6 mm. long E. macropodoides.

Leaves much larger.

Capsule pubescent, very densely so when young.

Leaves acute at the base, fleshy E. lancifolia.

Leaves rounded or very obtuse at the base, thin . . E. Oerstediana.

Capsule glabrous from the first.

Floral leaves white, the white leaves sometimes very small but

usually conspicuous E. scabrella.

Floral leaves green.

Larger stems conspicuously angulate, flexuous or somewhat

zigzag E. ephedromorpha.

92 FIELDIANA: BOTANY, VOLUME 24

Larger stems terete, neither flexuous nor zigzag.

Involucres glabrous or pubescent with appressed hairs, the

lobes entire or nearly so E. graminea.

Involucres densely pilose with short spreading hairs, the

lobes incised-dentate E. guatemalensis.

A. Leaves all opposite or verticillate.

Plants trees or shrubs. Leaves large, entire, mostly verticillate.

Floral leaves white, conspicuous E. leucocephala.

Floral leaves green, inconspicuous.

Involucres glabrous E. Schlechtendalii.

Involucres pubescent.

Appendages of the involucre cleft into 2 narrow lobes; leaves oblong-

ovate, 1-3 cm. wide E. chiapensis.

Appendages of the involucre very shallowly or not at all lobate; leaves

orbicular or rounded-ovate.

Plants erect; leaves mostly 5-8 cm. wide E. cotinifolia.

Plants scandent or subscandent; leaves less than 2 cm. wide.

E. verapazensis.

Plants herbaceous, annual or perennial, low. Leaves often dentate, usually

small.

Leaves entire.

Leaf blades not oblique at the base; involucres mostly solitary in the forks

of the branches • E. chaculana.

Leaf blades conspicuously oblique at the base; involucres axillary.

Plants erect; leaves thick and succulent, mostly 8-12 mm. long.

E. buxifolia.

Plants prostrate; leaves relatively thin, smaller.

Stems densely and finely pubescent E. Seleri.

Stems glabrous E. serpens.

Leaves dentate or serrulate, sometimes only near the apex, but the teeth

there usually evident under a lens.

Plants glabrous throughout or usually so, the pubescence, if any, very

inconspicuous, never of long spreading hairs, the capsule glabrous.

Plants prostrate E. Blodgettii.

Plants normally erect.

Leaves linear or lance-linear or very narrowly oblong.

Leaves mostly 2-3 cm. long E. hyssopifolia.

Leaves mostly 1-1.5 cm. long E. cumbrae.

Leaves oblong to oval or broadly obovate.

Seeds black or gray, often with paler angles; capsule 2-2.2 mm. in

diameter E. brasiliensis.

Seeds dark red or reddish; capsule 1.5 mm. in diameter.

E. glomerifera.

Plants usually conspicuously pubescent on the stems and sometimes on

the leaves, the stems densely pubescent or sometimes only with sparse

long spreading hairs; capsule usually pubescent, often very densely so,

sometimes only on the angles, or rarely glabrous in plants having long

spreading hairs on the stems.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 93

Appendages of the involucre large, conspicuous, and petal-like, usually

reddish.

Plants erect, suffrutescent below E. ciichumatanensis.

Plants prostrate, herbaceous.

Appendages of the involucre densely pubescent £7. densiflora.

Appendages of the involucre glabrous.

Transverse ridges of the seeds very blunt, mostly 5, the sulci

between them almost closed and very inconspicuous . E. dioica.

Transverse ridges of the seeds rather sharp-edged, commonly 6 or

more, the sulci between them conspicuous and somewhat

open E. rutilis.

Appendages of the involucre much reduced and narrow, not petaloid.

Involucres numerous, in usually crowded or subglobose, pedunculate

cymes E. hirta.

Involucres in small axillary clusters, not in conspicuous pedunculate

cymes, the clusters sessile, or the flowers sometimes in lax open

cymes.

Stems prostrate, lying flat on the ground and often forming dense

mats.

Leaves densely pubescent.

Stems and leaves densely hirsute with very long, rather stiff,

spreading hairs E. villifera.

Stems and leaves velutinous-pilosulous or pilose with short,

not stiffly spreading hairs.

Leaves thinly pilose with long weak hairs; upper leaves acute.

E. velleriflora.

Leaves densely and finely velutinous-pubescent; upper leaves

rounded at the apex.

Involucre densely tomentulose; appendages subentire,

broader than the gland E. pantomalaca.

Involucre glabrous to puberulent; appendages crenulate,

about as wide as the gland E. prostrata.

Leaves glabrous.

Capsule densely pubescent all over with short, appressed or

incurved hairs E. thymifolia.

Capsule pilose only on the angles with rather long, spreading

hairs E. prostrata.

Stems erect or ascending, rarely procumbent, never lying flat on

the ground or forming mats.

Stems glabrous, or puberulent or pilose with very short hairs;

capsule densely pubescent with short hairs. Plants annual.

E. hypericifolia.

Stems long-hirsute throughout or at least below the nodes or at

the nodes; capsule glabrous or sparsely pilose with long hairs.

Leaves finely and evenly serrate from the apex almost to the

base, mostly oval, the upper leaves not or scarcely narrowed.

E. anychioides.

Leaves obscurely dentate, usually dentate only near the apex,

mostly triangular-ovate, the upper leaves much reduced.

94 FIELDIANA: BOTANY, VOLUME 24

Involucres glabrous outside or bearing only a few scattered

hairs; leaves very sparsely long-pilose or glabrate.

E. villifera.

Involucres densely long-pilose; leaves very densely pilose with

very long, spreading, white hairs, these almost as long

as the breadth of the leaves E. senilis.

Euphorbia anychioides Boiss. Icon. Euphorb. 12. 1856.

Golondrina; Pocul (Huehuetenango).

Open pine-oak forest, sometimes in rocky places, 1,500-2,500

meters; Jalapa; Chimaltenango; Huehuetenango. Mexico; Hon-

duras.

Plants usually perennial from a slender or thick, hard, woody root, usually

erect or ascending, the stems often numerous, slender, often much-branched,

sparsely or densely pilose with rather weak, long or short, more or less spreading

hairs; leaves short-petiolate, mostly oval or rounded-oval, chiefly 8-15 mm. long,

rounded at the apex, obliquely rounded at the base, rather thick, often tinged

with red, paler beneath, finely and almost regularly serrate from the apex nearly

to the base, thinly pilose on both surfaces with rather long, weak hairs or glabrate;

stipules minute, 2-3-parted with setaceous segments; involucres very small, mostly

solitary in the uppermost leaf axils, pedicellate, campanulate, reddish, glabrous,

the appendages of the glands transverse-oblong, pink or dark red, shallowly and

irregularly dentate; capsule glabrous or sparsely pilose with weak spreading hairs;

seeds ovoid-tetragonous, irregularly rugulose.

A characteristic and rather common plant of the dry, oak and

pine forests at middle and rather high elevations.

Euphorbia astroites Fisch. & Mey. Ind. Sem. Hort. Petrop.

10: 44. 1845.

Wet to dry thickets, 450-1,000 meters; Jutiapa; Santa Rosa;

Escuintla. Southern Mexico.

A slender erect annual about 30 cm. high, usually much-branched, the branches

green or greenish, usually glabrate below, densely short-pilose above with spread-

ing gland-tipped hairs; lower leaves alternate, the upper ones opposite or verticil-

late, on rather long, filiform petioles, orbicular or ovate-orbicular, sometimes

broader than long, very thin, entire, 1-2 cm. long or many of the leaves smaller,

rounded or very obtuse at the apex, truncate or broadly rounded at the base, thinly

pilose with very slender, subappressed hairs; stipules obsolete; involucres terminal,

pedicellate, green, hirtellous, the lobes ovate, 4-5-dentate, the glands concave,

the appendages 3-parted; seeds reddish brown, ellipsoid, tuberculate.

This is very closely related to E. ocymoidea, of which it may be

only a variety.

Euphorbia Blodgettii Engel. ex Hitchc. Kept. Mo. Bot. Gard.

4: 126. 1893.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 95

Sandy fields or usually on sea beaches; British Honduras; Florida;

southern Mexico; West Indies.

Plants annual, prostrate, glabrous, the stems 10-40 cm. long, often much-

branched and forming mats; leaves opposite, rather distant, somewhat fleshy,

oblong or oblong-oval, 4-14 mm. long, obtuse or rounded at the apex, rounded-

oblique at the base, conspicuously or obscurely serrulate, at least near the apex;

stipules triangular, acute, ciliate; involucres solitary in the upper leaf axils, cam-

panulate, short-pedicellate, glabrous outside, the lobes triangular; glands 4, short-

stipitate, orbicular to ovate, the appendages almost as broad as the glands, entire

or 2-3-crenate; capsule glabrous; seeds ovoid-tetragonous, pinkish or brownish,

the angles prominent, the sides slightly transverse-rugose.

Euphorbia brasiliensis Lam. Encycl. 2: 423. 1786. Golondrina;

Pie de paloma.

Moist or rather dry, open or brushy plains or hillsides, frequently

in rocky places, on sandbars, or a weed in waste or cultivated ground,

sometimes in pine-oak forest, 2,200 meters or less; Pete"n; Alta

Verapaz; Izabal; Zacapa; Chiquimula; Jutiapa; Santa Rosa; Es-

cuintla; Guatemala; Sacatepe"quez; Chimaltenango; Retalhuleu; San

Marcos; Huehuetenango; doubtless in all or most of the other

departments. Southern United States; Mexico; British Honduras

to Salvador and Panama; West Indies; South America.

An annual, usually erect and 15-50 cm. high, sometimes decumbent or almost

prostrate, the stems usually branched, often densely so, slender, glabrous or nearly

so; leaves short-petiolate or almost sessile, mostly oblong, sometimes lance-oblong,

ovate, or obovate, 1-3.5 cm. long, rounded to subacute at the apex, obliquely

rounded at the base, serrulate, especially toward the apex, glabrous or nearly so,

the leaves of the inflorescences smaller and narrower; stipules small, ciliate;

cymes axillary, lax, small, the involucres few, mixed with linear bracts, small,

campanulate, glabrous; glands of the involucre oblong or orbicular, the appendages

rather conspicuous, white, transversely elliptic; capsule glabrous, 2 mm. in diam-

eter; seeds trigonous, black or gray, with 2-3 transverse ridges.

Called "wild pisabed" and "chicken-weed hembra" in British

Honduras. This is one of the most common weedy plants of the

lowlands of Central America, abundant in many places. This

species often has been keyed from its close relatives as having

terminal inflorescences but, as Kostermans correctly states, the

inflorescences are axillary, just as in its near relatives.

Euphorbia buxifolia Lam. Encycl. 2: 421. 1788.

On or near sea beaches; British Honduras (Stann Creek, W. A.

Schipp S-58); Florida; Yucatan Peninsula of Mexico and on the

coastal islands; Panama; West Indies; northern Central America.

96 FIELDIANA: BOTANY, VOLUME 24

Plants erect, usually 50 cm. high or less, glabrous, succulent, herbaceous or

suffrutescent below, the branches usually numerous and suberect; leaves opposite,

short-petiolate, ovate to broadly oblong, 8-14 mm. long, subacute, obliquely

cordate at the base, entire, often involute; involucres mostly clustered at the ends

of the branches, campanulate, 1.5 mm. long, pedicellate, the glands transverse-

oblong, the appendages reduced to an obscure line; capsule 2 mm. broad, glabrous;

seeds white, ovoid-tetragonous, 1 mm. long, the angles very obtuse, irregularly

transverse-ridged.

The plant usually is confined to the vicinity of sea beaches and

apparently is rare in Central America.

Euphorbia canariensis L. Sp. PI. 450. 1753. An arborescent

plant, cactus-like in habit, a native of the Canary Islands. Several

individuals of what are believed to be this species are in cultivation

in La Aurora Park, Guatemala, where they attract much attention

because of their large size and fantastic form.

Euphorbia chaculana Bonn. Smith, Bot. Gaz. 27: 441. 1899.

Open grassy slopes, 1,600-2,700 meters; endemic; Huehuetenango

(type from Chacula, Seler 3128).

Plants perennial, erect or diffuse, 8-17 cm. high, arising from a radish-shaped

tuberous root 2 cm. long and 1 cm. thick, sparsely pubescent, the stems slender,

trichotomous, simple or much-branched; stipular glands none; leaves on very short

petioles, linear-oblong, obtuse at each end, 1-1.5 cm. long, 3-5 mm. wide, opposite,

cartilaginous-marginate, 1-nerved, sparsely pubescent, the petioles 1 mm. long;

pedicels 7-12 mm. long, usually equaling the subtending leaves, solitary; involucre

campanulate, glabrous, the appendages oval, entire or erose; capsule glabrous,

3 mm. high; seeds 2 mm. long, oval-tetragonous, rugose.

Euphorbia chiapensis Brandeg. Univ. Calif. Publ. Bot. 6: 54.

1914. Flor de pascua (Huehuetenango).

Brushy hillsides or in pine-oak forest, 1,500-2,000 meters;

Chiquimula; Jalapa; Huehuetenango. Chiapas; Oaxaca.

Plants 2 meters high or less, woody throughout or at least below, glabrous

except in the inflorescences; leaves opposite or verticillate, sometimes very numer-

ous at a node, thin, green above, pale beneath, the very slender petioles often

almost as long as the blades; leaf blades ovate to lance-ovate, 3-6 cm. long, obtuse,

rounded or broadly cuneate at the base, entire; involucres in terminal and axillary,

leafy cymes, these umbellate at the ends of the branches, the cymes glabrous or

somewhat pubescent; involucres 1 mm. high, pedicellate, puberulent, the 5 glands

oblong, the appendages cleft into 2 linear or narrowly oblong lobes; styles 2-

parted; capsule glabrous, 2.5 mm. broad; seeds subglobose, glaucous-gray, densely

and coarsely obtuse-tuberculate.

Euphorbia cotinifolia L. Amoen. Acad. 3: 112. 1756. Hierba

mala.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 97

Moist or dry, brushy hillsides, common or abundant in roadside

hedges in many regions, 1,200-2,400 meters; Alta Verapaz; Jalapa;

Santa Rosa; Guatemala; Sacatepe"quez; Chimaltenango; Solola;

Quiche" ; Quezaltenango; San Marcos. Southern Mexico; Nicaragua;

Costa Rica; Panama; northern South America.

A shrub or small tree, mostly 9 meters high or less, with a rounded crown, the

trunk often thick, covered with whitish, almost smooth bark, the branches glabrous

or nearly so; leaves deciduous, opposite, the petioles very slender, often equaling

the blades; leaf blades rounded-ovate or orbicular, entire, 5-14 cm. long, broadly

rounded at each end, glabrous or sparsely pubescent beneath, pale beneath;

involucres in dense terminal cymes, usually forming large leafy panicles, white

and rather showy, broadly campanulate, pubescent or glabrate; appendages of

the glands white or cream-colored, broader than long, crenate; young capsules

densely pubescent, at least on the angles.

Sometimes called "mala mujer" in Mexico, and "Barrabas" in

Costa Rica. This is a very common shrub or small tree in many

parts of the Guatemalan mountains, but it is almost confined to

hedgerows. It is especially abundant about Antigua, where it may

be found in almost every hedge. Although it is difficult to decide

whether the plant is really native in Guatemala, or comes from an-

other region, it is hard to understand why it should have been

introduced, since the abundant milky sap causes blisters and in-

flammation of the skin and is much dreaded. One gets the impres-

sion that the trees are left in hedges because it is dangerous to cut

them, the milk issuing rapidly and in large amounts whenever the

bark is cut. However, the trees are very solid and do make good

living fence posts. They shed their leaves during the dry season

and are quite bare for a long time. It is said that the foliage is eaten

only by goats, the sap causing severe blisters in the mouths of other

animals. It is stated that the plant is an important source of honey,

blooming when other flowers are scarce and providing a yellow honey

of good flavor. The milk is reported to have been used for criminal

poisoning by some Central American Indians, and in South America

it supplies one of the barbascos or fish poisons. The seeds are

reported to have drastic purgative properties, a property common

in many genera of this family.

Euphorbia cuchumatanensis Standl. & Steyerm. Field Mus.

Bot. 23: 169. 1944.

Known only from the type, Huehuetenango, Sierra de los Cu-

chumatanes, between Nenton and Las Palmas, 800-1,200 meters,

Steyermark 51646.

98 FIELDIANA: BOTANY, VOLUME 24

An erect shrub 20-30 cm. high, densely and often intricately branched, frutes-

cent below, herbaceous above, the root thick and ligneous, the older stems fusees-

cent, terete, the young ones pilose with spreading, white, almost straight hairs,

the internodes longer than the leaves; leaves small, on petioles scarcely more than

1 mm. long, opposite, pale when dry, rather thick, obliquely deltoid-ovate or ovate-

oval, 5-10 mm. long, 3.5-7 mm. wide, very obtuse or rounded at the apex, some-

times subacute, obliquely subcordate or rounded at the base, inconspicuously

undulate-denticulate near the apex or often almost entire, softly pilosulous on both

surfaces with spreading whitish hairs; involucres numerous, axillary or terminal,

solitary, short-pedunculate, densely pilosulous, broadly turbinate, acute at the

base, the appendages white, glabrous, suborbicular, 1 mm. in diameter, broadly

rounded at the apex, spreading, conspicuous; capsule 2 mm. broad, pubescent;

seeds plump, brownish ochraceous, somewhat thickened along the angles, the

sides almost smooth.

A well-marked species, noteworthy for the erect frutescent habit,

grayish, densely pubescent leaves, and the conspicuous white ap-

pendages of the involucre. It is to be expected in southern Mexico,

but we have been unable to place the plant with any of the very

numerous species described from Mexico.

Euphorbia cumbrae Boiss. Icon. Euphorb. 16. 1856.

Dry open slopes, about 250 meters; Zacapa (along Rio Motagua

west of Teculutan, Steyermark 29205). Mexico.

A very slender, erect, glabrous annual, mostly 25 cm. high or less, sparsely or

much-branched, the ultimate branches almost filiform and the lower ones scarcely

1 mm. thick; leaves opposite, on very short petioles, pale beneath, narrowly lance-

oblong, mostly 1-1.5 cm. long, obtuse or subacute, very oblique at the base,

obscurely serrulate near the apex; stipules 2-4-fid; involucres very small, subsessile

in the forks of the branches and in the uppermost leaf axils, campanulate, glabrous

within, the lobes subulate; glands suborbicular, concave, the appendages rather

conspicuous, entire, white or pink; styles short, 2-parted; capsule scarcely 2 mm.

broad, glabrous; seeds tetragonous, reddish, almost or quite smooth.

Euphorbia densiflora (Klotzsch & Garcke) Klotzsch in Peters,

Reise Mossamb. 94. 1862. Anisophyllum densiflorum Klotzsch &

Garcke, Tricocc. 28. 1859. Chamaesyce densiflora Millsp. Field

Mus. Bot. 2: 391. 1914. Golondrina.

Moist, open or brushy fields and hillsides, frequently in waste

or cultivated ground, sometimes on limestone, 2,050 meters or less;

Pete"n; El Progreso; Zacapa; Jutiapa; Santa Rosa; Guatemala;

Sacatepe"quez; Chimaltenango; Retalhuleu; Quiche"; Huehuetenango.

Mexico; Honduras to Panama.

Annual or perennial, usually much-branched from the base, prostrate and

often forming dense mats, the root often ligneous, sometimes fibrous, the stems

densely pilose or villous with rather long, spreading, whitish hairs, often dark

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 99

red, usually very densely leafy; stipules subulate-aristate; leaves almost sessile,

broadly oblong or ovate-oblong, mostly 1-1.5 cm. long, conspicuously oblique,

obtuse or rounded at the apex, very oblique at the base, denticulate, crisp-pilose

on both surfaces, usually densely so, often tinged or spotted with red; involucres

axillary, densely crowded near the ends of the branches and in the leaf axils,

campanulate, pilose, the lobes broad, fimbriate; glands large, flat, the appendages

whitish to dark red, pilose on both surfaces, erose-dentate; style 3-parted, the

branches bifid for half their length; capsule densely pubescent; seeds ovoid, slightly

tetragonous, pinkish, the sides convex, rugose.

From the state of New Mexico far southward into South America

the name "golondrina" is applied commonly to all or most species

of Euphorbia of the group Chamaesyce, particularly to the prostrate

plants. Perhaps on account of their conspicuous milky sap, they

are much used in Central America and Mexico in domestic medicine.

This custom seems to be widespread, and the senior author remem-

bers that in his childhood days the plants were sometimes used in

decoction for treating affections of children as far north as the state

of Missouri. This species has been reported from Guatemala as

E. adenoptera Bertol., a species of Florida and the Greater Antilles.

The species of this alliance were treated monographically by Mill-

spaugh (Field Mus. Bot. 2: 383. 1914).

Euphorbia dentata Michx. Fl. Bor. Amer. 2: 211. 1803.

Open or brushy plains and hillsides, frequently on sandbars

along streams or a weed in cultivated fields, 200-2,000 meters;

Zacapa; Chiquimula; Jalapa; Guatemala; Retalhuleu; Huehue-

tenango. Widely distributed in the United States and Mexico.

An erect, rather stout annual, mostly 40 cm. high or less, simple or branched,

the branches erect or ascending, villous or pilose; stipules glanduliform; lower leaves

alternate, the upper opposite, slender-petiolate, or those at the base of the inflores-

cence sessile or nearly so; leaf blades broadly ovate to lanceolate, 1-7 cm. long,

usually acute or acuminate, acute at the base, coarsely dentate, villous or pilose,

paler beneath; involucres clustered at the ends of the branches, oblong-campanu-

late, 3 mm. high, 3-5-lobate, bearing 1-4 yellowish short-stipitate glands, these

not appendaged; capsule glabrous or minutely pubescent, 4-5 mm. broad; seeds

obovoid or ovoid-globose, gray, inconspicuously 4-angulate, irregularly tuberculate.

Euphorbia dioica HBK. Nov. Gen. & Sp. 2: 53. 1817. Cha-

maesyce dioica Millsp. Field Mus. Bot. 2: 384. 1914. ?E. bryophylla

Bonn. Smith, Bot. Gaz. 56: 62. 1913 (type from Santa Rosa, Baja

Verapaz, 0. F. Cook 225). Golondrina; llama (Chimaltenango).

Brushy plains or hillsides, sometimes on sandbars along streams,

1,800 meters or less; Baja Verapaz(?); Zacapa; Chiquimula; Guate-

100 FIELDIANA: BOTANY, VOLUME 24

mala; Chimaltenango; Retalhuleu. Mexico; Honduras; Nicaragua;

Dominican Republic.

Plants usually or at least sometimes perennial, much-branched from the

base, prostrate and forming mats, mostly 30 cm. long or less, the stems densely

pilose with spreading hairs, usually very densely leafy; stipules subulate, pilose;

leaves opposite, on very short petioles, oblong or ovate-oblong, 3-8 mm. long,

obtuse or subacute, oblique-cordate at the base, denticulate, pilose on both surfaces

or glabrate, especially above; involucres usually very numerous and densely

crowded in the leaf axils, pink or red, pyriform-globose, the lobes minute, lanceo-

late; glands in 2 pairs, the appendages conspicuous and petal-like, unequal; capsule

pubescent; seeds triangular-obovoid, pinkish gray, transversely 4-sulcate.

Euphorbia elata Brandeg. Univ. Calif. Publ. Bot. 6: 55. 1914.

Dense, moist or wet, mixed forest, 1,200-2,000 meters; Izabal

(Cerro San Gil); Suchitepe'quez (Volcan de Santa Clara); Quezal-

tenango (Finca Pirineos). Chiapas.

A stout glabrous shrub 2.5-3.5 meters high, the branches thick, densely leafy

at the ends, grayish; leaves alternate, almost sessile or on somewhat elongate,

broad, marginate petioles, rather succulent when fresh, oblanceolate, mostly 30-

45 cm. long and 6-9 cm. wide, obtuse or abruptly subacute, long-attenuate to the

base, the lateral nerves obsolete or very obscure, deep green above, paler beneath;

cymes axillary, long-pedunculate, cymose-corymbiform, often much longer than

the leaves, the bracts large and foliaceous, soon deciduous, the thick pedicels 5-8

mm. long; involucres campanulate, 9 mm. broad, the lobes rounded, lacerate;

glands 5, orbicular; capsule glabrous, 1.5 cm. long; seeds subglobose, 6-7 mm. in

diameter, fuscous, smooth or nearly so.

Euphorbia ephedromorpha Bartlett, Proc. Amer. Acad. 43:

54. 1907; Bull. Torrey Club 38: 343. /. 1-3. 1911.

Brushy rocky hillsides, often in ravines, 120-650 meters; endemic;

Zacapa (type from Gualan, C. C. Deam 232; collected also at other

localities) ; Chiquimula.

Plants perennial, herbaceous or somewhat suffrutescent near the base, the

stems wing-angulate, pale green, flexuous, sometimes 1.5 meters long, leafless

during the dry season, glabrous, usually pendent from banks or subscandent

over low shrubs; leaves on long slender petioles, alternate, ovate or broadly ovate,

2-4.5 cm. long, obtuse or acute, usually rounded at the base, paler beneath, entire,

thinly pilose, especially beneath, or almost glabrous; stipular glands minute;

involucres cymose, the cymes lax, axillary on old branches, expanding before the

leaves, densely glandular-pilose; involucres narrowly obconic, 3 mm. long, pedicel-

late, the 5 lobes very short, flabelliform, laciniate; glands 5, the appendages white,

oblong or subspatulate, entire, rounded at the apex; capsule 2 mm. long; seeds

pinkish gray, ovoid, foveolate.

Euphorbia Francoana Boiss. Icon. Euphorb. 22. 1856.

Wet thickets or open places; Alta Verapaz (Panjach^, W. C.

Muenscher 12561). Southern Mexico.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 101

An erect annual, about 50 cm. high or lower, usually much-branched, the

branches ascending, slender, somewhat angulate, inconspicuously pubescent or

almost glabrous; leaves alternate or the upper ones opposite, very thin, green above,

paler beneath, on long, very slender petioles, broadly rhombic-ovate to lanceolate,

mostly 3-6 cm. long, obtuse to acute, cuneate to almost rounded at the base, thinly

pubescent with very slender hairs or glabrate, the leaves of the inflorescence small

and narrower, attenuate into a long slender setiform tip; stipules minute, gland-

like; involucres very small, slender-pedicellate, axillary, short-turbinate, the lobes

truncate, 3-4-dentate, the glands transverse-ovate, the appendages purplish,

biparted, the segments oblong, subacute, twice as long as the glands; capsule

sparsely short-pilose; seeds ovoid-rounded, subacute, coarsely pitted.

Apparently very rare in Guatemala, and perhaps introduced at

the single known locality, where the plant was found along the rail-

road tracks.

Euphorbia glomerifera (Millsp.) Wheeler, Contr. Gray Herb.

127: 78. 1939. Chamaesyce glomerifera Millsp. Field Mus. Bot. 2:

377. 1913. Partilla (Suchitep<§quez).

Open or brushy fields and hillsides, often a weed in waste or

cultivated ground, 700 meters or less; Pet£n; Zacapa; El Progreso

(type from El Rancho, W. A. Kellerman 8053); Escuintla; Suchi-

tepe"quez. Southern United States; Mexico; British Honduras to

Panama; West Indies; South America.

A slender erect annual, usually glabrous throughout or nearly so, 75 cm. high

or less, simple or usually branched, the stems often tinged with dark red or purple;

stipules small, ovate, dentate and ciliate; leaves on very short petioles, oblong to

oval or obovate, mostly 1.5-3 cm. long, rounded or very obtuse at the base, very

oblique at the base, serrate, pale beneath; inflorescences cymose, axillary, peduncu-

late, usually dense and many-flowered; involucres turbinate, very small, glabrous

outside, hirtellous within, the lobes lance-triangular, lacerate-dentate; glands small,

suborbicular, short-stipitate, the appendages white or reddish, orbicular or ovate;

capsule glabrous, 1.5 mm. broad; seeds ovoid-tetragonous, dark red or reddish,

the angles conspicuous, the sides irregularly rugose.

Called "chicken-weed" and "wild pisabed" (British Honduras);

"golondrina," "pela-tripa" (Veracruz). This is the plant to which

the name E. hypericifolia has been applied by most authors. Ac-

cording to Wheeler, E. hypericifolia L. of the Linnaean Herbarium

is really the plant that has been known commonly as E. lasiocarpa

Klotzsch, and on technicalities, at least, should replace the latter

(see Contr. Gray Herb. 127: 73 et seq. 1939). It would seem that

among the many names that have been published for these closely

related species an earlier one than that of Millspaugh could be found

for this common and widespread, weedy plant but Wheeler, ap-

parently, found no earlier one. The name E. glomerifera has at least

102 FIELDIANA: BOTANY, VOLUME 24

the virtue of clarity, for the type is an excellent and unmistakable

specimen. The Maya name in Yucatan is recorded as "toplanxiu."

It and related species are eaten commonly by horses and other stock.

Euphorbia graminea Jacq. Sel. Stirp. Amer. 151. 1763. Leche-

trezna (fide Aguilar) ; Escorpion-xiu (Pete*n).

Wet to dry thickets or open forest, often in pine-oak forest,

frequently a weed in waste or cultivated ground, 2,300 meters or

less; Pete"n; Alta Verapaz; El Progreso; Izabal; Zacapa; Jalapa;

Santa Rosa; Escuintla; Guatemala; Sacatepe*quez ; Chimaltenango;

Retalhuleu; Quezaltenango; Huehuetenango. Southern Mexico;

British Honduras to Costa Rica; northern South America.

A perennial herb, erect or decumbent or procumbent, usually 50 cm. high or

less, the stems slender and weak, green, usually more or less pilosulous with spread-

ing or crisped hairs, often glabrate, dichotomous; stipules minute, subulate; lower

leaves alternate, the upper ones opposite, on long slender petioles, very thin, entire,

broadly ovate to oblong or lanceolate, very variable in shape even on the same

plant, acute to rounded at the apex, cuneate to rounded at the base, mostly 1-7 cm.

long, thinly or rather densely pilosulous or crisp-pubescent, often almost glabrous;

involucres very small, pedicellate in the forks of the branches and in small lax

terminal cymes, appressed-pubescent or sometimes glabrous, the lobes ovate,

fimbriate; glands 2-4, transverse-ovate, the appendage obovate, white or yellowish

white, broader than the gland, entire or nearly so; capsule glabrous, small; seeds

tuberculate.

The Maya name in Yucatan is recorded as "onobcax." This is

presumably the plant that has been reported from Huehuetenango

as E. montereyana Millsp. In Guatemala this species is highly vari-

able in pubescence and leaf shape, and it may well be that the ample

material represents more than a single species, but it is not obvious

how the forms are to be separated. In this group a great number

of species have been described, based upon apparently inconse-

quential characters. The nomenclature is consequently involved,

also the taxonomy, and it is impossible to make a satisfactory dis-

position of the forms until this special group has been monographed

with great care. In some parts of Guatemala this plant is reported

to be poisonous to stock. Some of the specimens from Huehue-

tenango approach E. biformis Wats., a Mexican plant with tuberous

roots. Guatemalan material has been referred to the related E.

xalapensis HBK., which is very close to E. graminea, and rather

doubtfully distinct.

Euphorbia guatemalensis Standl. & Steyerm. Field Mus.

Bot. 23: 119. 1944. Lechetrezno bianco; Pie de nino.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 103

Moist or dry, open or brushy, often rocky hillsides or banks,

1,050-2,200 meters; endemic; Zacapa; Jalapa; Jutiapa; Santa Rosa;

Guatemala; Chimaltenango (type collected along road between

Chimaltenango and San Martin Jilotepeque, Standley 80907).

A perennial herb, the stems erect or often elongate and sprawling on the

ground or pendent from banks, sometimes subscandent and a meter long or

more, terete, not jointed, greenish, sparsely or densely pilose with soft weak hairs;

stipules minute, gland-like; leaves alternate, thin-membranaceous, entire, paler

beneath, on very slender petioles 1-2.5 cm. long, broadly ovate or rounded-

ovate, 2.5-4.5 cm. long, 1.5-4 cm. wide, obtuse or rounded at the apex, rounded

at the base and usually very narrowly subpeltate, thinly and laxly pilose or glabrate

on both surfaces; inflorescence consisting of a long lax leafy panicle bearing very

numerous small involucres, the slender branches sometimes flexuous; involucres

cymose, slender-pedicellate, 2 mm. long, turbinate-campanulate, densely pilose

with spreading hairs; glands 5, the appendage broadly oblong, strigillose outside,

subtruncate at the apex, laciniate-dentate or obtuse-dentate; capsule glabrous,

2.5 mm. broad; seeds pale, deeply foveolate.

Euphorbia heterophylla L. Sp. PL 453. 1753. Viborana;

Hierba mala de chibola; Copal (fide Morales); Flor de pascua de

monte; Echua (Izabal, fide Blake); Pastorcita.

Wet to dry, open or brushy plains and hillsides, a common weed

in cultivated ground, 1,700 meters or less, most common at low

elevations; Pete"n; Alta Verapaz; El Progreso; Izabal; Zacapa;

Jutiapa; Santa Rosa; Escuintla; Guatemala; Sacatepe"quez; Chimal-

tenango; Quiche" ; Suchitepe"quez ; Retalhuleu; Quezaltenango; San

Marcos; Huehuetenango; probably in most of the other departments.

Southern United States; Mexico; British Honduras to Salvador and

Panama; West Indies; South America.

An erect annual, sometimes a meter high but usually half as high or lower,

simple or sparsely branched, almost glabrous or somewhat pubescent; leaves

mostly alternate, the upper ones opposite, slender-petiolate, highly variable on the

same plant, mostly oblong-lanceolate to ovate or frequently panduriform, some-

times suborbicular, entire or irregularly dentate, paler beneath, those subtending

the inflorescence often red or pink at the base; involucres mostly crowded at the

ends of the branches, the lobes fimbriate; glands usually solitary, concave, not

appendaged; capsule 6 mm. broad, glabrous; seeds ovoid, pointed, grayish, 2 mm.

long, minutely and irregularly tuberculate in transverse lines.

Called "redhead" (British Honduras); "chilamatillo," "hierba

del duende" (Salvador); "hoboncax" (Yucatan, Maya). One of

the common weedy plants of the Central American lowlands, widely

distributed and common but seldom abundant locally, the plants

scattered and often solitary in a given locality. The milky latex

is used in domestic medicine. The plant is a highly variable one in

104 FIELDIANA: BOTANY, VOLUME 24

pubescence, leaf shape, and color of the floral leaves. Numerous

segregates have been proposed in this group, and some authors

recognize as species E. cyathophora Murr. and E. geniculata Ortega,

whose claims to specific status are not well supported. The most

conspicuous variant is the following:

Euphorbia heterophylla var. graminifolia (Michx.) Engelm.

in Torr. Bot. Mex. Bound. 190. 1859. E. graminifolia Michx. Fl.

Bor. Amer. 2: 210. 1803. Pascua; Pascua, miniatura.

Savannas or grassy hillsides, sometimes in thickets or on lime-

stone, or a weed in cultivated ground, 1,000 meters or less; Alta

Verapaz; Zacapa; Santa Rosa; Guatemala; Suchitepe"quez; Huehue-

tenango. British Honduras. Ranges with the typical form.

Differing from the typical variety only in the shape of the leaves, which are

all or mostly linear or linear-lanceolate.

Euphorbia hirta L. Sp. PI. 454. 1753. E. pilulifera of many

authors, perhaps not of Linnaeus. Golondrina; Coliflorcito (Jutiapa) ;

Hierba de paloma; Cocmachpin (Coban, Quecchi); Sabana de la

Virgen (fide Aguilar).

Wet to dry, open or brushy fields and hillsides, an abundant weed

of waste and cultivated ground, 2,500 meters or less, most abundant

at or little above sea level; Pete'n; Alta Verapaz; Izabal; Zacapa;

Chiquimula; El Progreso; Jutiapa; Santa Rosa; Escuintla; Guate-

mala; Sacatepe"quez; Suchitepe"quez; Retalhuleu; Quezaltenango; San

Marcos; Huehuetenango; probably in all the departments. Florida;

Mexico; British Honduras to Salvador and Panama; West Indies;

tropical South America.

An annual, generally erect or ascending, sometimes prostrate, usually densely

pubescent almost throughout with appressed or spreading, often yellowish and

multicellular hairs, the stems mostly 40 cm. long or less, generally dichotomous;

leaves opposite, on very short petioles, ovate to oblong-lanceolate, asymmetric,

1-3 cm. long, often blotched with dark red, acute, very oblique at the base, serrate

above the middle; stipules small, aristiform, pubescent; involucres small, very

numerous, mostly in dense head-like cymes, these pedunculate in the leaf axils;

lobes of the involucre triangular, densely long-ciliate, the glands 4, stipitate, the

appendages small and inconspicuous; capsule pubescent; seeds salmon-pink, ovoid-

tetragonous, with acute angles, the sides transverse-rugose.

The Maya name in Yucatan is "xanabmucuy" ; "tianguis"

(Yucatan); "chicken-weed" (British Honduras). One of the most

abundant weeds of the Central American lowlands, usually found

everywhere about dwellings, in dooryards, and in streets. Like

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 105

other weedy species of the genus, this varies considerably in habit,

leaf form, and other characters, but the species as a whole is well

marked and usually may be recognized at a glance. It has been

stated that this plant harbors the organisms causing the characteristic

tropical ulcers on children's legs, but this is denied by others who have

investigated the matter. The plant is one of the common domestic

remedies for a variety of minor ailments. The milk-like latex is

applied to cauterize granulated eyelids and as a remedy for itch

and other cutaneous affections.

Euphorbia hypericifolia L. Sp. PI. 1: 454. 1753. Euphorbia

lasiocarpa Klotzsch, Nov. Act. Acad. Nat. Cur. 19: Suppl. 1. 414.

1843. Golondrina; Golondrina blanca; Chibolita; Cangrejo bianco.

Dry to wet, open or brushy plains and hillsides, often a weed in

cultivated ground, frequently in rocky places, 1,300 meters or less;

Pete"n; Chiquimula; Jalapa; Jutiapa; Santa Rosa; Guatemala;

Quiche". Southern Mexico; Salvador to Panama; West Indies;

South America.

An erect annual, sometimes almost a meter high, usually lower, branched, the

branches slender, terete, usually pale, finely pubescent; leaves opposite, on very

short petioles, oblong or oval-oblong, pale, 1-4 cm. long, obtuse or rounded at

the apex, oblique at the base, finely and closely serrulate, short-pilose on both

surfaces with very slender, pale hairs; stipules triangular, 1 mm. long; involucres

in small dense terminal cymes, pubescent, narrowly campanulate, 1-1.5 mm.

long, the lobes triangular; appendages of the glands white, suborbicular, conspicu-

ous, entire or shallowly lobate; capsule almost 2 mm. long, usually very densely

pubescent or tomentulose; seeds ovoid-tetragonous, glaucous-brownish, shallowly

rugose.

According to Wheeler (Contr. Gray Herb. 127: 73. 1939), the

proper name for this species is E. hypericifolia L. That name has

been applied previously very generally to a common weedy plant of

tropical America, E. glomerifera.

Euphorbia hyssopifolia L. Syst. ed. 10. 1048. 1759. E. steno-

meres Blake, Contr. U. S. Nat. Herb. 24: 13. 1922 (type collected

between Los Amates and Izabal, Izabal, S. F. Blake 7776). Golon-

drina.

Open, wet or moist, grassy places, frequently in lowland pine

forest or savannas, 1,400 meters or less; usually at low elevations;

Pete"n; Izabal; Huehuetenango. Florida; Mexico; British Honduras;

Costa Rica; Panama; West Indies; South America.

An erect annual, 50 cm. high or usually lower, glabrous throughout or nearly

so, simple or sparsely branched, the stems often dark reddish, slender and stiff,

106 FIELDIANA: BOTANY, VOLUME 24

the ultimate branches almost filiform; stipules deltoid, lacerate-lobate; leaves

opposite, on very short petioles, linear to oblong-linear or lance-linear, mostly

2-3 cm. long or more, serrulate, at least toward the apex, paler beneath, acute

or obtuse, oblique at the base; cymes terminal, few-flowered; involucres short-

pedicellate, glabrous, the lobes triangular, mostly entire; glands minute, stipitate;

capsule glabrous; seeds ovoid-tetragonous, black or olivaceous, 1 mm. long, the

angles rounded, often white-edged, the sides transverse-rugose.

Euphorbia lancifolia Schlecht. Linnaea 7: 143. 1832. Ixbut;

Sapillo; I shut.

Wet to rather dry thickets, sometimes in pine forest or in open

fields, 600-1,900 meters; Alta Verapaz; Izabal; Santa Rosa; Guate-

mala; Sacatepe"quez; Quezaltenango; San Marcos; Quiche"; Hue-

huetenango. Southern Mexico; British Honduras; Honduras.

A perennial herb, somewhat fleshy and succulent, the stems terete, pale green-

ish, glabrous or nearly so, ascending or procumbent or prostrate, sometimes greatly

elongate, as much as 2 meters long or more, and subscandent; leaves alternate, on

very short, stout petioles, mostly rhombic-lanceolate and 5-9 cm. long, acute or

acuminate, acute at the base, entire, green and glabrous above, pale beneath and

inconspicuously and sparsely pilosulous or glabrate, the lateral nerves obsolete;

involucres in small, almost naked, terminal cymes, campanulate-turbinate,

4-lobate, glabrous, the lobes obovate, fimbriate, the glands transverse-ovate, the

appendage semiorbicular, crenulate, white or whitish.

This plant is well known in Guatemala, and even outside the

borders of the country, by the name "ixbut," which probably is of

Quecchi derivation. It is said to double the quantity of milk given

by cows that eat it. An infusion or decoction of it often is given

to nursing women to increase their flow of milk, and it is claimed

that it will cause the milk to flow after it has ceased normally, or

even in women who have not given birth to a child. Rather curiously,

it is claimed in Coban that the plants often cause the death of cattle

and horses eating them, and this may be the result of inherent

properties of the seeds. In Guatemala the plant often is planted as

a curiosity or for medicinal use in regions where it does not grow wild,

and we have seen it in cultivation in other countries of Central

America. It seems always to be assumed that the plant comes from

Coban or Verapaz, but as a matter of fact it is plentiful in many

parts of Guatemala.

Euphorbia leucocephala Lotsy, Bot. Gaz. 20: 350. pi. 24- 1895.

Pascuas; Flor de pascua; Flor de leche.

Moist or dry, open or brushy, often rocky plains or hillsides,

often on cliffs, frequently in pine-oak forest, 600-2,000 meters; El

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 107

Progreso; Zacapa; Chiquimula; Jalapa; Baja Verapaz (fide Clover);

Jutiapa; Guatemala; Chimaltenango; Quiche"; Huehuetenango (type

from Cuilco, W. C. Shannon 305). Southern Mexico; Salvador.

A slender erect shrub 1.5-4 meters high, the stems disarticulating at the nodes

when dry, green, angulate, glabrous; leaves mostly verticillate, on long, slender,

almost filiform petioles, obovate to oblong, elliptic-oblong, or oblong-lanceolate,

8 cm. long and 3.5 cm. wide or usually smaller, obtuse or rounded at the apex and

apiculate, obtuse or broadly cuneate at the base, often tinged with red, deciduous,

usually glabrous above, more or less pilose beneath; inflorescences thinly pilose,

cymose, the cymes rather dense, umbellate at the ends of the branches, the bracts

spatulate, white, obtuse, 1-1.5 cm. long; involucres campanulate, almost sessile,

pilose, the lobes transverse-oblong, ciliate; glands 5, transverse-oblong, the ap-

pendages white, petaloid, ovate, obtuse, entire, glabrous; capsule glabrous, 5 mm.

long.

Called "pascuita" in Salvador; "punopuno," "flor de nino,"

"flor de pascua" (Chiapas). The inflorescences are large and showy

because of the abundance of white bracts, which often are tinged

with pink and remain for a long time on the branches. Bunches of

the flowering branches are much used as decorations in houses and

churches. The bushes are conspicuous even from a great distance,

blooming, as they do, at a season when other flowers are scarce.

Euphorbia macropodoides Rob. & Greenm. Amer. Journ. Sci.

50: 164. 1895. Chapup (Huehuetenango).

Grassy, moist or wet, alpine meadows, 3,000-3,700 meters;

Totonicapan (Desconsuelo) ; Huehuetenango (Sierra de los Cuchu-

matanes). High mountains of Oaxaca.

A low, somewhat succulent perennial, mostly 5-8 cm. high, arising from a

rather large tuber, this fusiform or in age irregular, as much as 3.5 cm. in diameter;

stems usually several, erect, dichotomous, glabrous or nearly so; leaves mostly

alternate or the upper ones opposite, very small, slender-petiolate, orbicular to

short-oblong, 4-6 mm. long, rounded at the apex and the subequal base, regularly

but obscurely serrulate, sparsely pilosulous or glabrous; involucres solitary in

the forks of the branches or in the leaf axils, sparsely puberulent or almost wholly

glabrous, slender-pedicellate; glands 5, reniform, the appendages greenish, obtuse;

capsule glabrous; seeds ovoid, gray, 2 mm. long.

Evidently an alpine plant (it was collected in Oaxaca at 3,000

meters), not found in Guatemala by either of the authors.

Euphorbia neriifolia L. Sp. PI. 451. 1753. Tirabuzon; Tuno

de San Antonio.

Said to be a native of the East Indies; much planted for hedges

or for ornament in Guatemala, especially in the Oriente, particularly

108 FIELDIANA: BOTANY, VOLUME 24

common in Zacapa and Jutiapa, grown chiefly in the warmer regions,

but frequent about Guatemala and at other localities of middle

elevation.

A stout shrub commonly 1-2 meters high, with few or numerous branches,

these deeply 5-angulate, the angles bearing clusters of short sharp dark-colored

spines, leaves, and inflorescences; leaves thick and fleshy, deciduous in age, gla-

brous, alternate, on short marginate petioles, obovate-oblong, 8-12 cm. long or

larger, rounded at the apex, long-attenuate to the base, entire, the lateral nerves

obsolete; involucres small, greenish, in small dense sessile cymes, mostly borne

in the upper leaf axils, hemispheric, the lobes large, ovate, fimbriate on the margin;

glands transverse-ovate.

Called "tuna francesa" in Salvador. The sap is said to be

poisonous, but in Guatemala it is used as a remedy for hemorrhoids.

This is one of the commonest hedge plants in the Oriente of Guate-

mala. It is a very solid, cactus-like plant, and serves well for a

hedge, although it has no pretensions to beauty.

Euphorbia ocymoidea L. Sp. PI. 453. 1753. Flor de pascua

(Huehuetenango) .

Wet to dry, open or brushy hillsides or fields, often in pine-oak

forest, 600-2,000 meters; Alta Verapaz; Baja Verapaz; Zacapa;

Chiquimula; Jalapa; Jutiapa; Guatemala; Sacatepe"quez; Chimal-

tenango; Huehuetenango. Southern Mexico; British Honduras;

Honduras; Costa Rica.

A slender erect annual, 50 cm. high or usually lower, commonly much-

branched, the stems pale green, generally glabrous; leaves small, alternate or the

upper ones opposite, on very long, filiform petioles, very thin, entire, orbicular or

rounded-ovate, the larger ones about 1 cm. long, rounded at each end, paler be-

neath, entire, sparsely pubescent with weak slender hairs or almost glabrous, the

smaller leaves often broader than long; stipules obsolete; involucres small, pedicel-

late, solitary in the forks of the branches and clustered at the ends of the branch-

lets, usually pubescent outside, the lobes ovate, fimbriate-dentate; glands trans-

verse-ovate, the appendage 3-4-parted, with subulate divisions; capsule very small,

thinly hirtous; seeds small, ovoid, tuberculate.

A slender weak plant that withers as soon as the rains cease.

Central American material has been referred to E. adiantoides Lam.

and confused with E. astroites Fisch. & Mey., also with E. Armourii

Millsp., which is doubtfully distinct. The Maya name in Yucatan

is reported as "cambal-sac-chacah."

Euphorbia Oerstediana (Klotzsch & Garcke) Boiss. in DC.

Prodr. 15, pt. 2: 59. 1862. Poinsettia Oerstediana Klotzsch & Garcke,

Monatsber. Akad. Berlin 1859: 253. 1859. E. enalla Brandeg. Univ.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 109

Calif. Publ. Bot. 6: 54. 1914 (type from Chiapas). E. tetradenia

Brandeg. loc. cit. as syn. Ixbut de cobra; Mielilla.

Moist or wet thickets or open or dense forest, sometimes in pine

forest, 2,500 meters or less; Pete"n; Alta Verapaz; El Progreso;

Zacapa; Chiquimula; Santa Rosa; Escuintla; Guatemala; Sacatepe"-

quez; Chimaltenango; Solola; Quiche"; Quezaltenango; Huehuete-

nango. Nicaragua; Costa Rica; West Indies; northern South

America.

A perennial herb, erect or ascending, the stems often several from a single

root, simple or generally branched, mostly 50 cm. high or less, glabrous or pilose;

stipules obsolete; lower leaves alternate, the upper ones opposite, slender-petiolate,

thin, broadly ovate to oblong or lanceolate, 2-8 cm. long, acute or obtuse, rounded

or obtuse at the base, entire, glabrous or often rather densely pilose beneath, usually

ciliate; involucres chiefly in small, lax or dense, terminal cymes, slender-pedicel-

late, narrowly campanulate, pubescent, 2.5 mm. long, the lobes subtruncate,

fimbriate; glands with small, whitish or greenish appendages; capsule 3-4 mm. in

diameter, sparsely or densely pilosulous with weak pale hairs; seeds ovoid-globose,

coarsely punctate or tuberculate, grayish.

Euphorbia orizabae Boiss. in DC. Prodr. 15, pt. 2: 147. 1862.

Open or brushy hillsides, or often in open or rather dense forest

of oak, pine, or Abies, 1,900-3,000 meters or even higher; Chimal-

tenango (Volcan de Acatenango); Quiche"; Huehuetenango; Totoni-

capan; Quezaltenango; San Marcos. Mountains of southern Mexico.

An erect perennial herb, generally 75 cm. high or less, sometimes a stiff shrub

of 1.5 meters, glaucous-green, the stems slender, glabrous or usually minutely

puberulent or pilosulous above, simple or usually branched, umbellately branched

at the ends; leaves mostly alternate, those at the base of the inflorescences

opposite or verticillate, short-petiolate or sessile, narrowly lanceolate, mostly

2-6 cm. long, acute or subacute, attenuate to the base, entire, glabrous, thick,

the lateral nerves obsolete; involucres in umbellate leafy-bracteate cymes, cam-

panulate, glabrous outside, pubescent within, the lobes ovate, ciliate; glands

truncate, obscurely bicornute; capsule depressed-globose, glabrous; seeds obscurely

punctate.

This has been reported from Guatemala as E. campestris Cham.

& Schlecht., a Mexican species. It is widely scattered in the moun-

tains of the Occidente but seldom occurs in abundance, and is a

rather rare plant.

Euphorbia pantomalaca Standl. & Steyerm. Field Mus. Bot.

23: 120. 1944.

Known only from the type, Dept. Baja Verapaz, Sierra de las

Minas, opposite El Rancho (Dept. El Progreso), W. A. Kellerman

5175.

110 FIELDIANA: BOTANY, VOLUME 24

A perennial herb, arising from a thick woody root, the stems numerous, pros-

trate, densely leafy, dichotomous-branched, terete, densely and softly pilosulous,

the hairs short, spreading, white; leaves opposite, almost sessile, obliquely oval

or oval-ovate, 5-9 mm. long, rounded or very obtuse at the apex, very oblique

at the base, thick, obtusely denticulate near the apex or subentire, densely and

finely velutinous-pilosulous on both surfaces; involucres axillary, almost sessile,

campanulate, densely white-tomentulose, scarcely 1 mm. long; glands small,

transverse-oval, deep red, the appendage paler, subentire, broader than the gland;

young capsules short-exserted, densely white-tomentulose; styles hirtous.

Euphorbia Peplus L. Sp. PI. 456. 1753. E. chamaepeploides

Lotsy, Bot. Gaz. 20: 351. 1895 (type from San Martin Jilotepeque,

Heyde & Lux 3481).

A weed in moist shaded ground, mostly about dwellings, some-

times in flower beds or coffee plantations, 1,500-1,900 meters;

Sacatepe"quez; Chimaltenango; Huehuetenango. Native of Europe

and Asia, introduced and naturalized in various regions of America.

A glabrous annual, erect or ascending, 30 cm. high or less, often much-

branched, the branches erect, terete, greenish; leaves mostly alternate, those at

the base of the inflorescence verticillate, slender-petiolate, entire, broadly obovate

to oblong-obovate, 1-2 cm. long, rounded at the apex, acute at the base, thin;

bracts of the inflorescence broadly ovate or deltoid-ovate; involucres subsessile,

campanulate, less than 2 mm. long; glands 4, crescentic, not appendaged, pro-

longed into slender horns; capsule 2-3 mm. broad, glabrous; seeds oblong-ovoid,

whitish, subterete, with 1-4 transverse rows of conspicuous pits.

The plant is rare and local in Guatemala, but it may have been

imported from Spain long ago. It is strange that so well known a

plant should have been described as a new species by Lotsy, but it

merely illustrates the credulity of European and many American

botanists regarding supposed deep differences that separate tem-

perate and tropical floras, and the seriousness with which they

face international boundary lines.

Euphorbia prostrata Ait. Hort. Kew. 2: 139. 1789.

Open moist soil, 200 meters or less; Pete"n (Uaxactun). South-

eastern United States; Mexico; British Honduras; West Indies;

South America; Old World.

Plants annual or perhaps sometimes perennial, prostrate, much-branched,

forming mats, the stems slender, compressed, densely short-pilose, mostly 15 cm.

long or less; leaves opposite, subsessile, oblong or broadly obovate, 4-7 mm. long,

rounded or very obtuse at the apex, oblique at the base, serrulate, pubescent on

both surfaces or glabrate; stipules broadly deltoid, ciliate; involucres axillary,

glabrous to puberulent, very small, the lobes elongate-triangular, ciliate; glands 4,

transverse-oval, the appendages about as wide as the glands, crenulate; capsule

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 111

pilose on the angles with spreading white hairs, elsewhere glabrous; seeds pink,

ovoid-tetragonous, closely transverse-ridged.

Called "chicken-weed" in British Honduras. For this plant L. C.

Wheeler recently proposed (Rhodora 43: 265. 1941) to substitute

the name Euphorbia Chamaesyce L., but in error. For a later discus-

sion of the subject see Croizat, Bull. Torrey Club 72: 312-318. 1945.

Euphorbia pulcherrima Willd. ex Klotzsch, Allg. Gartenz. 2:

27. 1834. Poinsettia pulcherrima Graham, Edinb. New Phil. Journ.

20: 412. 1836. E. erithrophylla Bertol. Fl. Guat. 419. 1840 (type

collected in Guatemala by Velasquez). Flor de pascua; Pascua;

Guacamayo (Santa Rosa).

Native probably of southern Mexico and perhaps also in Guate-

mala; abundantly cultivated for ornament in Guatemala every-

where from middle or even rather high elevations down to the coasts;

observed as perhaps wild in moist or wet, wooded ravines in Jalapa,

Santa Rosa, and Huehuetenango. Grown for ornament in most

civilized tropical regions.

A shrub, commonly 1-4 meters high, with few stout branches, the branches

terete, glabrous; leaves alternate or the upper ones opposite or verticillate, on

long slender petioles, membranaceous, usually broadly ovate or panduriform but

frequently entire, mostly 12-20 cm. long, acute or acuminate, broadly cuneate

at the base, paler beneath, not dentate, glabrous or sometimes pubescent beneath;

leaves of the inflorescence large, brilliant red; involucres green and yellow, cymose-

corymbose, on stout pedicels, campanulate, hirtous within, the lobes broad and

short, laciniate; gland 1, not appendaged.

The floral leaves are usually of a brilliant red but sometimes

they are pale pink, or of a dull, dirty unattractive red shade, or rarely

white or pale yellow (forma lutea Standl., the "pascua amarilla" of

Salvador). This is one of the favorite and most showy ornamental

shrubs of Guatemala, planted around almost every dwelling from

the coasts far up into the mountains. The plants, of course, do

not thrive in regions where there is heavy frost, and it is said they

do not grow at Quezaltenango, but there may well be some in pro-

tected places. There are many bushes in the bleak mountains of

Huehuetenango, although there they are sometimes cut down by

frost. Many of the little white houses on the bare hills of this region

have growing beside them a single large bush of "flor de pascua,"

which in December is visible from a long distance, and blazes with

color so as to suggest a bonfire. Often no other ornamental plant

grows around these isolated dwellings, and one is tempted to wonder

if the plant may not have had formerly, or perhaps even now, some

112 FIELDIANA: BOTANY, VOLUME 24

religious significance. The name "flor de pascua" ("Christmas

flower") is given everywhere in Central America to this shrub

because it is most brilliant about Christmas time. Sometimes it

begins to "flower" in October in some regions, and may persist into

March, but after middle January the bushes usually look ragged and

far from attractive. The inflorescences are used everywhere for

decorations, especially on altars, but they are unsatisfactory for this

purpose, since they wilt rapidly unless placed in water. The plant

is well known in the United States under the name "Poinsettia," and

is grown in vast quantities in hothouses for sale in pots at Christmas

time, many of the plants so sold being merely unrooted branches

set in pots, in which condition they often remain fresh for a long

time if carefully watered. The name "Poinsettia," incidentally, a

former generic name for the plant, was given in honor of Joel R.

Poinsett, United States minister to Mexico, who "discovered" the

plant there in 1828, and introduced it into cultivation in the North.

Few people have had a more gorgeous plant species named for them.

It seems to be uncertain where the Poinsettia is native, and in

Central America it usually is confined to hedges and gardens. How-

ever, we have found it in several departments of Guatemala, as

indicated above, in localities where it appeared to be a native plant,

remote from any dwelling, and in places where it seemed improbable

that any dwelling had ever been. It was growing in rather dense

forest in quebradas, on very steep, rocky banks or cliffs, and this

may be its native habitat. It is of course possible but not very

probable that the plant had been introduced by birds, or that it

had been planted about some former shrine, but this seems unlikely.

The scarlet bracts are said to give a red coloring principle. In Guate-

mala the milk is sometimes used as an emetic, also as a remedy for

toothache, or as a depilatory, and poultices of the leaves are applied

to relieve body pains. The shrub grows readily from cuttings, and

in the tropics it thrives with little or no care, often covered with

drying clothes or used as a perch by the household chickens.

Euphorbia rutilis (Millsp.) Standl. & Steyerm. Field Mus.

Bot. 23: 120. 1944. Chamaesyce rutilis Millsp. Field Mus. Bot. 2:

385. 1914.

Known in Guatemala only from the type, growing in sand along

the railroad, Fiscal, Guatemala, 1,110 meters, C. C. Deam 6189.

Yucatan Peninsula of Mexico.

Plants annual or perhaps also perennial, prostrate and forming mats, the

stems 40 cm. long or less, much-branched, the stems compressed, densely white-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 113

pilose; leaves opposite, almost sessile, oblong to broadly ovate-oblong, 5-8 mm.

long, very obtuse or subacute, oblique at the base, serrate, glabrous or glabrate

above, pilose beneath; stipules subulate-setaceous, pilose; involucres axillary,

short-campanulate, pubescent outside, glabrous within, the lobes broadly deltoid,

ciliate; glands transverse-oval, concave, the appendages dark red or reddish,

glabrous, crenulate; ovary densely pubescent; capsule densely pubescent; seeds

ovoid, buff, the sides transverse-rugose, with about 4 sulci, these broad and open,

the ridges narrow and rather sharp-edged.

Closely related to E. dioica and rather doubtfully distinct.

Euphorbia scabrella Boiss. in DC. Prodr. 15, pt. 2: 55. 1862.

E. microappendiculata Lotsy, Bot. Gaz. 20: 349. 1895 (type from

Laguna de Ayarza, Santa Rosa, Heyde & Lux 3850). Flor de pascua

(Huehuetenango) ; Tuhonon (Coban, Quecchi) ; Quilete de leche; Ojo

de tdbano.

Wet to moist or dry, open or brushy, often rocky hillsides or

fields, sometimes pendent from cliffs, frequently in pine or oak

forest, 800-2,100 meters; Alta Verapaz; Baja Verapaz; El Progreso;

Zacapa; Chiquimula; Jalapa; Santa Rosa; Guatemala; Sacatepe"-

quez; Chimaltenango; Huehuetenango; Quezaltenango; San Marcos.

Southern Mexico.

A perennial herb, erect, decumbent, or sometimes procumbent, the stems a

meter long or usually half as long or less, branched, rough-hirtellous, pilosulous,

or glabrate, slender, green; lower leaves alternate, the upper ones opposite, slender-

petiolate, very variable in form, broadly ovate to linear, entire, mostly 7 cm. long

or shorter, obtuse or acute, rounded to acute at the base, thin, deep green above,

pale beneath, thinly pilose on both surfaces or glabrate; stipular glands small

and inconspicuous; bracts of the inflorescence small or rather large, spatulate or

obovate, pure white, conspicuous; involucres in small lax terminal cymes, pedicel-

late, turbinate, pubescent, glabrous in the throat, the lobes orbicular, fimbriate;

glands 2-4, concave, the appendage oblong; capsule glabrous, small; seeds tuber cu-

late.

A very common plant in many parts of Guatemala, often some-

what weedy, in general appearance like E. graminea, from which it

is not always sharply differentiated. It may well be that more than

one species is represented by the material we have referred here,

but, as remarked under E. graminea, the taxonomy of this group of

species must remain obscure and unsatisfactory until the group has

been monographed with great care. To E. scabrella probably belong

Guatemalan collections that have been referred toE. arenaria HBK.

Euphorbia Schlechtendalii Boiss. Cent. Euphorb. 18. 1860.

E. Friedrichsthalii Boiss. in DC. Prodr. 15, pt. 2: 61. 1862 (type

said to have been collected in Guatemala by Friedrichsthal). E. ad-

114 FIELDIANA: BOTANY, VOLUME 24

inophylla Dorm. Smith, Bot. Gaz. 47: 261. 1909 (type from Salvador).

Huailintdm (Huehuetenango).

Dry or moist, brushy, often rocky hillsides, 300-2,300 meters;

Alta Verapaz; El Progreso; Zacapa; Santa Rosa; Quezaltenango ;

Huehuetenango. Southern Mexico; Salvador to Costa Rica.

A shrub 1-2.5 meters high or larger, often much-branched, glabrous outside

the inflorescence or nearly so, the branches brown or pale brown, readily dis-

articulating at the nodes; leaves thin, entire, on long, almost filiform petioles,

verticillate, broadly ovate to suborbicular, 1-3 cm. long, rounded or very obtuse

at each end, paler beneath; involucres in small cymes, terminal and axillary, short-

pedicellate, hemispheric, glabrous, the lobes obovate, short-fimbriate; glands

broad, the appendage entire, whitish.

Called "carafio" and "pascuita" in Salvador; "boxchacah,"

"zacchacah" (Yucatan, Maya).

Euphorbia Seleri Donn. Smith, Bot. Gaz. 27: 441. 1899.

Known only from the type, Chacula, Huehuetenango, 1,600

meters, Seler 3127.

Perennial from a rather thick, hard root, the stems prostrate, dichotomously

much-branched, densely pubescent, 10-15 cm. long; stipules setiform, 1 mm. long;

leaves opposite, distichous, almost sessile, broadly ovate, 5-7 mm. long, thick

and somewhat fleshy, obtuse, entire, subcordate and unequal at the base, finely

pubescent when young, glabrate in age; involucres short-pedicellate, solitary in

the upper leaf axils, campanulate, sparsely pubescent or almost glabrous, the

lobes triangular-subulate, short-ciliate; glands 4, transverse-oval, yellowish, the

appendages white, obovate, entire; capsule almost glabrous; seeds reddish, ovoid,

rugulose.

Euphorbia senilis Standl. & Steyerm. Field Mus. Bot. 23: 120.

1944.

Known only from the type, Dept. Huehuetenango, dry slopes

between San Ildefonso Ixtahuacan and Cuilco, 1,350-1,600 meters,

Steyermark 50739.

A perennial herb, arising from a long perpendicular woody root, the stems

numerous, slender, flexuous, prostrate, laxly dichotomous, densely hirsute with

long spreading white hairs; stipules small, the segments almost setaceous, hirsute;

leaves opposite, almost sessile, broadly ovate or very broadly oval, often almost

quadrangular, chiefly 5-7 mm. long and almost as wide, very obtuse or broadly

rounded at the apex, obliquely subcordate at the base, thick, obsoletely undulate-

dentate or often entire, densely hirsute on both surfaces with very long, spread-

ing, white, slender but stiff hairs; involucres small, in the upper leaf axils, densely

white-hirsute, scarcely 1 mm. long; glands red-purple, transverse-oval, the ap-

pendages narrow, entire, red-purple; capsule almost 2 mm. broad, deeply sulcate,

glabrous or with a few long hairs; seeds obtusely tetragonous, glaucous, almost

smooth, the sides flat or slightly concave.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 115

Closely related to E. villifera, but very different in appearance

because of the unusually dense indument of long spreading stiff

hairs.

Euphorbia serpens HBK. Nov. Gen. & Sp. 2: 52. 1817.

Moist or wet fields or along lake margins, 500 meters or less;

Pete"n; Jutiapa; Retalhuleu. United States; Mexico; British Hon-

duras; northern South America.

Plants annual or sometimes perennial, the stems prostrate and forming mats,

much-branched, glabrous throughout, as much as 30 cm. long; leaves short-

petiolate, ovate-orbicular to broadly oblong, 2-7 mm. long, entire; stipules small,

lacerate; involucres short-pedicellate, solitary in the leaf axils, 1 mm. long, the

lobes deltoid; glands transverse-oblong, ochroleucous, the appendages little wider

than the glands, white, glabrous, usually crenate; capsule glabrous, 1.2 mm. long;

seeds ovoid, with rounded angles, white or brownish.

Euphorbia splendens Bojer ex Hook. Bot. Mag. pi. 2902.

1829. Corona de Cristo.

Native of Madagascar, but grown commonly for ornament or

as a curiosity in other regions of the earth, out of doors in the tropics

and in hothouses in the North; a rather common garden or pot plant

of Guatemala.

A shrub, often or usually scandent, glabrous or nearly so, the branches thick,

dark reddish brown, densely armed with long stout spines; leaves small, thin,

obovate or oblong-spatulate, entire, obtuse or rounded at the apex, narrowed

to the base, short-petiolate; cymes produced in the upper leaf axils, long-peduncu-

late, the bracts bright red, petal-like, showy; involucre campanulate, the lobes

broadly ovate, fimbriate; glands fleshy, red, transverse-ovate; seeds ovoid, tubercu-

late.

The name "crown-of-thorns" often is given to this plant in the

United States.

Euphorbia Steyermarkii Standl. Field Mus. Bot. 23: 121. 1944.

Contra-rotura.

On rocks in dense forest, 200-2,200 meters; endemic; Zacapa

(type collected near Zacapa, Standley 74673); Chiquimula; Hue-

huetenango.

An erect perennial herb about 30 cm. high, glabrous throughout, the stems

several, simple or sparsely branched, the branches subterete, about 5 mm. thick,

sometimes subangulate, rather sparsely leafy; leaves on stout petioles 1 cm. long

or less, somewhat fleshy, entire, deep green above, paler beneath, alternate, those

at the base of the inflorescence ternate, the blades obovate, rhombic-obovate,

or rounded-elliptic, 5-6.5 cm. long, 2.5-4.5 cm. wide, very obtuse or rounded at

116 FIELDIANA: BOTANY, VOLUME 24

the apex, cuneate at the base, penninerved, the lateral nerves obscure; stipules

obsolete; inflorescence terminal, umbelliform, small, with few branches, much

shorter than the leaves, the bracts green, cordate-orbicular or reniform, sessile,

8-14 mm. broad, broadly rounded at the apex; involucres few, sessile at the ends

of the branches among the leaves and hidden by them, glabrous, broadly campanu-

late, 2 mm. long, the lobes short, obovate, fimbriate-dentate; glands 5, not ap-

pendaged, suborbicular, spreading, entire; capsule long-exserted, the pedicel

stout, suberect.

Euphorbia thymifolia L. Sp. PI. 454. 1753. E. rubrosperma

Lotsy, Bot. Gaz. 20: 349. 1895 (type from Santa Rosa, Santa Rosa,

Heyde & Lux 4271). Golondrina.

Moist, grassy or brushy fields or hillsides, often a weed in waste

or cultivated ground, frequent on sandbars along streams, 1,000

meters or less; Pete"n; Izabal; Zacapa; Jalapa; Jutiapa; Santa Rosa;

Escuintla; Suchitepe"quez; Retalhuleu; Quezaltenango; San Marcos.

Mexico; British Honduras to Salvador and Panama; West Indies;

South America; Old World tropics.

A prostrate annual, the numerous stems much-branched and forming mats,

pubescent; leaves opposite, on very short petioles, oblong, 10 mm. long or less,

rounded or very obtuse at the apex, oblique at the base, serrulate, sparsely

pubescent or glabrous; stipules 1 mm. long, lacerate; involucres very small, mostly

solitary in the leaf axils, puberulent or glabrate, the lobes triangular, ciliate;

glands small, the appendages narrow or none, very inconspicuous; capsule puberu-

lent throughout with subappressed or incurved, short hairs, 1 mm. long; seeds

oblong, tetragonous, reddish, transverse-rugose.

Called "chicken-weed" in British Honduras. One of the com-

monest small weedy plants of the Central American lowlands.

Euphorbia trichotoma HBK. Nov. Gen. & Sp. 2: 60. 1817.

British Honduras (on beaches, Freshwater Cay, W. A. Schipp

929); southern Florida; Quintana Roo; Cozumel Island; Cuba.

A glabrous perennial, herbaceous or suffrutescent below, usually branched

from the base, the stems simple or branched, erect or ascending, widely dichoto-

mous above, very densely leafy; leaves sessile or nearly so, alternate, glaucescent,

cuneate or oblong, 5-12 mm. long, minutely denticulate; inflorescences umbellate

or often greatly reduced; involucres campanulate, 2 mm. high, sessile or nearly

so; glands obreniform, 1 mm. broad, yellow; capsule 4 mm. broad, depressed;

seeds subglobose, 1.5 mm. long, white or pale gray, smooth.

Euphorbia velleriflora (Klotzsch & Garcke) Boiss. in DC.

Prodr. 15, pt. 2: 40. 1862. Anisophyllum velleriflorum Klotzsch &

Garcke, Abh. Akad. Berlin 1859: 28. 1860.

Dry soil, about 300 meters; El Progreso (El Rancho, W. A.

Kellerman 7841). Southern Mexico.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 117

A prostrate annual, or perhaps sometimes perennial, the stems slender, much-

branched, terete, densely pilose with short soft weak hairs; leaves opposite, almost

sessile, ovate to ovate-oblong, 8 mm. long or shorter, the upper ones usually acute,

the lower obtuse, very oblique at the base, inconspicuously denticulate, thinly

pilose on both surfaces with slender weak hairs; stipules 3-6-fid, the segments

setaceous; involucres solitary in the leaf axils or somewhat crowded at or near

the ends of the branches, almost sessile, cylindric-turbinate, hirtellous outside

and in the throat, the lobes obovate, fimbriate; glands transverse-oblong, purplish,

the appendage equally broad or narrower; capsule white-hirsutulous; seeds oblong,

subacute, tetragonous, obscurely reticulate-rugose.

The single Guatemalan collection was determined by Millspaugh.

It agrees moderately well with type material of E. velleriflora, but

possibly it represents a distinct species.

Euphorbia verapazensis Standl. & Steyerm. Field Mus. Bot.

23: 121. 1944.

Known only from the type, Rio Chiacte", Alta Verapaz, 480

meters, C. L. Wilson 272.

A scandent shrub, glabrous outside the inflorescence, the stems articulate,

sub terete, greenish, with elongate internodes; stipules none; leaves opposite or

ternate, thin-membranaceous, entire, the filiform petioles 8-15 mm. long; leaf

blades orbicular or oval-orbicular, 15-20 mm. long, 10-17 mm. wide, broadly

rounded or broadly emarginate at the apex, rounded or very broadly cuneate

at the base, somewhat paler beneath, the lateral nerves evident but not elevated,

divergent at a wide angle; involucres arranged in small pedunculate cymes, these

scarcely longer than the petioles, ternate at the nodes; involucres short-pedicel-

late, sparsely strigillose, almost 2 mm. long, acute at the base, the pedicels and

branches of cymes thinly strigillose; glands transverse-oval, the appendage

whitish, glabrous, broader than the gland, obtusely denticulate, shallowly 2-lobate

at the apex.

Euphorbia villifera Scheele, Linnaea 22: 153. 1849. E. villifera

var. nuda Engelm. ex Boiss. in DC. Prodr. 15, pt. 2: 45. 1862.

E. siguatepequensis Standl. Field Mus. Bot. 4: 313. 1929 (type from

Siguatepeque, Honduras).

Open, often rocky fields or hillsides, most common in oak-pine

forest, frequently on limestone, generally in rather dry situations,

200-2,100 meters; El Progreso; Zacapa; Chiquimula; Jalapa; Jutiapa;

Quich^ ; Huehuetenango. Texas; Mexico; Honduras.

Annual or perennial, usually erect and much-branched, the stems often several

and interlaced, usually branched, thinly hispid throughout with long spreading

stiff hairs, or sometimes glabrate, the branches very slender, often almost filiform,

often dark red; leaves opposite, almost sessile, ovate or broadly ovate, or the upper

ones narrower and revolute, mostly 8 mm. long or less, obtuse to acute, cordate

and very oblique at the base, subentire but usually with several conspicuous coarse

118 FIELDIANA: BOTANY, VOLUME 24

teeth, often undulate; stipules lanceolate, 2-3-fid, hispidulous; involucres small,

solitary in the forks of the branches and in the upper leaf axils, pedicellate, turbi-

nate-campanulate, very sparsely hispidulous or often glabrous, the lobes ovate;

glands concave, the appendage white or reddish, entire, broader than the gland;

capsule glabrous or nearly so; seeds ovoid-tetragonous, obscurely rugulose.

A variable plant as represented in Guatemala, but probably all

the material referred here represents a single species. The typical

plant is abundantly or sparsely hispid; in var. nuda the plants are

glabrous or practically so. Both are represented in Guatemala.

Several species have been described in this group, most of which

probably will have to be reduced to synonymy under E. villifera,

or perhaps some earlier name that we have overlooked. The plants

in age usually turn dark red and are then rather conspicuous in the

grassy places where they grow. They are especially plentiful in

the dry, oak and pine forests of the mountains of Huehuetenango.

There have been reported from Guatemala two other species,

E. stictospora Engelm. (Rio Negro, Heyde & Lux 3483) and E. hirtula

Engelm. (Rio Negro, Heyde &Lux 3483c). Neither of these species

is to be expected in Guatemala, and we do not know what the

proper determination of these collections may be.

GARCIA Rohr

Reference: F. Pax, Pflanzenreich IV. 147, i: 14-15. 1910.

Trees; leaves alternate, without stipules, petiolate, firm-membranaceous,

penninerved, entire; flowers rather large, 1-3 at the ends of the branches, monoe-

cious, petaliferous; staminate calyx membranaceous, globose in bud, valvately

ruptured in anthesis into 2-3 segments; petals 8-12, narrow, sericeous, longer

than the calyx; glands of the disk free or connate at the very base, glabrous;

stamens numerous, inserted on a convex pilose receptacle, the filaments free;

pistillate calyx like the staminate, caducous, the petals fewer than in the staminate

flower; hypogynous disk deeply lobate; ovary 3-celled, the style very short, the

stigmas thick, reflexed, emarginate-bifid; ovules solitary in each cell; capsule

large, separating into 2-valvate cocci, the endocarp somewhat ligneous; seed

globose, not carunculate.

The genus consists of a single species.

Garcia nutans Rohr, Skrivt. Naturh. Selsk. Kjoebenhavn 2:

217. pi. 9. 1792.

In forest at base of bluff, 300-500 meters; Alta Verapaz (west

of Cubilgiiitz, Steyermark 44974}- Western and southern Mexico;

Salvador; Costa Rica; West Indies; Colombia; Venezuela.

A tree about 9 meters high, the branchlets, petioles, and lower leaf surface

finely puberulent at first, soon glabrate; leaves on slender petioles 2-5 cm. long,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 119

oblong or oblong-obovate, mostly 10-15 cm. long and 4-6 cm. wide, abruptly

acuminate or short-acuminate, acute or obtuse at the base; cymes 1-3-flowered,

on a very short peduncle, the pedicels almost twice as long as the calyx; flowers

nutant; calyx 5 mm. long, pubescent; petals lanceolate, acute, 8-12 mm. long,

whitish-sericeous, purplish within; filaments villous to the middle; ovary pubescent;

capsule glabrous, 3-coccous, 2.5 cm. broad; seeds 12 mm. in diameter.

Called "huevo de gato" in Salvador. The wood is pale brown,

of medium density, rather fine- textured, not difficult to work.

GYMNANTHES Swartz

Reference: F. Pax, Pflanzenreich IV. 147, v: 81-88. 1912.

Glabrous trees or shrubs; leaves alternate, short-petiolate, coriaceous or sub-

coriaceous, penninerved, entire or remotely glandular-denticulate, 2-stipulate;

flowers usually monoecious, apetalous, small, green, the spikes terminal or axillary,

solitary or fasciculate, commonly bisexual; staminate flowers 3-nate in the axils

of the bracts, these usually adnate to the rachis, the pistillate flowers few at the

base of the spike, or sometimes in separate spikes, solitary within the bracts;

disk none; staminate calyx obsolete or rudimentary, usually of 1-2 minute sepals;

stamens 2-6, generally 3, the filaments free or connate only at the base; ovary

rudiment none; pistillate flower naked or with 2-3 minute sepals; ovary 3-celled,

sessile in the calyx or borne on a gynophore; styles free or connate only at the base,

simple, recurved; ovules solitary in the cell; capsule tridymous, separating into

2-valvate cocci, smooth, the columella persistent; seeds subglobose, carunculate,

the testa crustaceous; endosperm carnose, the cotyledons broad, flat.

Species about 11, all in tropical America and mostly in the

West Indies. Only two are known from Central America.

Leaves with conspicuous glands on the margins at the base of the blade, mostly

12-20 cm. long, abruptly acute or acuminate G. guatemalensis.

Leaves without glands on the margins, mostly 5-14 cm. long and obtuse.

G. lucida.

Gymnanthes guatemalensis Standl. & Steyerm. Field Mus.

Bot. 23: 122. 1944. Cacho de venado.

Moist or wet, mixed forest, 1,300-1,500 meters; endemic; Quezal-

tenango (type collected in quebrada between Finca Pirineos and

Finca Soledad, lower southern slopes of Volcan de Santa Maria,

Steyermark 33501); San Marcos (Volcan de Tajumulco).

A large shrub or a small tree, 6 meters high or more, glabrous except in the

inflorescence, the branches slender, ochraceous, lenticellate, the young ones

angulate, green, rather densely leafy; leaves large, on stout petioles 7-12 mm.

long, coriaceous, oblong-oblanceolate or narrowly oblanceolate, 11-20 cm. long,

3.5-7 cm. wide, abruptly and shortly acute or acuminate with an obtuse tip,

gradually cuneate-attenuate to the acute base, appressed-crenate-serrate, the

margin bearing several large glands near the base of the blade, the veins prominu-

120 FIELDIANA: BOTANY, VOLUME 24

lous and laxly reticulate on both surfaces, the lateral nerves about 11 pairs;

staminate spikes 3.5-5 cm. long, floriferous almost to the base, many-flowered,

the rachis densely puberulent; bracts broadly ovate, sessile, obtuse, puberulent,

3-flowered, the flowers on rather long pedicels, the sepal broadly ovate; stamens

3-5 in the middle flower, 3 in the lateral ones.

Gymnanthes lucida Swartz, Prodr. Veg. Ind. Occ. 96. 1788.

Pij (Maya).

In thickets or forest on limestone, at or little above sea level;

Pete"n; Baja Verapaz (?; at 1,500 meters). Yucatan Peninsula of

Mexico; British Honduras; West Indies.

A glabrous shrub or tree, sometimes 10 meters high, the stiff slender branches

grayish; leaves on petioles 6-10 mm. long, coriaceous, lustrous, oblong-obovate

or oblong-oblanceolate, 5-14 cm. long, 2.5-4 cm. wide, obtuse or often narrowed

to a narrow obtuse tip, obtuse to attenuate at the base, often highly variable in

size and shape, subentire or somewhat crenate-serrate, often 2-glandular at the

base, paler beneath, the lateral nerves very slender, numerous, ascending at a

narrow angle, the veins prominent-reticulate; flower spikes about 3 cm. long,

bisexual, densely many-flowered, red-brown, becoming yellowish green; bracts

ovate, obtuse, puberulent, subcochleate-incurved, the staminate ones 3-flowered;

staminate sepal 1, broadly ovate; pistillate sepals scale-like, scarcely 1 mm. long;

stamens 3-5 in the middle flower, 2-3 in the lateral ones; ovary borne on a rather

long gynophore; pistillate pedicels in fruit 1.5-2 cm. long or more, the gynophore

as much as 1 cm. long; capsule about 7 mm. long and 9 mm. broad, globose-

tridymous, obscurely verruculose; seeds blackish brown, globose, 4-5 mm. in

diameter, the caruncle pale.

Called "false lignum-vitae" in British Honduras. The heart-

wood is variegated in olive and dark brown, sharply demarcated

from the white sap wood, hard, heavy, strong, its specific gravity

1.10-1.20, fine-textured, not difficult to work, takes a high polish,

is durable. Some use is made of the wood in the West Indies, but

probably not in Central America. It is sometimes imported into

the United States for use as backs of mirrors and brushes, walking

sticks, umbrella handles, and veneers for marquetry.

Hevea brasiliensis Muell. Arg., the rubber tree, native of the

Amazon basin, is commercially by far the most important plant of

the Euphorbiaceae, being the source of almost all rubber of com-

merce. It has been planted experimentally on a small scale in

Guatemala, particularly in the lowlands of Alta Verapaz and Izabal,

and in Suchitepe"quez; occasional trees may be found elsewhere,

grown more or less as curiosities. Most of the Hevea rubber of

commerce has been obtained from vast plantations in the East Indian

region.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 121

HIERONYMA Allemao

Trees or large shrubs, the incjument mostly of closely appressed scales, simple

hairs also sometimes present; leaves alternate, petiolate, penninerved, entire;

stipules small or large, early deciduous; flowers dioecious, apetalous, racemose

or paniculate, short-pedicellate, the bracts minute; inflorescences axillary, the

staminate large, the pistillate ones shorter and less branched; staminate calyx

campanulate, 3-6-dentate, the teeth slightly imbricate; glands of the disk 5,

alternate with the sepals; stamens 3-6, borne upon the sepals, the filaments free;

anthers exserted, the cells pendulous, longitudinally dehiscent; ovary rudiment

small; pistillate calyx like that of the staminate flower, the hypogynous disk

entire or lobulate; ovary generally 2-celled, the styles short, shallowly 2-parted,

reflexed; ovules geminate in each cell; fruit drupaceous, often by abortion 1-celled,

1-seeded, the exocarp thin, the endocarp osseous; seed not carunculate, the endo-

sperm carnose, the cotyledons broad, flat.

About 20 species, in tropical America. One other Central Ameri-

can species is found in Costa Rica. The generic name was written

originally Hyeronima.

Leaves almost glabrous beneath, with only a few scattered, very remote scales.

H. oblonga.

Leaves densely lepidote beneath, the scales close together and persistent.

Leaves of fertile branches mostly 6-16 cm. wide, the scales of the lower surface

not crowded and overlapping; lower lateral branches of the inflorescence

equaling the terminal branch H. alchorneoides.

Leaves of fertile branches mostly 3-5 cm. wide, the scales of the lower surface

crowded and overlapping; lateral branches of the inflorescence usually

few, much shorter than the terminal branch H. guatemalensis.

Hieronyma alchorneoides Allemao, Diss. in Trab. Veil. Rio

Janeiro, ill. 1848.

Moist or wet, mixed forest, 900 meters or less; Pete"n; Alta Vera-

paz; Izabal. British Honduras, along the Atlantic coast to Pana-

ma; tropical South America.

A tree, commonly 12-15 meters high, the trunk 30 cm. or more in diameter,

the young branches densely lepidote; leaves large, on petioles 2-7 cm. long, mem-

branaceous or subcoriaceous, broadly elliptic to elliptic-orbicular, mostly 9-20 cm.

long and 6-16 cm. wide, abruptly acute or short-acuminate, rounded at the base,

very sparsely lepidote and green above, densely lepidote beneath, brownish when

dried, often pilose along the nerves and costa, the lateral nerves 7-10 pairs;

stipules petiolate, about 1 cm. long or often much larger on juvenile branches,

ovate-rounded, subacute, soon deciduous; panicles densely lepidote, often 15 cm.

long, usually with numerous branches, the pistillate smaller than the staminate

and less branched, the flowers short-petiolate, cream-colored; calyx cupular,

3-5-dentate, pubescent; ovary lepidote; fruit ovoid-globose, 4-5 mm. long, black

at maturity.

Called "curtidor" in Honduras, the bark used there for tanning

leather. The trunks are often buttressed at the base. The strong

122 FIELDIANA: BOTANY, VOLUME 24

durable timber is used in some regions for fence posts, railway cross-

ties, bridge timbers, miscellaneous construction, and furniture. The

sapwood is pinkish white, the heartwood'very dark brown or reddish

brown, rather hard and heavy, its specific gravity 0.70-0.80; texture

medium or coarse, sometimes uneven; not easy to cut or split;

finishes smoothly.

Hieronyma guatemalensis Bonn. Smith, Bot. Gaz. 54: 241.

1912; Pax, Pflanzenreich IV. 147, xv:/. 5. 1922. Tern (fide Aguilar).

Dense or wet, mixed forest, or in forest of pine or Liquidambar,

sometimes in wooded swamps, 1,300-2,600 meters; Alta Verapaz

(type from Coban, Tuerckheim 423); Zacapa; Guatemala(?); Hue-

huetenango. Mountains of Costa Rica.

A tree, usually 7-12 meters high, the branches very densely lepidote; leaves

on petioles 2-4 cm. long, obovate or oblong-obovate, mostly 7-12 cm. long and

3-7 cm. wide, abruptly short-acuminate, cuneate at the base, green above, sparsely

lepidote and somewhat rough to the touch, very densely lepidote beneath with

crowded, often overlapping scales, the lateral nerves about 7 pairs; stipules small,

narrow, caducous; panicles densely lepidote, sparsely branched, short-pedunculate,

11 cm. long or shorter, the branches stout, the pedicels 1 mm. long; staminate calyx

2 mm. broad, 5-lobate to the middle, densely lepidote, the lobes acute.

Hieronyma oblonga (Tulasne) Muell. Arg. Linnaea 34: 66.

1865. Stilaginella oblonga Tulasne, Ann. Sci. Nat. III. 15: 248. 1851.

Dense wet mixed forest, at or little above sea level ; British Hon-

duras; southern Mexico; Costa Rica; tropical South America.

A tree, commonly about 10 meters high, the branchlets densely lepidote at

first, glabrate in age; leaves on slender petioles 1-2 cm. long, thick-membran-

aceous, obovate to oblong or obovate-oblong, mostly 6-15 cm. long and 3-6 cm.

wide, abruptly short-acuminate, acute at the base, green above, almost glabrous,

lustrous, somewhat paler beneath, sparsely and inconspicuously lepidote, appear-

ing glabrous, the lateral nerves 5-7 pairs; stipules cochleate, 6 mm. long; panicles

3-10 cm. long, with few branches, densely lepidote, the pedicels 1 mm. long or

less, the flowers whitish; staminate calyx shallowly 5-dentate, lepidote; ovary

glabrous; fruit red, 3-5 mm. long, ellipsoid, subacute at each end.

HIPPOMANEL. Manchineel

Reference: F. Pax, Pflanzenreich IV. 147, v: 261-263. 1912.

Glabrous trees with copious milky latex; leaves alternate, long-petiolate,

coriaceous, subentire or denticulate, penninerved, 2-stipulate; flowers monoecious,

apetalous, in terminal spikes, greenish, the rachis thick; bracts small, peltate-

glandular on each side at the base; staminate flowers 8-many in each bract, sub-

sessile, the pistillate flowers solitary within the lowest bracts of the spike or often

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 123

lacking; disk none; staminate calyx small, shallowly 2-3-fid, the lobes imbricate;

stamens 2, the filaments connate into a short column, the anthers longitudinally

dehiscent; ovary rudiment none; pistillate calyx small, deeply 3-fid; ovary 6-9-

celled, the style short, connate at the base, spreading above, simple; ovules solitary

in the cells; fruit drupaceous, large, apple-like, indehiscent, yellow or reddish, the

endocarp very thick, osseous, sinuous-rugose, 6-9-celled; seeds not carunculate,

the cotyledons broad, flat.

The genus consists of a single species.

Hippomane Mancinella L. Sp. PI. 1191. 1753. Manzanillo.

Frequent on sandy beaches of both coasts, growing only at the

inner edge of the beaches, and found nowhere else; Retalhuleu;

doubtless in all the coastal departments. Southern Florida; southern

and western Mexico; British Honduras and Salvador to Panama;

West Indies; northern South America (Colombia, Venezuela).

A glabrous tree, in some regions as much as 18 meters high with a trunk 90 cm.

in diameter, in Central America generally much smaller, the branches spreading,

forming a rounded crown, the bark thin, scaly, grayish or reddish brown; leaves

on petioles 3-5 cm. long, the petioles slender, 1-glandular at the apex; leaf blades

ovate or oval-ovate, 5-10 cm. long, abruptly acute or short-acuminate, obtuse or

rounded at the base or often emarginate, green and lustrous above, somewhat paler

beneath, rather conspicuously reticulate-veined but the venation not or scarcely

elevated; flower spikes 5-13 cm. long, stout, laxly flowered, usually bearing 1-2

pistillate flowers at the base; bracts very broadly ovate, entire; staminate calyx

scarcely 1 mm. long, the lobes denticulate; pistillate calyx 3 mm. long, the lobes

acuminate from an ovate base; fruit globose or depressed-globose, 2.5-3.5 cm.

broad, smooth.

The usual English name is "manchineel," a corruption of the

Spanish term manzanillo ("little apple"), and the specific name has

the same derivation. The tree is one of the most publicized ones of

Middle America, and has received much more notice than it deserves.

Growing upon the sea beaches, it naturally was about the first plant

to greet the Spaniards, who promptly gave it the name Manzanillo,

because the yellow and red fruit resembles a small apple. Always

seeking and expecting in the New World the plants of the Old,

they assumed they had found wild apples, and sampled the fruit,

sometimes, it is reported, with fatal results. The milky sap is

poisonous if taken internally, and upon contact with the skin causes

severe inflammation in some persons. Others are apparently im-

mune to the external effects of the juice. Smoke from the burning

wood is dangerous to the eyes. It is reported that the latex was

used by the Caribs for poisoning arrows. The early Spaniards had

such unfortunate experiences with the tree that they gave it a dark

reputation indeed, claiming that a person who even sat beneath a

124 FIELDIANA: BOTANY, VOLUME 24

tree would be blinded or would die, statements long ago proved

false. It is possible that people resting or passing beneath the trees

during rain might be harmed. The wood is lustrous yellowish brown

with markings of brown and black, and has long been used in some

parts of tropical America, mostly in the West Indies, for making

good furniture. It has a specific gravity of 0.60-0.68 and weighs

38-43 pounds per cubic foot; of fine and uniform texture; easy to

work and finishes smoothly; high in durability. The wood must be

handled with great care, even when dry.

HURAL. Sandbox

Reference: F. Pax, Pflanzenreich IV. 147, v: 271-274. 1912.

Large trees, glabrous, the trunk covered with short hard conic prickles and

resembling a grater; leaves alternate, long-petiolate, 2-stipulate, broad, more or

less cordate, crenate-serrate, penninerved; flowers rather large, monoecious, apetal-

ous, the staminate spikes terminal, pedunculate, oblong, thick, the flowers sessile,

solitary within the bracts; bracts membranaceous, at first enclosing the buds

and closed, irregularly ruptured at maturity; pistillate flowers solitary in the

uppermost leaf axils or at the base of the staminate spike, stout-pedicellate;

disk none; calyx eglandular, the staminate short-cupular, with a truncate margin,

denticulate; stamens usually numerous, the filaments continuous with the con-

nective, united into a thick column, the anthers longitudinally dehiscent; ovary

rudiment none; pistillate calyx coriaceous, broadly cupular, truncate, entire, laxly

surrounding the ovary; ovary 5-20-celled, the styles connate into a long column,

radiately spreading at the apex, simple; ovules solitary in each cell; capsule large,

somewhat depressed, the cocci verticillate, ligneous, elastically separating from

the persistent columella; seeds laterally compressed, large, not carunculate, the

testa crustaceous; endosperm carnose, the cotyledons orbicular, flat.

One other species is known, H. crepitans L., occurring in Central

America in Costa Rica and Panama, and found also in the West

Indies and tropical South America.

Hura polyandra Baill. Etud. Euphorb. 543. 1858. Jabillo;

Tetereta;Caquibach (Quecchi);Jaguili6 (Quiche*, fide Tejada) ; Haba;

Arbol del diablo.

Usually on forested plains or on dry, often rocky, thinly forested

hillsides, 1,200 meters or less; often found in cleared pastures, about

dwellings, or along roadsides; El Progreso; Zacapa; Chiquimula;

Jutiapa; Santa Rosa; Quiche"; Huehuetenango; doubtless in several

other departments, and probably in all the lowland areas. Mexico,

and extending southward, mostly along the Pacific slope, to Costa

Rica.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 125

A giant tree, often 30 meters high or more, with a trunk a meter or more in

diameter, the trunk usually straight and regular and free of branches usually for

a great height, the branches and trunk densely covered with short, very sharp,

hard prickles, the crown broad and spreading, the bark grayish or pale brownish;

leaves on petioles 8-15 cm. long, rounded-ovate, thick-membranaceous, 11-20 cm.

long and wide, rather deeply and narrowly cordate at the base, cuspidate-acuminate

or cordate-acuminate, coarsely crenate-serrate, the lateral nerves conspicuous

beneath, elevated, connected by faint transverse veins; staminate spikes long-

pedunculate, 5-8 cm. long, with a thick rachis, the flowers whitish or reddish;

stamen column as much as 15 mm. long, the anthers in 8-10 verticels; fruit de-

pressed-globose, 8-10 cm. broad, about 15-celled; seeds much compressed, brown,

3 cm. long.

The name used in Yucatan is "solimanche," a combination of

Spanish and Maya. This is one of the four or five largest trees of

Central America, reaching its best development on the plains or in

the foothills of the Pacific coast, where it often occurs in great abun-

dance, and in some regions is a dominant tree. This species has been

reported from Guatemala as H. crepitans L., but the latter is not

known to extend north of Costa Rica, although it might conceivably

extend to Guatemala. In H. crepitans the stamens are in only 2-3

verticels, while in H. polyandra the verticels are numerous, this dif-

ference being a very conspicuous one when flowers of the two species

are compared. The two species can not be separated by foliage

alone. The milk-like latex exuding copiously from the cut bark or

from the broken stems or leaves is caustic upon the skin, in some

persons producing blisters and intense inflammation. This latex,

mixed with meal or sand, is much used in Mexico as a barbasco or

fish poison, and the same use may be made of it along the Pacific

coast of Central America. The seeds are known to be dangerously

poisonous, but in Guatemala the pulverized kernels sometimes are

administered in small doses to human beings as a purgative and to

expel intestinal parasites. It is unnecessary to state that such use is

dangerous. They are used in the Oriente of Guatemala to poison

noxious animals. In Honduras it is claimed that a person who sleeps

under one of the trees amanece muerto, or that at the least the whole

body will be covered with blisters. The first portion of this statement

is undoubtedly untrue, and there probably is little basis for the

second part. The name "jabillo" or "jabilla" is a derivative of

Haba, the Spanish name for the horse bean, Vicia Faba, given be-

cause there is some resemblance between the seeds of the two plants.

An aldea of the Department of Guatemala bears the name "Javilla."

The dry leaves of this tree, which often cover the ground during the

dry season, are eaten by cattle.

126 FIELDIANA: BOTANY, VOLUME 24

The fruit of the sandbox is curious in appearance and structure,

somewhat resembling a diminutive pumpkin or cantaloupe, with the

cocci arranged like the sections of an orange. When thoroughly dry,

the fruit explodes elastically with great force and a loud report,

scattering the seeds to some distance. The force of the explosion

is great enough to burst small boxes in which the fruits may have

been stored, or to cause wounds if they strike any part of the body.

The dry fruits were sometimes used in early days in the British

West Indies as containers for fine sand with which letters were

dried or "blotted," hence the customary name of sandbox. The

seeds apparently are a favorite food of the brilliantly colored macaws

of Central America. Sandbox wood varies in color from lustrous

creamy white to yellowish brown or olive-gray; it is light and soft,

medium-textured, more or less woolly, the specific gravity 0.36-0.44;

weight 23-27 pounds per cubic foot; easy to cut, takes stains and

glues well. It is used in Central America to some extent for con-

struction.

JATROPHA L.

Herbs, shrubs, or trees; leaves alternate, petiolate or sessile, simple and

entire or often palmate-lobate, glabrous or pubescent, often with glands of various

kinds; stipules small and inconspicuous or setaceously dissected, sometimes in-

durate and spinose; flowers small, usually monoecious, petaliferous, in dichot-

omous cymes; sepals of the staminate flowers more or less connate at the base,

imbricate; petals 5, contorted-imbricate, free; disk entire or 5-glandular; stamens

usually 8-10, in 2 or more verticels, monadelphous, the outer 5 epipetalous;

staminodia present and filiform or none; calyx and corolla of the pistillate flower

like those of the staminate flower; ovary 2-3-celled, rarely 4-5-celled; styles

connate at the base, entire or 2-fid; ovules solitary in the cells; fruit capsular, the

seeds carunculate.

Species about 115, in tropical regions of both hemispheres.

Probably all the Central American ones are included in the following

enumeration.

Leaves peltate; stipules indurate and spine-like J. podagrica.

Leaves not peltate; stipules not spinose.

Petioles bearing numerous long-stalked glands; plants herbaceous; leaves lobate

to the middle or more deeply J. gossypiifolia.

Petioles without stipitate glands; plants shrubs or trees; leaves entire or only

very shallowly lobate.

Leaf blades rounded at the base; corolla bright red and showy. . .J. hastata.

Leaf blades cordate at the base; corolla whitish, small and inconspicuous.

Leaves entire J. Gaumeri.

Leaves shallowly angulate-lobate J. Curcas.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 127

Jatropha Curcas L. Sp. PI. 1006. 1753. Pindn; Tempate;

Yupur, Tempacte (fide Tejada), Sakilte (Quecchi).

Moist or dry thickets on plains and hillsides, most plentiful in

hedges and often planted for living fence posts, 1,500 meters or lower,

most common at low elevations; Pete"n; Alta Verapaz; El Progreso;

Izabal; Zacapa; Jalapa; Jutiapa; Santa Rosa; Escuintla; Sacatepe"-

quez; Suchitepe"quez; Retalhuleu; San Marcos; Huehuetenango;

probably in all or most of the other departments. Mexico; British

Honduras to Salvador and Panama; West Indies; South America;

cultivated and sometimes naturalized in the Old World tropics.

A shrub or small tree, sometimes 8 meters high but usually lower, the bark

pale and almost smooth; leaves long-petiolate, the slender petioles about as long

as the blades; leaf blades rounded-ovate in outline, mostly 7-16 cm. long and of

about the same width, openly cordate at the base or sometimes truncate, shallowly

3-5-lobate or angulate, not dentate, palmately 5-7-nerved from the base, almost

glabrous but more or less pilose beneath on the nerves, at least near the base of

the blade; cymes small, dense, long-pedunculate, many-flowered, the bracts

lanceolate or linear; sepals ovate-elliptic, 4 mm. long, glabrous; petals whitish,

oblong-obovate, almost free, densely pilose within, in the staminate flower twice

as long as the sepals, in the pistillate flower almost equaling the sepals; stamens 8,

the outer filaments free, the inner ones connate; ovary glabrous; capsule 2.5-4 cm.

long, 2-3-celled, ellipsoid; seeds about 2 cm. long and 1 cm. broad, pale, oblong-

ellipsoid, with conspicuous black lines.

Maya names reported from Yucatan are "xcacalche" and"sicilte."

This is one of the best-known and most common plants of the low-

lands of Central America where it is planted abundantly for hedges

and living fence posts, principally because it is not eaten by stock

of any kind. The shrub may not be native in Guatemala, since it

is found principally in hedges, but if not, doubtless it has been in

cultivation for a long time. In Mexico it has long been used as a

host plant for certain lac-producing scale insects known by the name

Axi or Axin, and it is quite probable that the same use may have

been made of the shrub in Guatemala. The lac thus produced is

highly esteemed as varnish for finishing guitars and other articles

of wood. In Guatemala an infusion of the leaves is used commonly

by some of the Indians for setting the dyes of cotton and perhaps

other textiles. The milky sap is applied commonly to wounds or

sores, to hasten healing, and it is placed in cavities in teeth to relieve

toothache. About Coban the heated leaves sometimes are applied

to the breasts of nursing women in the belief that this increases the

flow of milk. The properties of the large and rather well-flavored

seeds are well known in Guatemala and other parts of Central

America. When the seeds are pressed they yield a large amount

128 FIELDIANA: BOTANY, VOLUME 24

of clear oil that is used locally for making soap and for illumination.

This oil, or the whole seed, has drastic purgative properties, and if

the seeds are eaten the results sometimes are dangerous or even

fatal, at least in the case of small children. Some years ago the United

States Department of Agriculture not very wisely attempted to

introduce the plant into cultivation in Florida, as it had been found

that the kernels of the seeds after very thorough roasting are of

good flavor and may be eaten without harm. However, some persons

were made very sick by eating only partially roasted seeds. Since

the seeds are dangerous when taken as food, their use for this purpose

is to be avoided. The shrub sheds its leaves during the dry months

and has little to recommend it as a hedge plant.

Jatropha Gaumeri Greenm. Field Mus. Bot. 2: 256. 1907.

In thickets, at or little above sea level; Pete"n (Carmelita, F. E.

Egler 42-270); Zacapa(?). Yucatan Peninsula of Mexico; British

Honduras.

A shrub or small tree, sometimes 8 meters high with a trunk 30 cm. in diameter,

the branchlets thick; leaves long-petiolate, the slender petioles mostly 4-14 cm.

long, glabrous; leaf blades firm-membranaceous, broadly ovate or rounded-ovate,

5-18 cm. long and 5-15 cm. wide, acute or abruptly acuminate, deeply cordate

to subtruncate at the base, palmately 7-nerved, entire, glabrous above, more or

less pubescent beneath along the nerves near the base of the blade; inflorescences

cymose, pedunculate, many-flowered, glabrous or nearly so, the bracts triangular-

ovate, acute, the flowers sessile, whitish or cream-colored; calyx 2-3 mm. long,

glabrous, 5-lobate; corolla 6-7 mm. long, tubular for two-thirds its length, glabrous

outside, the 5 lobes erect or slightly spreading, rounded at the apex; stamens 8,

included; capsule oblong-globose, 15-18 mm. long and broad; seeds oblong, 13

mm. long, 11 mm. broad, slightly roughened.

Known in British Honduras by the names "wild physic nut,"

"pinon," and "pomolche"" (Maya). A single sterile collection from

Zacapa has leaves like those of this species, but much smaller than

in Yucatan specimens. They are densely and softly pilose beneath.

It is quite possible that the Zacapa plant is a distinct and undescribed

species.

Jatropha gossypiifolia L. Sp. PI. 1006. 1753.

Moist or dry fields or thickets, frequently on sand or gravel along

streams, 200-400 meters; Zacapa; Chiquimula; in the Jardin

Botanico, Guatemala, where planted or perhaps an accidental weed.

Southern Mexico; Honduras and Salvador to Panama; West Indies;

South America; western Africa (introduced?).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 129

Plants probably annual, usually herbaceous throughout, sometimes suffrutes-

cent below, usually a meter high or less, branched; stipules dissected into numerous

gland-tipped filiform divisions; leaves long-petiolate, membranaceous, the petioles

bearing numerous branched hairs with gland-tipped divisions; leaf blades 7-15 cm.

long and wide, cordate at the base, deeply 3-5-lobate, more or less pubescent or

glabrous, the lobes acute or acuminate, denticulate, glandular-ciliate; flowers

small, greenish, in small pedunculate cymes, the bracts linear-oblong, glandu-

liferous on the margins; sepals ovate, glandular-ciliate, pubescent outside, 5-7

mm. long; petals 5, obovate, purple, somewhat longer than the sepals; stamens

usually 8; ovary pubescent; capsule 1 cm. in diameter, 3-sulcate, glabrate; seeds

oblong, brown, carunculate.

Called "frailecillo" in Salvador. The seeds contain an oil having

drastic purgative and emetic properties.

Jatropha hastata Jacq. Stirp. Sel. Amer. 256. pi. 173, f. 54.

1763.

Native of Cuba; cultivated in Pete"n, and possibly at times be-

coming naturalized; planted also in Panama and perhaps elsewhere

in Central America.

A slender shrub, glabrous or somewhat pubescent on the young parts; leaves

on long slender petioles, membranaceous, variable in shape, mostly ovate or oblong-

ovate, sometimes panduriform or hastate-lobate, mostly 8-14 cm. long, abruptly

acuminate, rounded at the base, somewhat paler beneath, palmate-nerved at

the base, penninerved above; cymes long-pedunculate, corymbiform, many-

flowered, the bracts linear-lanceolate, entire; staminate sepals ovate, obtuse,

3 mm. long; petals 10-12 mm. long, oblong, obtuse, bright red; ovary glabrous.

Jatropha multifida L. is said to be cultivated sometimes in Guate-

mala and probably is, although we have not seen it. In habit it is

similar to J. podagrica, but the long-stalked leaves are epeltate and

divided almost to the base into numerous narrow segments which

are again lobate.

Jatropha podagrica Hook. Bot. Mag. pi. 4376. 1848. Rui-

barbo; Capa de rey.

Cultivated commonly for ornament in gardens at low and middle

elevations; native on dry rocky hillsides, in open places or in dense

forest, 200-700 meters; Zacapa; Chiquimula. Apparently native

also in Honduras, and perhaps in Quiche", Guatemala; cultivated

widely for ornament in tropical America.

Plants erect, usually 60 cm. high or less, sometimes as much as 1.5 meters

when growing wild, generally with a short thick woody stem armed with short

sharp-pointed, spine-like bases of the stipules, the basal part of the stem greatly

enlarged and suggesting the top of a turnip, the more slender portion of the stem

usually 20 cm. long or shorter; leaves large, few, deciduous during the driest

130 FIELDIANA: BOTANY, VOLUME 24

months, long-petiolate, the petioles mostly 10-18 cm. long; leaf blades sub-

orbicular in outline, peltate near the middle, green above, very pale beneath, gla-

brous, shallowly or deeply 3-5-lobate, 10-25 cm. long and wide, the lobes very

broad, rounded at the apex and often abruptly short-acuminate; stipules glanduli-

ferous at first, becoming much indurate in age; cymes rather dense and many-

flowered, glabrous, on stout peduncles 35 cm. long or shorter, the flowers small,

brilliant red; staminate sepals 2 mm. long, entire, glabrous, rounded-ovate;

petals spatulate, glabrous, 6 mm. long, obtuse; stamens 6-8; ovary glabrous;

capsule about 15 mm. long and almost as broad, 3-sulcate; seeds 11 mm. long,

fuscous, smooth.

This plant is seen rather infrequently in gardens of Guatemala

and other parts of Central America, where it attracts attention be-

cause of the large peltate leaves, the small but showy, bright red

flowers, and especially because of the turnip-like base of the stem,

suggesting a turnip only half buried in the soil. The name "ruibarbo"

("rhubarb") is applied to it generally, and the plant is said to have

been used as a drug in Europe under the name "Guatemalan rhu-

barb." So far as we know, no medicinal use is made of it in Guate-

mala. The plant is certainly native on the very dry, rocky hills

about the divide on the road between Zacapa and Chiquimula. The

dry hills of eastern Guatemala and adjacent Honduras are apparently

the only known native habitat of this curious and much cultivated

plant, which has reached many distant parts of tropical America.

JULOGROTON Martius

Annual or perennial herbs or shrubs, abundantly pubescent; leaves alternate,

petiolate, 2-stipulate, usually denticulate or crenate, generally 3-5-plinerved;

inflorescence terminal, bisexual, densely spicate-racemose, the flowers 2-bracteo-

late, the pistillate ones at the base of the spike; bracts 1-flowered, dentate;

staminate sepals imbricate, the pistillate sepals imbricate and usually very unequal,

the outer ones usually with dorsal glands or appendages; petals 5; disk more or

less developed; stamens mostly 11, the anthers refracted in bud, dehiscent by

longitudinal slits; ovary of the pistillate flower 3-celled, the 3 styles connate below,

dichotomous; ovary rudiment none; ovules 1 in each cell; fruit capsular, the

2-valvate cocci separating from the persistent columella; seeds carunculate.

In general appearance the plants are exactly like some species

of Croton, and it would perhaps be more rational to unite them with

that genus. About 30 species, in tropical America, mostly in South

America. One other species is known from Central America, J. deca-

lobus (Muell. Arg.) Benth., which was described as a plant of Guate-

mala but is rather a native of Costa Rica.

Plants annual; inflorescences densely clustered at the ends of the branches; leaves

grayish or silvery when dried J. argenteus.

Plants frutescent; inflorescences mostly solitary; leaves brownish when dried.

J. conspurcatus.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 131

Julocroton argenteus (L.) Didrichs. PI. Mus. Univ. Hafn. 42.

1857. Croton argenteus L. Sp. PI. 1004. 1753.

Wet to dry fields or thickets, often in dried mud about former

pools, sometimes a weed in waste ground, 1,200 meters or less;

Jutiapa; Escuintla; Retalhuleu; Huehuetenango. Mexico; British

Honduras; Salvador to Costa Rica; northern South America.

An erect annual, usually less than a meter high, sparsely or much-branched

above and sometimes branched from the base, covered throughout with a whitish

or silvery pubescence of fine, dense, closely appressed hairs, the branches terete

or obtusely angulate, dichotomous or trichotomous above, the leaves appearing

verticillate at the forks of the branches, elsewhere alternate, on long slender

petioles or the upper ones subsessile; stipules setaceous; leaf blades 5-nerved,

broadly ovate or rhombic-ovate or the upper ones oblong-ovate, mostly 4-8 cm.

long, obtuse or subacute, obtuse to subcordate at the base, rather thick, obscurely

undulate-dentate or entire, usually silvery on both surfaces but sometimes green

above; racemes short and often head-like, usually numerous and crowded at the

ends of the branches and subtended by large leaves; staminate flowers slender-

pedicellate, in bud 1.5 mm. in diameter; pistillate sepals lance-obovate, acuminate,

incised-dentate above, 6-7 mm. long; style branches twice divided; capsule 4 mm.

long, stellate-tomentulose; seeds 3 mm. long, lustrous, smooth, minutely caruncu-

late, fuscous and somewhat mottled.

During the wet months this plant grows commonly in fields on

the Pacific plains, but it soon withers after the rains cease, being

found then only occasionally and usually in dry mud where there

were shallow pools during earlier months. There it usually is asso-

ciated with Glinus and one or more species of Heliotropium.

Julocroton conspurcatus (Schlecht.) Klotzsch in Wiegm.

Archiv 7: 193. 1841. Croton conspurcatus Schlecht. Linnaea 7: 380.

1832. J. triqueter var. conspurcatus Muell. Arg. in DC. Prodr. 15,

pt. 2: 705. 1862. J. montevidensis var. guatemalensis Muell. Arg. op.

cit. 703 (type said to have been collected in Guatemala by Friedrichs-

thal). Colipavo.

Wet or dry thickets or forest, about 1,000-1,600 meters; Alta

Verapaz; Chiquimula; Santa Rosa; Sacatepe"quez ; Quiche". Southern

Mexico.

A branched shrub, commonly 3 meters high or less, the branches densely

stellate-pilose with mostly fulvous or brownish hairs, these often loose and spread-

ing but sometimes more or less appressed; leaves membranaceous or thicker,

the lower ones long-petiolate, the upper ones on shorter petioles, ovate-lanceolate

to ovate, mostly 5-14 cm. long, long-acuminate to acute, rounded at the base,

3-nerved at the base, penninerved above, denticulate or almost entire, minutely

stellate-scaberulous and green on the upper surface, very rough to the touch,

densely stellate-pubescent or tomentose beneath, the pubescence usually rather

132 FIELDIANA: BOTANY, VOLUME 24

lax and spreading; flower spikes strobile-like, narrowly ovoid or oblong, short-

pedunculate, mostly 2-3 cm. long, very dense and many-flowered, densely and

coarsely stellate-tomentose with mostly brownish hairs; bracts triangular-ovate,

pinna tisect; pistillate sepals incised-lacerate.

This plant has been reported from Guatemala as J. triqueter

(Lam.) Baill.

MABEA Aublet

Reference: F. Pax, Pflanzenreich IV. 147, v: 26-42. 1912.

Shrubs or trees with milky sap, often sarmentose or somewhat scandent, the

branches generally very slender; leaves alternate, mostly oblong, entire or denticu-

late, penninerved, short-petiolate, often glaucous beneath; indument of branched

hairs, usually scant; flowers monoecious, apetalous, the racemes terminal, many-

flowered, sometimes paniculately branched, the bracts generally with 2 conspicuous

glands; staminate flowers numerous, mostly ternate within the bracts, often umbel-

late or spicate; pistillate flowers few at the base of the raceme, pedicellate, solitary

within the bracts; staminate calyx small, globose in bud, open long before anthesis,

3-5-lobate, the lobes broad, slightly imbricate; stamens 10-70, rarely 2-3, inserted

on a convex receptacle, the anthers almost sessile, extrorse, longitudinally dehiscent;

ovary rudiment none; pistillate sepals 5, rarely 6 or 3, often very unequal, im-

bricate; ovary 3-celled, the styles connate to form an elongate column, free above

and simple; ovules solitary in each cell; capsule globose or somewhat tridymous,

separating into 2-valvate cocci, the endocarp hard; seeds ovoid, carunculate, the

testa crustaceous, smooth; endosperm carnose, the cotyledons broad, flat.

Species about 30, in tropical America, all much alike and difficult

to distinguish. One other species is recorded from Panama.

Lateral nerves of the leaves united remote from the margin to form a distinct but

irregular collective nerve; seeds about 7 mm. long and 5 mm. broad.

M. occidentalis.

Lateral nerves of the leaves irregularly joined close to the margin, not forming a

distinct collective nerve; seeds about 9 mm. long and 7 mm. broad.

M. excelsa.

Mabea excelsa Standl. & Steyerm. Field Mus. Bot. 23: 123.

1944.

Moist or wet forest or thickets, 200-850 meters; endemic; Santa

Rosa; Retalhuleu; Quezaltenango (type collected near Colomba,

A. F. Skutch 2008) . Chiapas.

A tall glabrous tree as much as 30 meters high with a trunk 65 cm. in diameter,

the bark close, slightly flaky, cinnamon-colored, the branches very slender, terete,

green when young; leaves chartaceous, on slender petioles 6-12 mm. long, narrowly

oblong or lance-oblong, 8-13 cm. long, 2-4 cm. wide, narrowly long-acuminate

or abruptly acuminate, acute to rounded at the base, slightly lustrous above,

glaucescent beneath, the lateral nerves about 17 pairs, divergent at a wide angle,

almost straight, irregularly joined close to the margin; fruiting pedicels 18 mm.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 133

long, stout; columella 1 cm. long, thickened at the base and apex; capsule 1.5 cm.

long, very densely but minutely tomentulose, the valves hard and ligneous; seeds

thick, scarcely compressed, olivaceous, lustrous, smooth, 9 mm. long and 7 mm.

broad, the caruncle 1.5 mm. long.

Mabea occiden tails Benth. in Hook. Journ. Bot. 6: 364. 1854.

M. belizensis Lundell, Field & Lab. 13: 4. 1945 (type from British

Honduras, Toledo District, upper reaches of Golden Stream, P. H.

Gentle 4595).

Moist or wet thickets or forest, at or little above sea level; Izabal.

Chiapas and Tabasco; British Honduras, along the Atlantic coast

to Panama; southward to the Amazon Valley.

A shrub or small tree, 8 meters high or less, the trunk as much as 8 cm. in

diameter, glabrous outside the inflorescence or practically so, the branches usually

very slender; leaves chartaceous or coriaceous, on petioles 5-6 mm. long, oblong

to elliptic-oblong or lance-ovate, mostly 6-14 cm. long and 3-6 cm. wide, cuspidate-

acuminate or caudate-acuminate, obtuse or rounded at the base and often some-

what oblique, obscurely denticulate or almost entire, green and lustrous above,

very glaucous beneath, the lateral nerves 11-14 pairs; panicles about 10 cm. long

and 1.5-3 cm. broad, lax, the staminate branches 4-5 mm. long, umbellately 3-

flowered, bearing very large and conspicuous glands near the base, the pedicels

rufous-tomentulose, the flowers dark red or purple; staminate sepals semiorbicular,

obtuse, the pistillate triangular-ovate, acuminate, 3 mm. long; stamens about 25;

ovary tomentulose; style column almost 2 cm. long, the capsule slightly 3-sulcate,

1.5 cm. long, densely tomentulose; seed brown, smooth, 7 mm. long, 5 mm. broad,

with a small caruncle.

Reported from Guatemala as M. montana Muell. Arg. It is

stated (from British Honduras) that this plant — presumably the

milky sap — causes irritation and itching of the skin.

MANIHOT Adanson

References: F. Pax, Pflanzenreich IV. 147, ii: 21-99. 1910. Leon

Croizat, A study of Manihot in North America, Journ. Arnold Arb.

23: 216-225. 1942.

Shrubs, trees, or often herbs, usually glaucescent and glabrous or nearly so;

leaves alternate, petiolate, sometimes peltate, undivided or often deeply lobate,

the lobes entire or lobate, the stipules generally small; flowers rather large, monoe-

cious, apetalous, racemose or paniculate, the inflorescences terminal or arising

in the uppermost axils, simple or sparsely branched, with 1-few pistillate flowers

at the base, the pedicels often elongate; bracts small or large, entire or dentate;

staminate calyx sometimes colored, campanulate, shallowly or deeply 5-fid, the

lobes imbricate or contorted; stamens 10 and 2-verticillate, the filaments free,

the anthers dorsifixed, longitudinally dehiscent; ovary rudiment none or small;

pistillate calyx like that of the staminate flower, the hypogynous disk entire or

lobate; ovary 3-celled, the styles short-connate at the base, variously dilated or

134 FIELDIANA: BOTANY, VOLUME 24

lobate; ovules solitary in each cell; capsule separating into 2-valvate cocci; seed

carunculate, the testa crustaceous; cotyledons broad and flat.

About 130 species, all in tropical America, the great majority of

them Brazilian. One or two additional species may occur in southern

Central America.

Leaves peltate; trees or large shrubs with woody steins M. Glaziovii.

Leaves not peltate; stems herbaceous.

Lobes of the leaves entire.

Calyx pubescent inside, glabrous outside.

Ovary 6-angulate, the capsule winged; anthers very short. . . .M. esculenta.

Ovary terete, the capsule not winged; anthers elongate M. dulcis.

Calyx glabrous within and without M. aesculifolia.

Lobes of the leaves lobate or sinuate.

Lobes of the leaves linear or at least greatly elongate and narrow, their lateral

lobes acute or acuminate M. parvicocca.

Lobes of the leaves broadest at or near the apex, the terminal portion of the

lobe broadly rhombic, the lateral lobes rounded or very obtuse.

Leaves small, the middle lobe 4-6 cm. long; capsule 1 cm. long . M. ludibunda.

Leaves large, the middle lobes commonly 7-15 cm. long; capsule 1.5 cm.

long M. gualanensis.

Manihot aesculifolia (HBK.) Pohl, PI. Bras. Icon. & Descr.

1: 55. 1827. Janipha aesculifolia HBK. Nov. Gen. & Sp. 2: 107. pi.

109. 1817.

Moist thickets, at or little above sea level; Suchitepe"quez;

Quich^ ; San Marcos. Yucatan Peninsula of Mexico; Atlantic low-

lands of Honduras.

A coarse herb 1-3 meters high, perhaps sometimes suffrutescent below, gla-

brous throughout; leaves on petioles 6-12 cm. long, membranaceous, cordate at

the base, concolorous or somewhat glaucescent beneath, deeply 5-7-parted, the

lobes oblanceolate-oblong or obovate-oblong, 8-15 cm. long, acuminate, narrowed

to the base, entire, the outermost lobes very short; stipules caducous; racemes

5 cm. long, or in fruit sometimes much longer, the pedicels arcuate-recurved in

fruit, 2-3 cm. long; staminate calyx almost 2 cm. long, 5-fid to about the middle,

glabrous, greenish outside; disk and stamens glabrous; ovary glabrous, terete;

fruit globose, about 12 mm. high; seeds whitish or grayish, 7-8 mm. long, 7 mm.

broad.

The Maya names in Yucatan are reported as "batul" and "chac-

che"; "yuca cimarrona" (Yucatan).

Manihot dulcis (J. F. Gmel.) Pax, Pflanzenreich IV. 147, ii:

71. 1912. Jatropha dulcis J. F. Gmel. Onom. Bot. 5: 7. 1772-78.

M. palmata Muell. Arg. in DC. Prodr. 15, pt. 2: 1062. 1866. Yuca

duke.

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 135

Native of Brazil, but cultivated rather widely in other regions

of the tropics for its edible roots; sometimes planted in Guatemala.

Plants with large tuberous roots, herbaceous, commonly 1-2 meters high, more

or less puberulent on the young parts but in age almost wholly glabrous; leaves

large, long-petiolate, membranaceous, concolorous or glaucescent beneath, deeply

3-13-parted, the lobes lanceolate, oblanceolate, or lance-elliptic, acuminate or

acute; stipules narrow, setaceous-acuminate, entire or sparsely laciniate; panicles

many-flowered, the bracts small, lanceolate, entire; calyx 12-14 mm. long, glabrous

outside, puberulent within in the upper portion, the staminate calyx globose-

campanulate, 5-fid almost to the middle; pistillate calyx 5-parted; anthers several

times longer than broad; disk and filaments glabrous; ovary terete, slightly

costulate; capsule subglobose, not winged, 1.5 cm. long, rugose when dry.

The Maya name in Yucatan is listed as "cicitsin." It is stated

that the plant does not flower there. This form of cassava has

"sweet" roots, that is, they do not possess the poisonous properties

characteristic of M. esculenta. How extensively the plant is grown

in Guatemala we do not know, but it is apparently utilized much

less than M . esculenta,

Manihot esculenta Crantz, Inst. Herb. 1: 167. 1766. Janipha

Manihot L. Sp. PI. 1007. 1753. M. utilissima Pohl, PI. Bras. 1: 32.

pi. 24- 1827. Yuca; Tzin (Quecchi) ; Cassava.

Presumably native of tropical Brazil and neighboring regions,

but now grown widely in other parts of tropical America, also in

the Old World tropics; planted commonly and on a rather large

scale in the lowlands of Guatemala, and occasionally even at middle

elevations.

Plants arising from large tuberous roots, erect, 1-3 meters high, branched,

glabrous or nearly so; leaves long-petiolate, membranaceous, green and glabrous

above, usually very glaucous beneath, glabrous or minutely puberulent on the

nerves, 3-7-parted, the lobes 8-15 cm. long, spatulate-lanceolate or linear-lanceo-

late, acuminate, gradually attenuate to the base; stipules 5-7 mm. long, triangular-

lanceolate, setaceous-acuminate, entire or with 1-2 laciniations; peduncles mostly

5-6 cm. long, slender, equaling the panicle, the bracts linear-lanceolate, entire;

staminate pedicels 4-7 mm. long, the pistillate deflexed-spreading, as much as

2.5 cm. long; calyx campanulate, glabrous and pruinose outside, puberulent

within in the upper portion, 1 cm. long, 5-parted; disk and filaments glabrous;

anthers only 1.2 mm. long, hispidulous at the apex; ovary glabrous, acutely

6-angulate; capsule 1.5 cm. long, subglobose, rugose, narrowly 6-winged.

Sometimes called "yuca amarga" or "yuca brava"; the Maya

name in Yucatan is recorded as "tsin" or "tsiim." Yuca or cassava

is one of the most important of all food plants, supplying "bread" to

most of Brazil and other regions, and useful starches to other parts

of the earth, particularly in the form of tapioca. The plant is be-

136 FIELDIANA: BOTANY, VOLUME 24

lieved to have been introduced into tropical North America in

precolumbian times, perhaps transported by the Caribs from northern

South America to the West Indies and to the mainland. Among

the Caribs of the Atlantic coast of Central America cassava is still

an important foodstuff, and some of them still use the curious

elongate wicker baskets for expressing the juice from the grated

root, which is then made into large thin cakes. These are used in

place of the maize tortillas eaten as bread in most parts of Central

America. Outside the Carib settlements, yuca is used in Central

America only as a vegetable, boiled and eaten like sweet potatoes.

It is rather heavy and not too easily digestible, coarse and rather

hard, and has little to recommend it where other and better starchy

foods are available. Yuca is cultivated very extensively in all the

warmer parts of Guatemala, in the North Coast, the lowlands of

Alta Verapaz and Pete"n, in the irrigated Motagua Valley and else-

where in the Oriente, and in the largest amounts on the Pacific

plains, where extensive fields of it are planted. The large heavy

roots, looking somewhat like sweet potatoes, are transported from

the lowlands to the markets of the uplands. A great deal of starch

is made from the roots in Guatemala, to be used in starching clothes,

also in the preparation of atol, especially for invalids, and various

desserts. One of the last, much like blanc mange of the United

States, is prepared by boiling yuca starch with sugar and the in-

evitable cinnamon.

As a vegetable yuca must be used with some discretion. There

appear to be various races of M. esculenta, in some of which the roots

are intensely poisonous when raw, while in others they are sweet

and harmless. The poisonous property is stated to be hydrocyanic

acid, although others report that it is a peculiar principle, mani-

hotoxine. The poisonous property may be removed by expressing

the juice or by thorough cooking. If human beings or domestic

animals eat the raw roots of the bitter variety, death is likely to

ensue within a few hours. The names "yuca" and "cassava" are

of Antillean origin, and variants of the latter are "casabe" and

"casabi." The Brazilian name is "mandioca." The Nahuatl name

of the plant is "quauhcamotl" ("tree potato"). "Yuca," apparently,

was the Haitian name for the plant, "casabi" the term for bread

made from its roots. In some regions numerous varieties of the

cassava plant are recognized by cultivators. In Guatemala marked

forms are apparent in some of the plantations, differing in color of

the stems and shape of the leaves, but little or no attention is paid

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 137

to them. The plants are propagated from cuttings, and are cul-

tivated much like potatoes or any other similar root crop, but they

require a longer time for maturity than most similar crops.

Manihot Glaziovii Muell. Arg. in Mart. Fl. Bras. 11, pt. 2:

446. 1874.

Native of Brazil, but often planted in other tropical regions as

a source of rubber or for shade or as a curiosity; collected near Los

Verdes, Santa Rosa, in a hedge remote from dwellings, but probably

planted there; sometimes cultivated in Alta Verapaz, but not

commercially.

A small tree, usually 10 meters high or less, with a dense crown, glabrous

throughout or nearly so; leaves long-petiolate, membranaceous, green above,

glaucous beneath, peltate shortly above the base, deeply 3-5-lobate, about 12 cm.

long and 16 cm. wide, the lobes oblong-obovate or elliptic, rounded to abruptly

short-acuminate at the apex; stipules 4-6 mm. long, lance-ovate, acute, denticulate;

panicles 7-9 cm. long, broadly pyramidal, the bracts 2.5 mm. long, lanceolate, the

pedicels about 1 cm. long, the flowers nutant; calyx glabrous, campanulate, the

staminate 5-fid, 9 mm. long, green tinged with violet; pistillate calyx 5-parted,

1 cm. long; capsule globose, 2 cm. in diameter, not winged; seeds 1.5 cm. long,

1 cm. wide, gray mottled with brown.

This plant is the source of the Ceara rubber of commerce, formerly

an article of commercial importance. The trees were planted ex-

tensively in the East Indies, but their product was unable to compete

with Hevea rubber. In Salvador the tree is called "caucho bianco."

Manihot gualanensis Blake, Contr. U. S. Nat. Herb. 24: 13.

1922. Yuca cimarrona; Yuquilla.

Moist or wet thickets, 900 meters or less; Zacapa (type from

Gualan, S.F.Blake 7688); Chiquimula; Santa Rosa; Huehuetenango.

British Honduras; Honduras; Nicaragua.

A coarse herb 1-2 meters high, glabrous throughout or essentially so, the stems

stout, often much-branched; leaves on petioles 20 cm. long or less, 30 cm. wide

or smaller, membranaceous, about 9-lobate almost to the base, green above,

glaucous beneath, the lobes obovate or rhombic-obovate, usually shallowly lobate

above the middle and panduriform, acuminate, tapering to the base; panicles

about 10 cm. long, many-flowered, glabrous, the flowers nutant, short-pedicellate

or the staminate ones in age long-pedicellate; bracts linear-subulate, entire, 2-6

mm. long; pistillate calyx greenish, glaucous, 12 mm. long, the segments lance-

oblong, obtuse; ovary subglobose, glabrous; staminate calyx 7 mm. long, glabrous,

the lobes oblong, obtuse; stamens glabrous; capsule subglobose, 1.5 cm. long,

rugulose, somewhat 5-angulate at the base.

This species was described originally as a shrub, and other

species have been characterized in the same manner. As a matter

138 FIELDIANA: BOTANY, VOLUME 24

of fact, all the species enumerated here except M. Glaziovii are herbs.

Most of them grow luxuriantly during the rainy season but wither

when the rains cease, and nothing is seen of them during the long

verano.

Manihot ludibunda Croizat, Journ. Arnold Arb. 23: 219. 1942.

Yuca cimarrona.

On limestone bald knobs (pelones or pajales), 800-1,400 meters;

endemic; Huehuetenango (type from Guaxacana, C. & E. Seler

2814; collected also between Nenton and Miramar and at San Antonio

Huista).

Plants herbaceous, low, glabrous throughout; leaves on long slender petioles,

4-6 cm. long, mostly 5-parted almost to the base, the lobes spatulate-oblong,

constricted at about the middle and panduriform, the apex dilated-quadrangular,

short-acuminate, the lateral lobes rounded, green above, beneath concolorous or

somewhat paler; inflorescences apical or lateral, 3-5 cm. long; staminate flowers

on pedicels 7-10 mm. long, the calyx 1 cm. long, the short lobes ovate-acuminate;

pistillate sepals 5, lance-elliptic, 4 mm. long; ovary costulate; capsule 1 cm. long,

not winged, rugose-tuber culate; seeds 6-7 mm. long and almost as broad, smooth,

lustrous, mottled with gray and olive-brown, the caruncle large.

The type material of this was reported once as M . carthagenensis

(Jacq.) Muell. Arg.

Manihot parvicocca Croizat, Journ. Arnold Arb. 23: 219. 1942.

Open or brushy hillsides or plains, 1,200-2,400 meters; Baja

Verapaz; Guatemala (Fiscal); Quiche"; Huehuetenango. Chiapas,

the type from Siltepec; Honduras; Salvador.

Plants apparently perennial, erect, 30-100 cm. high, often much-branched,

glabrous throughout; leaves 5-7-lobate almost to the base, long-petiolate, mem-

branaceous, 16 cm. broad or smaller, the lobes unequal, the outermost much

shorter and smaller, the middle ones linear or linear-oblong, mostly 10 cm. long

or less, long-attenuate, all or most of them with a few small lobes or teeth, these

usually acute or acuminate, concolorous or nearly so; inflorescence terminal,

10 cm. long or less, lax, the pistillate pedicels stout in fruit and somewhat refracted;

staminate perianth campanulate, 1 cm. long, the lobes ovate-triangular; capsule

globose, 8-10 mm. long, the columella 3-4 mm. long; seeds 5-6 mm. long and

almost as broad, ochraceous or grayish mottled with olive, the caruncle rather

large and conspicuous.

OMPHALEA L.

Reference: F. Pax, Pflanzenreich IV. 147, v: 14-22. 1912.

Shrubs or large trees, sometimes scandent shrubs; leaves alternate, stipulate,

penninerved or 3-5-nerved from the base, the petioles 2-glandular at the apex;

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 139

flowers small, monoecious, apetalous, cymulose, the cymules staminate, with a

single central pistillate flower, disposed in terminal panicles; bracts usually elongate

and subfoliaceous, petiolate, the petiole 2-glandular; disk none or obsolete; stami-

nate sepals 4-5, broad, strongly imbricate; stamens 2-3, the filaments connate

into a short column, the connective thick and broad, connate to form a peltate

pileiform 2-3-lobate mass, the cells peripheral, longitudinally dehiscent; ovary

rudiment none; pistillate calyx like that of the staminate flower; ovary 2-3-celled,

the style columnar, thick, obtuse or shallowly 2-3-lobate; ovules solitary in each

cell; fruit large, somewhat carnose, the endocarp hard, indehiscent or finally

separating into 2-valvate cocci; seeds large, almost globose, not carunculate;

cotyledons broad, flat.

Fifteen species, 3 in the Old World tropics, the rest American.

One other known from Central America (0. diandra L.) may well

occur in the Atlantic lowlands of Guatemala.

Omphalea oleifera Hemsley, Pharm. Journ. Trans. XV. 13:

301. 1882. Palo de queso.

Dry, thinly forested hillsides, mostly at 500 meters or less;

Jutiapa; Escuintla. Salvador.

A large or medium-sized tree, often 15 meters high or more, with a tall pale

trunk and a rounded or spreading crown; leaves very large, on long stout petioles,

membranaceous, deciduous, rounded-cordate, often 30 cm. wide and somewhat

longer, abruptly acute or short-acuminate from an almost rounded apex, deeply

cordate at the base, entire or nearly so, 7-nerved at the base, when young some-

what stellate-puberulent but in age glabrous; flowers monoecious, greenish, in

large leafy-bracted panicles, the branches puberulent; bracts oblong, petiolate,

as much as 2.5 cm. long, puberulent; sepals 4, decussate, orbicular, ciliate; stamens

2; ovary glabrous; fruit subglobose, green or yellowish, 3 cm. or more in diameter;

seeds 3, very large, blackish.

Known in Salvador by the names "hoja de queso," "chiran,"

"shilan," "palo de jabon," "tambor," and "castafiete." The large

thin leaves are used there for wrapping cheeses. The young fruits

as well as the mature seeds are boiled and eaten, but in case of the

seeds it is stated that the embryos must be removed. The seeds

are rich in oil, which is used in Salvador for cooking, illumination,

and manufacture of soap.

OPHELLANTHA Standley

Large shrubs or small trees, the pubescence of simple hairs; leaves alternate,

slender-petiolate, membranaceous, penninerved, remotely denticulate; stipules

2, small, spinose, persistent; flowers monoecious, long-pedicellate, solitary or

fasciculate on axillary spurs; staminate calyx 5-parted, the lobes slightly im-

bricate; petals 5, distinct, entire, much longer than the calyx, sessile, ciliate;

disk large, densely short-hirsute; stamens numerous, 50 or more, irregularly

140 FIELDIANA: BOTANY, VOLUME 24

inserted over the disk, the filaments elongate, filiform, glabrous; anthers small,

2-celled, dehiscent by 2 introrse slits, each cell bearing at the apex a short filiform

appendage; ovary rudiment none; pistillate sepals 5, accrescent and foliaceous

after anthesis; margin of the disk very shallowly 5-lobate; ovary 2-3-celled, sessile;

styles 2-3, very stout, nearly or quite distinct, shortly 2-fid; ovules solitary in

each cell; capsule separating into 2-3 bivalvate cocci, the columella persistent;

seeds large, not carunculate, smooth or nearly so; endosperm carnose, the coty-

ledons broad, flat.

One other species is known, ranging from Oaxaca to western

Costa Rica. It is to be expected in Guatemala.

Ophellantha Steyermarkii Standl. Field Mus. Bot 23: 123.

1944.

Known only from the type, Dept. Huehuetenango, trail between

Santa Ana Huista and Nenton, over Rio Azul and La Laguna, 800-

900 meters, Steyermark 51398.

A tree of 6 meters, the branches slender, terete or brownish, when young

pilosulous with lax spreading hairs, the internodes elongate but shorter than the

leaves; stipules indurate, spine-like, sharp-pointed, 2-2.5 mm. long; leaves on

slender petioles 10-13 mm. long, membranaceous, bright green when dried,

lanceolate or lance-oblong, 5-6.5 cm. long, 1.5-2.5 cm. wide, narrowed to the

subacute apex, cuneately acute or obtuse at the base, subentire, densely pilose on

both surfaces with very slender, pale, spreading hairs, slightly paler beneath, the

lateral nerves about 5 pairs; pistillate pedicels in fruit slender, 2.5 cm. long;

sepals foliaceous, ovate or oblong-elliptic, about 17 mm. long and 7 mm. wide,

narrowed to the obtuse apex, 5-nerved, densely pilosulous on both surfaces, entire;

styles shortly connate at the base, the free portion thick, densely appressed-

pilosulous, 2-fid to the middle, the ultimate branches very stout; disk densely

hirsutulous; capsule 3-celled, subglobose, densely velutinous-pilosulous with

spreading hairs, about 12 mm. high; seeds 3, oval, very thick, 8 mm. long, 6 mm.

broad, smooth, lustrous, fuscous brown.

The only other species, 0. spinosa Standl., is called "limoncillo"

in Salvador. Occurring there and in Oaxaca, it is confidently to be

expected in Guatemala. 0. Steyermarkii may be only a pubescent

form of 0. spinosa, but it appears to differ definitely from the latter,

in which the leaves and other parts are almost glabrous, the scant

pubescence consisting of appressed hairs. In the original material

of 0. spinosa the united portion of the styles is much longer and the

ovary 2-celled, but the latter character probably is not a constant one.

PEDILANTHUS Poiteau

Reference: C. F. Millspaugh, The genera Pedilanthus and Cuban-

thus and other American Euphorbiaceae, Field Mus. Bot. 2: 353-

377. 1913.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 141

Mostly erect shrubs, sometimes herbs, the stems fleshy, green, exuding a

copious milk-like latex; leaves alternate, succulent, broad, entire, early deciduous,

the plants often leafless; stipules gland-like or none; flowers small, surrounded

by a usually shoe-shaped and colored involucre, naked, monoecious; involucres

borne in terminal cymes or often in the axils of the uppermost leaves; involucre

pedunculate, its tube cleft on the upper side, bifid at the apex of the lower of the

2 lips, with 1 middle and 2 lateral lobes more or less closing the fissure, the tube

bearing at the base on the upper side a spur-like appendage, this 2-fid or 2-3-

lobate and extending anteriorly above the upper side of the basal part of the

tube; flowers pedicellate within the involucre, the staminate numerous, the pistil-

late solitary, naked or sometimes with linear bractlets at the base; style of the

pistillate flower elongate, this finally protruding, usually declinate; stigmas 3,

long-connate, often distinct at the apex; ovary 3-celled, the cells 1-ovulate; fruit

capsular; seeds not carunculate.

Species 30 or more, mostly in tropical America, a few in tropical

Africa. One other Central American species has been described from

Nicaragua. The species of this genus are all much alike in general

appearance, and separated chiefly by floral characters which, while

not difficult to see, are of doubtful significance. The true number

of valid species probably is smaller than has been generally recognized.

Involucres rather densely tomentulose P. camporum.

Involucres glabrous.

Lobe of the appendage at the base of the tube 2-parted. Leaves glabrous.

P. macradenius.

Lobe of the appendage entire.

Leaves glabrous; lobe of the appendage glabrous on the margins or nearly so.

P. tithymaloides.

Leaves pubescent beneath, often densely so; lobe of the appendage conspicu-

ously and usually densely ciliate P. Deamii.

Pedilanthus camporum Standl. & Steyerm. Field Mus. Bot.

23: 124. 1944.

Known only from the region of the type locality, dry brushy

plains, 120 meters or less, between Nueva Linda and Champerico,

Retalhuleu, Standley 87781.

An erect shrub about a meter high, the stems thick, terete, dark green, when

young sparsely tomentulose, soon glabrate; leaves deciduous, not seen; cymes

short, dense, clustered at the ends of the branches; bracts small, cucullate, densely

ochraceous-tomentose, the peduncles 6 mm. long or less, sparsely tomentulose or

glabrate; involucre 8-10 mm. long, 3 mm. wide at the middle, densely tomentulose

or puberulent, red and green, glabrous within, cleft along the upper margin to

the base, shallowly cleft below, the apices rounded or very obtuse; appendage

very small, scarcely more than 4 mm. long, cucullate-dilated at the base, the lip

short, liguliform, obscurely retuse; staminate pedicels glabrous, exserted, the

pistillate puberulent or tomentulose; ovary densely whitish-tomentulose, the

142 FIELDIANA: BOTANY, VOLUME 24

slender glabrous style 6-7 mm. long; capsule globose-trigonous, 6 mm. long and

broad, sparsely puberulent; seeds smooth, dirty brown, 3-3.5 mm. long.

Pedilanthus Deamii Millsp. Field Mus. Bot. 2: 356. 1913.

Tithymalus Deamii Croizat, Amer. Journ. Bot. 24,: 704. 1937. Pie

de nino.

Dry rocky plains or brushy hillsides, 200-1,300 meters; Chiqui-

mula; Jutiapa; Santa Rosa; Guatemala (type from Fiscal, C. C.

Deam 6081). Chiapas.

A stiff erect shrub 0.5-1.5 meters high, the branches suberect, stout, fleshy,

dark green, tomentulose when young, flexuous; leaves sessile or short-petiolate,

fleshy, oblong-ovate to broadly ovate or suborbicular, 4.5-8 cm. long, 2.5-3.5

cm. wide, acute or acuminate, obtuse or cuneate at the base, puberulent or glabrous

above, puberulent or pilosulous beneath, often densely so, the costa elevated, thin

and wing-like; bracts small and inconspicuous, cucullate; peduncles glabrous, 5-6

mm. long; involucres bright or deep red or rose, about 12 mm. long, glabrous, the

upper fissure extending to the appendage, the principal lobes ovate, erose-dentate;

appendage small, less than one-third the length of the tube, the lip ligulate, retuse;

staminate pedicels puberulent at the apex, the filaments pilose; capsule glabrous;

seeds grayish olive, almost 4 mm. long.

This plant sometimes is planted for hedges in the Oriente. In

this and other species the costa on the lower surface of the leaf is

greatly elevated and projects as a ruffled wing. It is questionable

whether P. Deamii is more than a pubescent variety of P. tithy-

maloides.

Pedilanthus macradenius Donn. Smith, Bot. Gaz. 19: 263.

1894. Tithymalus macradenius Croizat, Amer. Journ. Bot. 24: 704.

1937.

Known only from the type, Canibal, Huehuetenango, 950 meters,

W. C. Shannon 412.

Plants glabrous outside the inflorescence; leaves obovate-oblong, 10-15 cm.

long, 5-7.5 cm. wide, obtuse at each end, on petioles 6 mm. long; cymes arising

in the upper leaf axils, the bracts oblong-ovate, longer than the peduncles, as much

as 12 mm. long, reddish, pubescent; peduncles pubescent; involucre red, 10-15

mm. long, 6 mm. broad, glabrous outside, pubescent within, the principal segments

ovate, acute, pilose at the apex; pedicels glabrous; appendage almost half as long

as the tube of the involucre, cleft one-third its length into 2 broadly ovate lobes;

capsule depressed-globose, 8 mm. in diameter, the cocci carinate; seeds trigonous-

globose.

One sterile collection (Steyermark 51103) from Rio Trapichillo,

Huehuetenango, with the vernacular name "pie de nino" is probably

referable here, but the leaves are softly pilose beneath.

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 143

Pedilanthus tithymaloides (L.) Poiteau, Ann. Mus. Paris 19:

390. 1812. Euphorbia tithymaloides L. Sp. PI. 453. 1753. Pie de

nino; Pie de santo (Pet&i); Itamo real, Dictamo, Tomo real (fide

Tejada).

Dry to wet thickets, mostly at 300 meters or less; often planted

in hedges or gardens, and sometimes escaping; Pete"n; Izabal;

Escuintla; Retalhuleu; often planted elsewhere. Mexico; British

Honduras to Panama; West Indies; Colombia and Venezuela.

An erect branched shrub 1.5 meters high or less, the branches thick or rather

slender, dark green, terete; leaves almost sessile, thick and fleshy, broadly ovate

to oblong, 4-7.5 cm. long, acute or obtuse, cuneate to rounded at the base, glabrous;

cymes terminal, dense, the bracts ovate, acuminate, slightly longer than the

peduncles, caducous; involucre red or pink and green, 11-13 mm. long, glabrous;

appendage 4-glandular, the lobe short, linear; staminate pedicels glabrous, the

pistillate pubescent; capsule about 7.5 mm. long and 9 mm. broad, the cocci

carinate; seeds ovoid, 5 mm. long.

The cultivated plants often have dark red or purple leaves, but

many of the plants usually are nearly or quite leafless. They are

much planted for low hedges in the lowlands or even at middle eleva-

tions in Central America. They make a dense compact growth,

and are rather handsome or at least novel in appearance. The

abundant milky sap flowing from the broken stems is generally

considered poisonous.

PERA Mutis

Reference: F. Pax, Pflanzenreich IV. 147, xiii: 2-13. 1919.

Trees, the indument lepidote or stellate-lepidote, rarely of simple hairs;

leaves alternate, short-petiolate, entire, penninerved; flowers dioecious or rarely

monoecious, apetalous, enclosed in globose involucres, these 1-2-bracteolate at

the base, open laterally in anthesis or valvate-bifid, usually unisexual; staminate

flowers 3-4 perfect ones and a few sterile ones; pistillate flowers 3-4, sometimes

with rudimentary central staminate flower; flowers sessile, the involucres fascicu-

late in the leaf axils; disk none; staminate calyx small, valvate, turbinate, some-

times reduced or none; filaments short and free or longer and connate below,

the anthers dorsifixed at or above the base, the cells parallel, longitudinally

dehiscent; pistillate flowers naked, the ovary 3-celled, the style very short, the

stigma peltate, disk-like or 3-lobate; ovules solitary in each cell; capsule 3-coccous,

the cocci 2- valvate, separating from the persistent columella; seeds ovoid or

obovoid and compressed, carunculate, the testa black, smooth, lustrous; endosperm

carnose, the cotyledons broad, flat.

About 20 species, in tropical America, only two in North America.

Leaves not barbellate beneath in the axils of the nerves, the pubescence wholly

lepidote P. arborea.

Leaves densely barbellate beneath in the axils of the nerves, also lepidote.

P. barbellata.

144 FIELDIANA: BOTANY, VOLUME 24

Pera arborea Mutis, Svensk. Vet. Akad. Handl. Stockholm 5:

299. pi. 8. 1784.

Reported by Lundell from British Honduras, Stann Creek,

Mullins River-Stann Creek road, in broken ridge, P. H. Gentle 3361 ;

Panama; Colombia.

A large or medium-sized tree, sometimes 15 meters high or more, the branch-

lets terete, densely lepidote at first, glabrate in age; leaves on petioles 1-1.5 cm.

long, oblong or obovate-oblong, mostly 7-16 cm. long and 2.5-5 cm. wide, obtuse

or acute, rounded to broadly acute at the base, coriaceous, lustrous and glabrous

above, slightly paler beneath and sparsely and minutely lepidote; involucres borne

mostly at defoliate nodes, lepidote, the staminate 3-flowered, the pistillate 4-5-

flowered; staminate calyx turbinate, acutely dentate, the stamens 4-5; ovary

glabrous or sparsely lepidote, the stigma 3-lobate; capsule about 12 mm. long and

10 mm. broad, obo void-globose, undulate-rugulose, very sparsely lepidote; seeds

black, lustrous, slightly compressed, acutish at the apex.

The wood in this genus varies in color from light to very dark

brown, of fine or coarse texture, of medium or very high density.

It is little used except for poles and fuel.

Pera barbellata Standl. Field Mus. Bot. 8: 19. 1930.

Mixed forest, 300 meters or less; Pete"n. Southern Mexico

(Oaxaca; Tabasco); British Honduras (type from Mullins River

road, W. A. Schipp 201).

A tree of 9-20 meters with narrow crown, the trunk round, 40-70 cm. in

diameter, tall, the bark chocolate-brown with grayish patches and with small

scales, the inner bark deep yellow, the branchlets slender, densely lepidote when

young; leaves on petioles 5-7 mm. long, chartaceous, oblong-elliptic or lance-

oblong, mostly 4-7 cm. long and 2.5-3.5 cm. wide, acuminate, often abruptly so,

with an obtuse tip, usually acute at the base or abruptly contracted, sometimes

obtuse, minutely puberulent above on the costa, lustrous, sparsely stellate-

lepidote beneath, densely short-barbate in the axils of the nerves, the lateral

nerves about 13 pairs; flowers fasciculate in the leaf axils or at defoliate nodes,

the fruiting pedicels 2-3 mm. long, densely stellate-puberulent; staminate in-

volucres very small, scarcely more than 3 mm. long, densely lepidote; pistillate

involucres globose-obovoid, 6-8 mm. long, very densely stellate-puberulent,

rounded at the apex; capsule 8-10 mm. long, densely stellate-puberulent.

Called "palo prieto" in Oaxaca. The sapwood is white to deep

yellow, darkening slightly on exposure to air; heartwood well defined,

medium brown to chocolate-brown; has a slightly unpleasant odor

when first cut. The wood is said to be used in Oaxaca for railroad

ties.

PHYLLANTHUS L.

Mostly herbs or rather small shrubs, rarely trees, variable in habit; leaves

small or large, generally alternate, entire, often distichous, the branchlets with

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 145

their leaves often simulating compound leaves; flowers small, greenish or whitish,

axillary or rarely at defoliate nodes, the staminate mostly numerous and glomerate,

subsessile or pedicellate, the pistillate flowers in the same axils or on distinct

branchlets, few or solitary; flowers monoecious or rarely dioecious, apetalous,

with or without a disk; staminate sepals generally 5 or 6, distinct or short-con-

nate, imbricate, more or less 2-seriate, herbaceous or subpetaloid; stamens mostly

3, the filaments free or connate into a central column; anther cells parallel or

divergent, longitudinally dehiscent, sometimes transversely 2-valvate or confluent;

ovary rudiment none; ovary 3-many-celled, the styles distinct or connate, entire

or usually 2-fid, subulate or dilated; ovules 2 in each cell; fruit capsular, dry or

carnose, usually separating into 2-valvate cocci; seeds somewhat 3-angulate, con-

vex dorsally, not strophiolate; endosperm carnose, the cotyledons flat.

Species about 500, in both hemispheres, most numerous in the

tropics. Two or three others are known from southern Central

America. The genus has not been monographed in recent years,

and is in need of critical attention, although the Central American

species are not numerous, and rather well understood.

Plants annual or essentially so, the stems soft and herbaceous throughout; leaves

less than 2 cm. long, most of them less than 1 cm. long; stems rarely suffru-

tescent, but the leaves then all less than 1 cm. long.

Leaves very minutely ciliolate; ovary minutely tuberculate P. Urinaria.

Leaves not ciliolate; ovary not tuberculate.

Stems conspicuously thickened at the base; plants growing in marshes, often

in shallow water P. diffusus.

Stems not thickened at the base; plants not growing in very wet soil.

Upper parts of the stems conspicuously compressed and 2-edged.

P. compressus.

Upper parts of the stems terete or nearly so, not 2-edged.

Branches erect or strongly ascending, the plants somewhat fastigiately

branched P. carolinensis.

Branches spreading or recurved, the plants not at all fastigiately branched.

Capsules about 2 mm. broad; seeds smooth; leaves with very faint

and indistinct lateral nerves P. Niruri.

Capsules about 3 mm. broad; seeds somewhat scaberulous; leaves

with 4-5 pairs of distinct lateral nerves P. lathyroides.

Plants perennial, shrubs or trees, rarely low but then with hard and distinctly

woody stems, usually much more than 2 cm. long and often 10 cm. or more.

Flowers in elongate racemes or panicles; fruit large, often very large, often fleshy.

Leaves mostly 3-5 cm. long, almost as wide as long P. acidus.

Leaves mostly 9-20 cm. long, usually much longer than wide. .P. glaucescens.

Flowers fasciculate or solitary in the leaf axils.

Leaves very obtuse or rounded at the apex, not at all narrowed to the apex.

Leaves 8-15 mm. wide; sepals obovate P. ferax.

Leaves mostly 20-30 mm. wide; sepals broadest at or near the base.

P. micrandrus.

Leaves acute or acuminate, or rarely narrowed to an obtuse apex.

Capsule 4-celled; staminate flowers with 4 sepals P. nobilis.

146 FIELDIANA: BOTANY, VOLUME 24

Capsule 3-celled; staminate flowers with 5-6 sepals.

Older branches very rough and thick, sparsely or densely beset with the

large persistent indurate stipules, the young branches resembling

pinnate leaves.

Leaves glabrous P. Purpusii.

Leaves densely pilose beneath P. minarum.

Older branches not much thicker than the young ones, not rough, the

stipules usually early deciduous, the young branches not resembling

pinnate leaves.

Pistillate flowers densely fasciculate at the base of the pistillate flower;

branches conspicuously angulate P. brasiliensis.

Pistillate flowers solitary, or with 1-2 staminate flowers at their base,

the flowers often dioecious.

Leaves chartaceous, acuminate to an obtuse tip P. longipes.

Leaves thin-membranaceous, acuminate or attenuate to a very acute

tip.

Capsule about 7 mm. long P. Austinii.

Capsule about 3 mm. long.

Petioles glabrous and smooth; leaf blades narrowly rounded at

the base P. capillipes.

Petioles papillose-puberulent; leaf blades acute at the base.

P. Bartlettii.

Phyllanthus acidus (L.) Skeels, U. S. Dept. Agr. Bur. PI. Ind.

Bull. 148: 17. 1909. Averrhoa acida L. Sp. PI. 428. 1753. Cicca

disticha L. Mant. PI. 1: 124. 1767. P. distichus Muell. Arg. in DC.

Prodr. 15, pt. 2: 413. 1866. Grosella.

Moist or dry thickets, sometimes in second growth, 500 meters

or less; Jutiapa (Lago de Giiija); Escuintla (San Jose"). Native of

India, often planted in other tropical regions, and thoroughly

naturalized in some parts of Central America (mostly along the

Pacific lowlands) and elsewhere in tropical America.

A shrub or tree, mostly 2-9 meters high, with pale bark, the older branches

stout, the young ones very slender, with their leaves resembling pinnate leaves,

sometimes deciduous with the leaves; leaves distichous, almost sessile, broadly

ovate to suborbicular, mostly 3-6 cm. long, acute to rounded at the apex, rounded

or even emarginate at the base, thick and firm, pale beneath, with 6-9 pairs of

lateral nerves, glabrous; flowers monoecious, short-pedicellate and axillary or

usually in many-flowered, raceme-like panicles; sepals 4 in the staminate flower,

obovate, the stamens 4, with free filaments; pistillate sepals elliptic; ovary usually

4-celled, the 4 styles free, reflexed, 2-parted; fruit 1 cm. or more in diameter, green

or yellowish, the pericarp fleshy and juicy, very acid, deeply 6-8-sulcate vertically.

Sometimes called "guinda" and "piemiento" in Salvador; "wild

plum" (British Honduras). English names are "Otaheite goose-

berry" and "West Indian gooseberry." The fruit may be as much

as 2 cm. in diameter and is distinguished by its usually green coloring

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 147

and several vertical ridges. It is intensely acid and somewhat

astringent, with a flavor suggestive of a green gooseberry. Little

or no use is made of the sour fruit in Central America, but in other

regions it is sometimes made into preserves or pickles. The wood

is described as rather hard and fine-grained, with a specific gravity

of about 0.57. In some parts of Central America this small tree

has become thoroughly naturalized, especially in land about ponds

and lakes that is inundated during the rainy months but very dry

in the dry season.

Phyllanthus Austinii Standl. Field Mus. Bot. 22: 38. 1940.

Dense wet mixed forest, at sea level; endemic; Izabal (type from

Escoba, across the bay from Puerto Barrios, Standley 72868; collected

several times in the same region).

A very slender shrub 1-1.5 meters high, sometimes more elongate and even

subscandent, glabrous throughout, the branches terete, greenish or ochraceous,

with mostly elongate internodes; stipules greenish, erect, linear-subulate, 2.5-3

mm. long; leaves on petioles 5-17 mm. long, membranaceous, ovate or lance-

ovate, 6-10 cm. long, 2-5.5 cm. wide, narrowly long-acuminate, rounded or broadly

obtuse at the base, glaucescent beneath, the veins prominent and reticulate on

both surfaces, the lateral nerves 5-6 pairs; flowers dioecious, axillary, solitary or

few and fasciculate, the staminate pedicels filiform, scarcely more than 4 mm. long,

the greenish flowers 3 mm. broad; pistillate pedicels capillary, 2-4 cm. long,

flexuous; styles 3, very short, recurved; capsule obo void-globose, about 8 mm.

long, 3-carinate; seeds 3, pale brownish, 4 mm. long, densely and minutely verru-

culose.

The species was named for Mr. George B. Austin of the United

Fruit Company, Puerto Barrios, to whom the authors are deeply

obligated for his substantial and courteous assistance in their

explorations of Guatemala.

Phyllanthus Bartlettii Standl. Carnegie Inst. Wash. Publ. 461 :

68. 1935.

Moist or wet forest on limestone, at or little above sea level;

endemic; British Honduras (type from river bluffs, El Cayo, H. H.

Bartlett 11441; collected also at various other localities).

A slender erect shrub a meter high or less, glabrous almost throughout, the

older branches ferruginous, terete, the young ones very minutely puberulent or

papillose-scaberulous, with short or elongate internodes; leaves on slender petioles

5-8 mm. long, the petioles papillose-scaberulous; leaf blades lance-oblong or ovate-

oblong, 5-7.5 cm. long, 1.7-2.8 cm. wide, acuminate or long-acuminate, with an

acute or attenuate tip, acute or obtuse at the base, glabrous, membranaceous,

somewhat paler beneath, the lateral nerves about 5 pairs, the veins inconspicuous;

flowers probably dioecious, the staminate axillary, fasciculate, the filiform pedicels

148 FIELDIANA: BOTANY, VOLUME 24

glabrous, 2-3 mm. long; sepals white, 1.5 mm. long, rounded at the apex, glabrous;

stamens 5; pistillate pedicels 2-2.5 cm. long, capillary, axillary, solitary; capsule

4 mm. broad, rather deeply 3-lobate, the columella slightly more than 2 mm. long;

seeds 2 mm. long, brown or stramineous, deeply impressed-punctate and some-

what corrugate longitudinally.

Phyllanthus brasiliensis (Aubl.) Poir. Encycl. 5: 296. 1804.

Conami brasiliensis Aubl. PI. Guian. 927. pL 354- 1775. P. acuminatus

Vahl, Symb. Bot. 2: 95. 1791. P. Conami Swartz, Prodr. Veg. Ind.

Occ. 28. 1788.

Wet to dry thickets, often in second growth, 1,200 meters or

less; Izabal; Zacapa; Jutiapa; Santa Rosa; Escuintla; Chimalte-

nango; Suchitepe"quez; Retalhuleu; San Marcos. Mexico; British

Honduras to Salvador and Panama; West Indies; tropical South

America.

A slender shrub or small tree, mostly 1.5-3 meters high, sometimes taller, the

branches slender, spreading, green, angulate, puberulent on the angles; leaves

membranaceous, on petioles scarcely more than 2 mm. long, oblong-ovate to

rounded-ovate, mostly 2-4.5 cm. long and 1-3 cm. wide, acute or acuminate and

mucronulate, usually rounded or very obtuse at the base, minutely ciliolate on

the margins, elsewhere glabrous or nearly so, usually pale beneath; stipules small,

linear-subulate; flowers very small, greenish white, in dense axillary fascicles,

one pistillate, the others staminate, on pedicels 2 mm. long or less; staminate sepals

6, ovate or oblong, the stamens 3; pistillate pedicels about 5 mm. long or in fruit

somewhat elongate, the sepals 6; ovary 3-celled, smooth, the styles free, 2-branched,

reflexed; capsule green, 3-4 mm. long, 3-sulcate, reticulate- veined; seeds reddish

brown, cristate dorsally.

Known in Salvador by the names "pimientillo" and "palo de

zope"; "ciruelillo" (British Honduras); "xpibul," "cahyuc," "xpay-

hul" (Yucatan, Maya). This is one of the commonest shrubs in

second growth thickets in the lowlands of all Central America, often

occurring in great abundance. The main branches bear all or most

of their lateral branches and leaves in a single plane, in such a

manner that they resemble the large fronds of a tree fern.

Phyllanthus capillipes Blake, Contr. U. S. Nat. Herb. 24: 10.

1922.

Known in Guatemala only from the type, in gravelly places,

Quebradas, Izabal, S. F. Blake 8614. Atlantic coast of Honduras.

A slender sufifrutescent plant scarcely more than 40 cm. high, glabrous through-

out, branched, the branches very slender and wiry, glabrous, somewhat flexuous

or zigzag; stipules subulate, 1.5 mm. long; leaves membranaceous, on slender

petioles 3-13 mm. long, lance-ovate or lanceolate, 3-5.5 cm. long, 1-2 cm. wide,

acuminate or attenuate-acuminate, rather narrowly rounded at the base, pale

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 149

beneath, with about 6 pairs of lateral nerves; flowers axillary, usually 1 pistillate

and 2-3 staminate in each axil, the staminate pedicels 5-7 mm. long, capillary;

sepals 5, scarcely 1 mm. long; stamens 5, the anthers vertically dehiscent; pistillate

pedicels capillary, 2-3 cm. long, the 5 sepals broadly ovate, subacute, 1-1.3 mm.

long, pale-marginate; styles 3, united at the base, 2-parted; capsule 3-celled,

glabrous, 3 mm. broad; seeds brownish, curved, 1.8 mm. long, verrucose in about

5 lines.

Phyllanthus carolinensis Walt. Fl. Carol. 228. 1788.

Moist or wet fields or thickets, often on open banks or on sand-

bars along streams, frequently in cultivated ground, 1,500 meters

or less; Pete"n; Alta Verapaz; Jutiapa; Santa Rosa; Escuintla;

Guatemala; Suchitepe"quez; Retalhuleu; San Marcos. Southeastern

United States; Mexico; British Honduras to Panama; West Indies;

northern South America.

Plants annual, erect, 30 cm. high or less, very slender, soft-stemmed, glabrous,

rather densely branched from above the base, the branches terete, erect or ascend-

ing, densely or rather sparsely leafy; stipules triangular-lanceolate, usually with

1-2 coarse teeth; leaves on petioles 1 mm. long or less, membranaceous, narrowly

to broadly obovate, mostly 1-1.5 cm. long and 2-8 mm. wide, sometimes some-

what larger, obtuse or rounded at the apex, cuneate at the base, pale green and

usually somewhat glaucous beneath; flowers mostly geminate in the leaf axils,

1 staminate, the other pistillate, the pedicels short, scarcely exceeding the calyx;

fruiting calyx 3 mm. broad, persistent, the sepals linear-obovate; ovary smooth;

capsule 2 mm. broad, depressed-globose; seeds dark brown, puncticulate-scaberu-

lous.

The Maya name in Yucatan is recorded as "cababesinixte."

Phyllanthus compressus HBK. Nov. Gen. & Sp. 2: 109.

1817.

Grassy open places or in wet fields, 200-1,000 meters; Zacapa;

Jutiapa; Alta Verapaz; reported from Escuintla. Southern Mexico;

British Honduras; reported from Peru, and perhaps elsewhere in

South America.

Plants annual, rather stiffly erect, mostly 30 cm. high or less, branched from

above the base or sometimes simple, the branches erect or strongly ascending, the

older ones brownish, the younger ones strongly compressed and 2-edged; stipules

cordate at the base and usually produced on one side below the point of insertion;

leaves almost sessile, oblong-elliptic or oblong-obovate, mostly 6-14 mm. long,

very obtuse or rounded at the apex, acute or obtuse at the base, slightly paler

beneath, the lateral nerves inconspicuous; flowers monoecious, on very short

pedicels, often subsessile, the pedicels aggregate; calyx 2 mm. broad, the sepals

broad, obtuse; capsule 3 mm. broad, 3-celled; seeds brown, very minutely puncticu-

late-scaberulous.

150 FIELDIANA: BOTANY, VOLUME 24

Phyllanthus diffusus Klotzsch in Seem. Bot. Voy. Herald 105.

1857.

About pools in forest or in marshes, often in shallow water, at

or near sea level; British Honduras; Honduras; Panama; West

Indies; tropical South America.

Plants essentially annual but perhaps sometimes more enduring, erect, 50 cm.

high or less, the main stem thickened and somewhat fistulous at the base, branched

above, the branches very slender, terete, ascending or somewhat spreading, gla-

brous throughout; stipules small, subulate, scarcely broadened at the base; leaves

almost sessile, oblong to obovate or elliptic-oblong, mostly 3-6 mm. long and 2-3

mm. wide, rounded or obtuse at each end, pale green or glaucescent, the lateral

nerves inconspicuous; flowers monoecious, solitary or binate in the leaf axils, on

very short pedicels; staminate sepals 5-6, ovate or obovate; stamens 3, the fila-

ments connate; pistillate sepals 6, oblong or obovate, green, obtuse or subacute;

ovary smooth, the styles short, 2-cleft; capsule 2-3 mm. broad, depressed-globose;

seeds brown, with 5-6 obscure dorsal lines and numerous transverse ones on the

dorsal surface.

Phyllanthus ferax Standl. Field Mus. Bot. 11: 134. 1932.

Moist or dry, rocky, brushy places, on limestone, 300 meters or

less; Pete"n (type from Uaxactun, H. H. Bartlett 12157). Yucatan;

British Honduras.

A slender glabrous shrub a meter high or less, rather densely branched almost

throughout, the branches ferruginous, flexuous, wiry, hard, rather stiffly ascend-

ing, abundantly leafy; stipules attenuate from an ovate base; leaves almost sessile,

membranaceous, obovate-elliptic or oblong-elliptic, 1.5-3 cm. long, 8-15 mm. wide,

rounded at the apex, acute at the base, glaucescent beneath, the lateral nerves

inconspicuous; flowers monoecious, solitary or geminate, the pedicels 3 mm. long

or less; staminate sepals 6, oblong, obtuse, 1.5 mm. long; pistillate sepals 6, broadly

obovate, green, obtuse or rounded at the apex; capsule 3-celled, depressed-globose,

3-3.5 mm. broad; seeds dark brown, smooth, dull.

Phyllanthus Galeottianus Baill. was reported from Huehuetenango

by Loesener on the basis of Seler 2161 and 3000 from Chacula,

the determination by John Donnell Smith. The collections probably

are referable to one of the species listed here, but it is possible that

they do represent the Mexican P. Galeottianus.

Phyllanthus glaucescens HBK. Nov. Gen. & Sp. 2: 115. 1817.

P. laxiflorus Benth. PI. Hartweg. 89. 1842 (type from "Monte

Pineda" near Guatemala, Hartweg 612). Manzana de raton.

Moist thickets or mixed forest, 900-2,000 meters; reported from

Alta Verapaz; Jutiapa; Santa Rosa; Guatemala; Sacatepe"quez;

Huehuetenango. Mexico; Salvador.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 151

A glabrous shrub, generally 1.5-3 meters high, sometimes a tree of 9 meters,

the branches stout, ferruginous, subterete or somewhat angulate; stipules linear-

lanceolate from an ovate base, long-acuminate; leaves on petioles 8-10 mm. long,

rounded-elliptic to oblong-elliptic, mostly 9-20 cm. long and 5-11 cm. wide,

obtuse to acuminate, usually rounded at the base, deep green above, glaucous

beneath, with conspicuous nerves and veins, the areolae densely and minutely

venulose; flowers small, pale green, laxly paniculate, monoecious, very numerous,

on pedicels 5-10 mm. long; staminate flowers scarcely 2 mm. long; stamens united

to form a column, the anthers connate; staminate sepals oblong-elliptic, penni-

nerved, 2.5-4 mm. long, obtuse; styles connate only at the base; capsule 3-celled,

depressed-globose, about 5 cm. broad, green, with a thick fleshy pericarp when

fresh, this becoming hard when dry; seeds ovoid, brown, 1 cm. long, smooth,

pointed at one end, broadly rounded at the other, the raphe 5 mm. long.

The Maya name in Yucatan and British Honduras is "pixton";

"monkey rattle" (British Honduras). The shrub is rather common

in the central region, especially about Antigua. The fruits are very

different from those of most plants of the family, somewhat sugges-

tive of a small green apple, and one seeing them for the first time,

detached from the plant, is likely to have difficulty in determining

their relationship.

Phyllanthus grandifolius L. is reported from Pansamala, Alta

Verapaz, on the basis of Tuerckheim 1085. We have not seen material

of this species from continental North America, and the true determi-

nation of the plant is uncertain.

Phyllanthus lathyroides HBK. Nov. Gen. & Sp. 2: 110. 1817.

Sarin de suelo (fide Aguilar).

Moist or wet thickets or mixed forest, often on open banks, fre-

quently a weed in waste ground, especially in cafetales, 500-1,700

meters; Alta Verapaz; Zacapa; Chiquimula; Santa Rosa; Escuintla;

Guatemala; Sacatepe"quez; Chimaltenango; Suchitepe"quez; Retal-

huleu; Quezaltenango. Mexico; Salvador to Panama; West Indies;

tropical South America.

A slender annual, usually erect, 50 cm. high or less, glabrous, branched above

and sometimes from the base, the branches short, spreading, weak, terete or nearly

so; stipules linear-lanceolate; leaves almost sessile, distichous, oblong or oblong-

elliptic, mostly 6-14 mm. long and 2.5-6 mm. wide, rounded and obscurely apicu-

late at the apex, oblique and often subcordate at the base, pale green, glaucescent

beneath, the 5-6 pairs of lateral nerves rather conspicuous beneath; flowers

monoecious, the pistillate solitary, the staminate solitary or binate, the pedicels

5 mm. long or usually much shorter; staminate sepals 5, broadly ovate, obtuse;

stamens 3, connate below; pistillate sepals 5, oblong-obovate, somewhat venose;

ovary smooth, the 3 styles 2-cleft; capsule 3 mm. broad, depressed-globose, 3-

celled; seeds puncticulate-scabrous, the minute projections in numerous longitudinal

152 FIELDIANA: BOTANY, VOLUME 24

A curious and conspicuous but probably unimportant form of

this species is the following: Phyllanthus lathyroides f. decoratus

Standl. & Steyerm. (forma nova. A forma typica speciei non nisi

sepalis purpureis differt. Type, Juvenal Valeria Rodriguez 3576 from

Zamorano, Dept. Morazan, Honduras, in Herb. Chicago Nat. Hist.

Mus.). Apparently this is a local form, for search in the herbarium

has revealed only two other collections, one collected in Chiquimula

(pine forest, Socorro Mountain, Steyermark 30979), and one from

Siguatepeque, Honduras (T. G. Yuncker 5556). The purple-red

sepals must be conspicuous in the living plants, which ordinarily

are of a pale green throughout.

Phyllanthus longipes Steyermark in Standl. Field Mus. Bot.

22: 153. 1940.

Known only from the type, on hillside, El Cayo District, British

Honduras, P. H. Gentle 2619.

A glabrous tree, the trunk 12 cm. in diameter, the branches slender; leaves on

petioles 2.5-4 mm. long, chartaceous, oblong-elliptic or oblong-ovate, 4-8 cm.

long, 1-3.5 cm. wide, acuminate with an obtuse tip, acute at the base, glaucescent

beneath, the lateral nerves about 8 pairs; flowers apparently dioecious; pistillate

flowers axillary, fasciculate, the pedicels filiform, 4.5-5 cm. long; capsule 3-celled,

castaneous, smooth, 5 mm. long and broad, deeply emarginate; seeds castaneous,

smooth, 4 mm. long.

Noteworthy for the deeply emarginate capsule whose cocci are

definitely sulcate longitudinally.

Phyllanthus micrandrus Muell. Arg. Linnaea 32: 27. 1863.

Chabin-te (Huehuetenango).

Moist or dry thickets or open forest, often in pine-oak forest,

1,700 meters or less; Pete'n; Jalapa; Jutiapa; Chimaltenango; Suchi-

tepe"quez; Retalhuleu; Huehuetenango; Quezaltenango. Mexico;

Venezuela.

A branched shrub 1-2 meters high, glabrous throughout or nearly so, the

branches slender, terete, grayish or brownish, the young ones often somewhat

angulate; stipules triangular-ovate; leaves membranaceous or somewhat thicker,

on petioles 2-3 mm. long, orbicular to rounded-elliptic or ovate-orbicular, mostly

1.5-3 cm. long and 2-3 cm. wide, rounded at the apex or subemarginate, apiculate,

rounded or very obtuse at the base; flowers monoecious, fasciculate in the leaf

axils, the pedicels capillary, the pistillate ones thickened at the apex; calyx green,

2 mm. long, the sepals oblong-elliptic, obtuse; anthers free; styles short, 2-parted;

capsule subglobose, 3 mm. long; seeds 2 mm. long, fuscous, minutely and densely

puncticulate.

Some of the Guatemalan collections are noteworthy in having

the leaves pubescent beneath, and the branches and pedicels also

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 153

may be pubescent. Possibly more than a single species is represented

by the specimens.

Phyllanthus minarum Standl. & Steyerm. Field Mus. Bot.

23: 125. 1944.

Known only from the type, El Progreso, Sierra de las Minas,

hills between Finca Piamonte and slopes southeast of the finca,

2,400-2,500 meters, Steyermark 43385.

A shrub 1.5-2.5 meters high, the older branches terete, ochraceous, roughened

by the persistent indurate stipules, about 12 mm. thick, bearing numerous very

large scars left by leaves and branches, the young branches with their numerous

leaves simulating a large pinnate leaf, only 2-3 mm. thick, white- villosulous;

stipules more or less persistent, ferruginous, in age indurate, at first often reflexed,

ovate-triangular or lance-triangular, as much as 8 mm. long, acuminate, sub-

cordate at the base; lowest leaves suborbicular or rounded-deltoid and 1.5-3 cm.

long, the principal leaves oblong-ovate or deltoid-oblong, 4.5-7 cm. long, 3 cm.

wide, obtuse or subacute at the apex and conspicuously cucullate, broadly rounded

at the base or the lowest leaves subcordate, membranaceous, on very short petioles,

green above, puberulent on the nerves, pale beneath, densely pilose with hyaline

hairs; flowers probably dioecious, the staminate pedicels capillary, flexuous, 2-2.5

cm. long, mostly solitary in the leaf axils, densely pilose; sepals 6, very unequal,

the inner ones 5.5 mm. long, oblong-oval, very obtuse, the outer ones 3.5 mm. long,

elliptic or ovate, subacute; glands free, about equaling the stamen column, broadly

ovate; stamens 3, the filaments connate into a short column, the anthers short,

coherent.

The cucullate leaf apices in this plant are curious, but it is un-

certain whether they constitute a natural character or are an

abnormality.

Phyllanthus Niruri L. Sp. PI. 981. 1753. Moco coquillo (fide

Aguilar).

Moist or wet thickets or fields, sometimes on open or rocky

banks, in cultivated ground, or rarely in forest, 1,600 meters or less,

mostly at very low elevations; Alta Verapaz; Izabal; Santa Rosa;

Escuintla; San Marcos. Mexico; British Honduras to Salvador and

Panama; West Indies; tropical South America; Old World tropics.

A slender glabrous annual 50 cm. high or less, usually erect, generally branched,

the branches weak and spreading or even somewhat pendent, terete or obscurely

angulate; leaves almost sessile, oblong or obovate-oblong, mostly 6-15 mm. long

and 2-6 mm. wide, rounded at each end, membranaceous, the lateral nerves ob-

scure; stipules subulate, with a broad triangular base; flowers monoecious, greenish

white, solitary, or one of each sex in the same axil, on very short pedicels; staminate

sepals 5-6, ovate or obovate; glands of the disk 5-6, small; stamens 3, the fila-

ments connate; pistillate pedicels 1-2 mm. long, the 6 sepals oblong; ovary smooth;

154 FIELDIANA: BOTANY, VOLUME 24

capsule 1.5-2 mm. in diameter, depressed-globose, 3-sulcate; seeds with 5-6 very

fine and inconspicuous longitudinal lines on the dorsal surface.

Both this and P. lathyroides are common weeds through most of

the lowlands of Central America, at least in moist or wet regions.

They are inconspicuous plants and seldom noticed.

Phyllanthus nobilis (L. f.) Muell. Arg. in DC. Prodr. 15, pt.

2: 414. 1866. Margaritaria nobilis L. f. Suppl. PI. 4298. 1781.

Cicca antillana Juss. Euphorb. Tent. pi. 4, f- 13 B. 1824. P. nobilis

var. antillanus Muell. Arg. op. cit. 415. P. nobilis var. hypomalacus

Standl. Carnegie Inst. Wash. Publ. 461: 68. 1935. Mapahuite

(Huehuetenango; probably an erroneous name).

Moist or dry thickets or thin forest, on plains or hillsides, often

in second growth, 900 meters or lower; Pete"n; Alta Verapaz; Chi-

quimula; Jutiapa; Escuintla; Suchitepe"quez; Solola; Huehuete-

nango; probably in all the Pacific coast departments. Mexico;

British Honduras to Salvador and Panama; West Indies; tropical

South America.

A shrub or small tree, sometimes as much as 15 meters high but usually 7

meters or less, the trunk slender, often crooked, as much as 15 cm. in diameter,

the crown small and narrow, the bark dark brown, thin, the inner bark deep pink,

the slender branches glabrous or pilosulous; leaves on very short petioles, mem-

branaceous, elliptic or lance-elliptic, mostly 6-13 cm. long and 2.5-5 cm. wide,

acuminate or cuspidate-acuminate, acute or subacute at the base, mostly glabrous,

sometimes puberulent or pilose beneath, green or glaucescent beneath; flowers

dioecious, greenish white, the staminate fasciculate along a short peduncle, the

pistillate solitary or 2-4-nate in the leaf axils; staminate pedicels slender, 3-5 mm.

long, the 4 sepals 2-seriate, rounded or elliptic; stamens 4, the filaments free;

pistillate pedicels rather stout, 10-15 mm. long; sepals 4; ovary usually 4-celled,

the styles thick, united below, shortly 2-cleft; capsule about 1 cm. in diameter,

green, subglobose, the pericarp at first fleshy, in age dry and dehiscent; seeds mostly

4, complanate, trigonous, olivaceous, 3 mm. long.

Called "clawberry" and "ramon macho" in British Honduras;

"nistamal" (Salvador); "icinche" (British Honduras, Maya);

"xnabalche" (Yucatan, Maya). This is a common weedy shrub in

many parts of the Guatemalan lowlands and elsewhere in Central

America. The plants are leafless during the dry season. The sap-

wood is pale yellow, the heartwood brownish, darkening upon ex-

posure, sometimes with a pinkish tinge or almost black. It is little

used unless for firewood and minor construction purposes. In the

typical form the leaves are glabrous; in var. hypomalacus they are

sparsely or densely pubescent beneath. The variety is widely dis-

tributed with the typical form, and probably is of only minor im-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 155

portance. The fresh seeds are somewhat fleshy and colored dark

blue.

Phyllanthus Purpusii Brandeg. Univ. Calif. Publ. Bot. 6: 55.

1914.

Steep brushy slopes or in moist mixed forest, 2,100-2,700 meters;

Solola; Suchitepe'quez; San Marcos. Chiapas, the type from Cerro

del Boqueron.

An erect stout shrub 1.5-3.5 meters high, the main trunk simple or with a

few stout branches 1 cm. thick or more, bearing at the apex crowded slender

leafy branches that resemble pinnate leaves and are deciduous like leaves, the

main stems roughened by the large persistent indurate stipules, the young branches

subterete, minutely puberulent, often reddish; leaves crowded and very numerous,

almost sessile, oblong-lanceolate, 2-7.5 cm. long, 8-22 mm. wide, long-acuminate

or attenuate to a very acute tip, somewhat oblique at the base and rounded or

very obtuse, glabrous, membranaceous, usually glaucous beneath; stipules lanceo-

late, longer than the petioles, often reflexed; flowers probably dioecious, the

staminate pedicels 4-6 mm. long, the sepals unequal, the larger ones lanceolate,

5 mm. long, the smaller ones less than half as long; filaments united, the anthers

coherent; pistillate pedicels about 2 cm. long, the sepals lanceolate or lance-ovate,

as much as 5 mm. long; styles connate only at the base, short-bifid; capsule sub-

globose, 3-celled, shallowly sulcate, 5 mm. in diameter, smooth; seeds brown,

minutely rugulose.

This shrub is common on Volcan de Santa Clara (Suchitepe'quez),

mostly at about 2,400 meters.

Phyllanthus Urinaria L. Sp. PI. 982. 1753.

Moist thickets or wet fields, at or near sea level; Izabal; perhaps

introduced in Central America. British Honduras; Atlantic coast

of Honduras; northern South America; Old World tropics.

Plants annual, erect or nearly so, glabrous, 50 cm. high or less, often much-

branched, the lateral branches short, rather weak, spreading or ascending, terete,

sometimes minutely hispidulous; leaves almost sessile, oblong, 5-18 mm. long,

2-4 mm. wide, rounded to acute and mucronulate at the apex, rounded at the base,

membranaceous, glaucescent beneath, minutely ciliolate, the lateral nerves 5-6

pairs, prominent beneath; stipules broad, auriculate, acuminate; flowers monoe-

cious, solitary, subsessile, the pistillate on the lower part of the branchlet, the

staminate above; staminate flowers very small, the 6 sepals obovate-elliptic, green;

stamens 3, the filaments connate; pistillate sepals 6, oblong, white-marginate,

persistent and reflexed in fruit; ovary densely and minutely verruculose, the

short styles 2-fid; capsule 2 mm. in diameter, depressed-globose; seeds conspicu-

ously transverse-rugose.

PLUKENETIA L.

Reference: F. Pax, Pflanzenreich IV. 147, ix: 12-17. 1919.

156 FIELDIANA: BOTANY, VOLUME 24

Woody vines; leaves alternate, petiolate, 2-stipulate, 3-nerved or penninerved;

flowers monoecious, apetalous, the inflorescences spike-like, with 1-2 pistillate

flowers at the base, the upper flowers staminate, the staminate bracts subtending

a few-flowered branchlet; staminate calyx in bud globose, in anthesis valvately

4-parted; stamens 12-30, inserted on a conic receptacle, the filaments free, rather

stout, the anthers terminal, sometimes more or less cruciately 4-celled; glands of

the staminal disk few and minute or none; sepals 4, small; ovary 4-celled, 4- winged,

the styles connate into an elongate column; stigmas short, entire or 2-lobate;

ovules solitary in each cell; capsule large or medium-sized, dehiscent, the cocci

carinate or appendaged; seeds lentiform or globose, not carunculate; cotyledons

ovate, 3-nerved at the base.

About 7 species, in tropical America. One other is known from

southern Central America (Panama).

Plukenetia penninervia Muell. Arg. Linnaea 34: 158. 1865.

P. angustifolia Standl. Field Mus. Bot. 4: 314. 1929 (type from

Lancetilla Valley near Tela, Honduras).

Moist or wet thickets, often in open pine forest, 350 meters or

less; Alta Verapaz; Izabal; Retalhuleu. Southern Mexico; British

Honduras; Honduras; Venezuela.

A small woody vine, twining, sparsely puberulent on the younger parts but

in age glabrous or nearly so outside the inflorescence; leaves on petioles about 1 cm.

long, chartaceous or almost coriaceous, often very lustrous, oblong-elliptic to lance-

oblong, acuminate or cuspidate-acuminate, rounded to broadly cuneate at the

base, with 2 conspicuous glands on the upper surface at the base, penninerved,

denticulate or almost entire; racemes sessile, short, few-flowered, hispidulous, the

flowers very small, green; stamens 30 or fewer; capsule depressed-globose, deeply

4-lobate, 1-1.5 cm. broad, green, glabrous, the cocci carinate dorsally and obtusely

tuberculate at about the middle, the style persistent, thick, 1.5 mm. long.

The Central American material exhibits some variation, but not

more than might be expected within a species. It is possible that

when more South American material is available for comparison,

P. angustifolia may be found tenable, since at present it seems to

be isolated geographically from P. penninervia, but so far as can be

determined at present, the two names are synonymous.

RICINUSL. Castor bean

Tall annuals, herbaceous or becoming somewhat woody, often persisting for

more than a single season, glabrous; leaves alternate, long-petiolate, peltate, pal-

mately lobate, the lobes dentate; stipules large, united, covering the buds; flowers

monoecious, apetalous, in paniculate racemes at the ends of the branches, the

lower flowers staminate, the upper ones pistillate, short-pedicellate; disk none;

staminate calyx globose in bud, 3-5-valvate in anthesis; stamens numerous, the

filaments repeatedly branched; anther cells subglobose, divaricate, attached

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 157

separately to the connective; pistillate calyx spathaceously cleft, caducous; ovary

3-celled, the style spreading, generally 2-cleft; ovules 1 in each cell; capsule splitting

into 3 bivalvate cocci, usually echinate, sometimes smooth; seeds large, carunculate;

endosperm carnose.

The genus consists of a single species.

Ricinus communis L. Sp. PI. 1007. 1753. Higuerillo; Higuerillo

bianco; Higuerillo rojo; Aceite (Coban); Ixcoch (Pete"n, Maya);

Raxten (Quiche").

Planted commonly and also thoroughly established and often

abundant in thickets on plains and hillsides, often in open places,

hedges, or about cultivated ground, sometimes plentiful along

streams; found from the coasts up to the limit of cultivation, but

most plentiful at low elevations; native of the Old World tropics,

perhaps of Africa, but now found in all tropical regions; Pete*n;

Alta Verapaz; Izabal; El Progreso; Zacapa; Chiquimula; Jutiapa;

Santa Rosa; Escuintla; Guatemala; Suchitepe'quez; Retalhuleu;

Quezaltenango; San Marcos; Huehuetenango; doubtless found in

all the departments.

A coarse stout erect herb or often tree-like and as much as 6 meters high, with

a thick trunk, the plants pale and glaucous or often tinged with red or purple;

petioles often equaling or exceeding the leaf blades; blades almost orbicular in

outline, 10-60 cm. broad, deeply palmate-lobate, the lobes ovate-oblong or lanceo-

late, acute or acuminate, irregularly glandular-dentate; staminate calyx 6-12 mm.

long, the pistillate ones 4-8 mm. long; ovary densely fleshy-tuberculate; capsule

1.5-2.5 cm. long, oval, densely echinate; seeds ellipsoid, somewhat compressed,

10-17 mm. long, smooth, mottled and highly variable in color, or entirely black,

conspicuously carunculate.

The Maya name in Yucatan is "coch" or "xcoch." In Guate-

mala, as well as elsewhere in Central America, there are recognized

two common varieties, the bianco and the rojo, Colorado, negro, or

morado. In the former the stems and leaves are rather pale green,

in the latter brilliantly tinged with red or purple. On Volcan de

Tajumulco two types are found: the large black-seeded kind is used

for lubricating oil for machinery, while the smaller, mottled light

and dark brown variety is used for medicinal castor oil.

Occasional plants are dark red or dark purple throughout, and

consequently very conspicuous. The two forms sometimes grow

together, but often one or the other prevails in a region. The plant

is important economically as the source of castor oil — "aceite de

ricino," "aceite de castor," or "aceite de palma-Cristi," and it is

grown on a small scale in Guatemala on this account. Some castor

oil is imported into Guatemala, but much is extracted locally in

158 FIELD IANA: BOTANY, VOLUME 24

hand presses, especially at Antigua. Its best-known use is as a

purgative medicine, but it is highly esteemed as a lubricant, in soap

manufacture, in dyeing and printing cotton goods, and for dressing

tanned hides. In India and China silkworms are fed on the leaves.

The stems have been utilized for paper making, and the ground seeds

from which the oil has been separated are a valuable fertilizer. In

Guatemala the oil is sometimes used for illumination, especially in

churches. It is placed on food exposed for the purpose of poisoning

cockroaches. It is applied by veterinarians to wounds in stock, and

employed also to give luster to the hair of people. The leaves

moistened with vinegar are applied as poultices to relieve headache,

and the oil mixed with turpentine is sometimes administered to

expel tapeworms.

SAPIUM P. Browne

Reference: F. Pax, Pflanzenreich IV. 147, v: 199-258. 1912.

Large trees or shrubs, glabrous, with copious white latex; leaves alternate,

petiolate, entire or serrate, often glandular, the apex often glandular and with a

cucullate inflexed tip, the petiole or the base of the blade usually bearing 2-4

large glands; flowers spicate, monoecious, the spikes terminal, staminate or with

pistillate flowers only at the base, the bracts usually with glands on each side of

the base; flowers apetalous, the disk wanting; staminate calyx usually 2-fid; ovary

generally 3-celled, the cells 1-ovulate; styles free or connate at the base; fruit

capsular, globose, pyriform, or trigonous; seeds subglobose, not carunculate.

About 100 species, in tropical regions of both hemispheres. A

substantial number of other species is found in southern Central

America. Some of the South American species are or have been of

some commercial importance as a source of rubber. The woods are

light and soft to moderately so, whitish or yellowish, staining readily,

easy to work, sawing woolly; they are suitable for packing boxes,

interior construction, and paper pulp. The available Guatemalan

material of this genus is insufficient and unsatisfactory for study.

The species are difficult of separation, being distinguished by minor

characters whose value is questionable. It is probable that the

number of recognized species is greater than the facts warrant.

Until much ampler and better material has been collected, it will be

impossible to decide just how many are represented in Central

America. The names "chilicuate," "amate de hule," and "cuxchon-

quic" are reported from Guatemala for trees of this genus whose

specific identity is uncertain.

Tip of the leaf blade strongly recurved against the upper surface of the blade.

S. Schippii.

Tip of the leaf blade flat, not recurved.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 159

Petioles short, only 5-8 mm. long S. Tuerckheimianum.

Petioles elongate, mostly 2-3 cm. long or often much longer.

Principal lateral nerves of the leaves mostly 6-10 pairs, remote. .S. nitidum.

Principal lateral nerves of the leaves usually much more numerous and close

together.

Lateral nerves of the leaves divergent at an acute angle, or at least strongly

ascending; spikes mostly or all lateral S. lateriflorum.

Lateral nerves of the leaves divergent at almost a right angle; spikes, at

least the fertile ones, terminal S. macrocarpum.

Sapium lateriflorum Hemsl. in Hook. Icon, sub pi. 2680. 1901.

Chilamate.

Wet to dry forest, often along stream banks, 1,300 meters or less;

Alta Verapaz; Jutiapa; Escuintla; Guatemala; Huehuetenango.

Southern Mexico.

A small or large tree, often 15 meters high or more, glabrous throughout,

the bark grayish or whitish, the crown spreading, dense, the branches thick,

subterete; leaves on petioles 1.5-4 cm. long, the petiole bearing at the apex 2 small

conic glands; leaf blades oblanceolate-oblong or elliptic-oblong, mostly 10-20 cm.

long and 4-8 cm. wide, on fruiting branches often smaller, obtuse or acute at the

apex, the tip flat or nearly so and obtuse, cuneate or obtuse at the base, coriaceous

or subcoriaceous, often lustrous, obscurely denticulate or subentire, the lateral

nerves usually 11-17, slightly curved, divergent or ascending at an angle of little

more than 45 degrees; stipules auriculiform, very small, persistent; flowers dioe-

cious, the staminate spikes slender, 7-9 cm. long, naked at the base, the bracts small,

reniform-ovate, denticulate, with a peltate gland on each side at the base, 5-9-

flowered; stamens 2, exserted; pistillate spikes axillary or sometimes perhaps

terminal, rather few-flowered; capsule borne on a stout pedicel 8 mm. long or

shorter, about 1.5 cm. long, very thick and hard; seeds 6-7 mm. long.

In Oaxaca called "palo de la flecha," probably because the sap

was used for poisoning arrows; "amatillo" (Veracruz). This perhaps

is the species that has been reported from Guatemala as S. bi-

glandulosum var. Klotzschianum Muell. Arg. All the local species

are much alike, doubtless have the same properties, and no distinc-

tion between them is made by the people. All are commonly called

"chilamate." In Guatemala, Salvador, and Honduras the trees have

the reputation of being highly poisonous, the copious milky latex

causing blisters and inflammation upon the skin. On this account

they often are left when land is cleared. In Panama, on the other

hand, boys sometimes collect and coagulate the latex, then chew it

to prepare bird lime. It is possible that different species differ in

their poisonous properties. In the Oriente of Guatemala the latex

is used as a barbasco or fish poison.

160 FIELDIANA: BOTANY, VOLUME 24

Sapiuni macrocarpum Muell. Arg. Linnaea 32: 119. 1863.

S. mexicanum Hemsl. in Hook. Icon. pi. 2680. 1901. Matapalo

(probably an erroneous name); Chilamate; Higuerillo.

Wet to dry forest, often in open fields, frequent along streams,

1,500 meters or less, most common at low elevations; Alta Verapaz;

Jutiapa; Santa Rosa; Escuintla; Sacatepe"quez; Solola; San Marcos.

Southern Mexico; Salvador.

A small to large, glabrous tree, sometimes 25 meters high, the branches thick,

often brown or brownish; petioles mostly 2.5-3 cm. long, sometimes longer, bearing

at or near the apex 2 thick subglobose glands; leaf blades oblong-lanceolate or

oblong, mostly 10-20 cm. long and 3-6 cm. wide but often larger, obtuse or

abruptly acute at the apex, the tip flat, eglandular, obtuse, cuneate or rounded

at the base, minutely callous-denticulate, subcoriaceous or coriaceous, the lateral

nerves very numerous, divaricate at almost a right angle, curved toward the

margin of the blade; stipules scale-like, persistent, small; spikes commonly terminal

and solitary, 10-14 cm. long, androgynous, bearing 3-4 pistillate flowers at the

base; bracts small, very broadly ovate, entire, with an oval gland at the base on

each side, the staminate bracts 9-12-flowered; staminate calyx 2-lobate, the

pistillate 3-parted; stamens 2; capsule on a very short pedicel, subglobose, ligneous,

when opened 3.5 cm. broad; seeds ovoid, acute, almost 1 cm. long, the aril bright

red.

Sapium nitidum (Monachino) Lundell, Amer. Midi. Nat. 29:

477. 1943. S. biglandulosum var. nitidum Monachino, Bull. Torrey

Club 67: 771. 1940. Palo de tuerto (Huehuetenango) ; Amate (Hue-

huetenango, probably an erroneous name).

Moist or wet forest, sometimes in swampy places, 1,700 meters

or less; Pete*n; Izabal; Solola; Huehuetenango. British Honduras

(type from Santa Rosa pasture near El Cayo, J. B. Kinloch 340) ;

Atlantic coast of Honduras.

A tree 9-18 meters high, glabrous throughout, the bark thick, light gray,

smooth or nearly so, the branches slender or stout, the older ones brown; stipules

small, ovate to oblong, persistent; petioles mostly 1.5-3.5 cm. long, bearing 2

short but conspicuous glands at or near the apex; leaf blades obovate to oblong

or elliptic-oblong, mostly 8-18 cm. long, obtuse or subacute, the tip flat, obtuse,

acute to rounded at the base, obscurely denticulate or practically entire, charta-

ceous or subcoriaceous, usually lustrous, the lateral nerves remote, commonly 6-10

pairs, arcuate and ascending; spikes terminal, solitary, the staminate slender, as

much as 15 cm. long; pistillate spikes many-flowered, short, producing numerous

capsules; capsules borne on short thick pedicels, about 8 mm. in diameter, hard

and ligneous, 2-3-celled; seeds 6 mm. long and broad, the aril red.

This has been reported from British Honduras as S. jamaicense

Swartz, a species of doubtful occurrence in northern Central

America. Called "leche Maria" in British Honduras.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 161

Sapium Schippii Croizat in Lundell, Amer. Midi. Nat. 29: 477.

1943.

Known only from the type, Forest Home, Toledo District, British

Honduras, near sea level, W. A. Schipp 1049.

A tree of 18 meters, the trunk 45 cm. in diameter; petioles slender, 2-4 cm.

long, bearing at the apex 2 stout-pedicellate, very conspicuous glands; leaf blades

elliptic or elliptic-oblong, 6-11 cm. long, 2-4.5 cm. wide, rounded or very obtuse

at the base, rounded and abruptly tipped at the apex, the tip bearing a conspicuous

gland, recurved upon the upper surface of the blade, subentire, the lateral nerves

about 15 pairs, arcuate, divergent at a broad angle; capsule subglobose, ligneous,

9 mm. long, borne on a stouf pedicel 6-8 mm. long; seeds 7 mm. long, 5 mm. broad,

surrounded by a red aril.

Sapium Tuerckheimianum Pax & Hoffm. Pflanzenreich IV.

147, xiv: 61. 1919.

Known only from the type, Cubilgiiitz, Alta Verapaz, 350 meters,

Tuerckheim 11.941.

A glabrous tree with slender branches; petioles short, 5-8 mm. long, eglandular;

leaf blades lanceolate or oblanceolate, 8-11 cm. long, 2.5-3.5 cm. wide, caudate-

acuminate, with a flat tip, cuneate-acute at the base and bearing at the very base

on the upper surface 2 patelliform glands, subcoriaceous, entire or obscurely

denticulate, the lateral nerves about 15 pairs, curved, ascending at an angle of

about 45 degrees; stipules very small, deltoid-auriculiform, caducous; flowers

dioecious, the staminate spikes axillary, 4 cm. long, the bracts small, reniform-

ovate, acute, with an oblong gland at the base on each side, 5-9-flowered; staminate

calyx shallowly 2-lobate; stamens 2 or rarely 3.

We have seen no material of this species.

SEBASTIANIA Sprengel

Reference: F. Pax, Pflanzenreich IV. 147, v: 88-153. 1912.

Mostly shrubs or trees; leaves alternate, short-petiolate, often more or less

coriaceous, penninerved, serrulate or rarely entire, the stipules small; flowers

generally monoecious, apetalous, spicate, the spikes slender, terminal on leafy

branchlets or opposite the leaves, rarely axillary, the bracts 2-glandular at the

base; disk none; staminate flowers very small, several or solitary within the bracts,

sessile or subsessile; calyx small, open before anthesis, usually 3-lobate or 3-parted;

stamens commonly 3, the filaments free or connate at the base, the anthers longi-

tudinally dehiscent; pistillate flowers few or solitary at the base of the spike or

few in distinct spikes; sepals 3, the ovary usually 3-celled, the styles spreading

or revolute, simple, free or rarely connate; ovules solitary in each cell; capsule

tridymous or subglobose, smooth or tuberculate, the 2-valvate cocci separating

from a central columella; endocarp crustaceous; seeds globose to oblong or cylindric,

carunculate, smooth; endosperm carnose, the cotyledons broad, flat.

162 FIELDIANA: BOTANY, VOLUME 24

Species about 75, mostly in Brazil, 3 in the Old World tropics.

One other Central American species, a small annual, occurs in

Panama.

Bracts of the staminate flowers short-stipitate; lowest teeth of the leaves not

glandular S. adenophora.

Bracts of the staminate flowers closely sessile; lowest teeth of the leaves glandular.

Staminate flowers usually 3 in each bract, the stamens 2-3; capsule generally

not more than 6 mm. long and 10 mm. broad; leaves acuminate or narrow-

acuminate : S. confusa.

Staminate flowers usually 6-9 in each bract, the stamens 2-6; capsule generally

8-11 mm. long and 10-13 mm. broad; leaves narrowly long-caudate.

S. longicuspis.

Sebastiania adenophora Pax & Hoffm. Pflanzenreich IV. 147,

v: 145. 1912.

Moist or dry forest, at or little above sea level; British Honduras;

Yucatan Peninsula of Mexico.

A glabrous tree 5-7 meters high, the trunk 10 cm. or less in diameter; leaves

on petioles 3-8 mm. long, chartaceous, oblong-ovate to elliptic-oblong, 4-8 cm.

long, 1.5-3 cm. wide, acuminate, obtuse or rounded at the base, this 2-glandular

and revolute, the margin finely serrulate; flower spikes terminal on short leafy

shoots, bisexual or unisexual, 3-5.5 cm. long, or the pistillate ones shorter, the

bisexual ones with 1-3 pistillate flowers at the base; staminate flowers mostly 5 to

each bract, the bracts short-stipitate, 2-glandular; sepals lanceolate or ovate;

stamens mostly 2, sometimes as many as 5; capsule smooth, 5-6 mm. long, 6-7

mm. broad.

"Chechem bianco," "sacchechem," "canchunup" (Yucatan,

Maya). All the local species of this genus are similar in general

appearance and their supposedly distinctive characters remain to

be confirmed. The milky latex is reported as highly poisonous in

contact with the skin, causing irritation and eruption after the fashion

of Rhus radicans.

Sebastiania confusa Lundell, Lloydia 2: 99. 1939.

Dry or moist forest, often on limestone 1,650 meters or less;

Pete"n; Jalapa; Huehuetenango. Tabasco; British Honduras.

A glabrous shrub or tree 1-15 meters high with slender branches, the trunk

20 cm. or less in diameter; leaves firm-membranaceous, lustrous, on slender petioles

4-10 mm. long, lance-oblong to oblong-elliptic, 5-11 cm. long, 2-3.5 cm. wide,

acuminate, usually abruptly so, rounded at the base, finely serrate, the basal

teeth glandular, the lateral nerves 10-14 pairs; stipules small, ovate; flower spikes

terminal on short leafy branchlets, unisexual or bisexual; staminate bracts sessile,

broadly ovate, 2-glandular at the base, 3-flowered, the flowers short-pedicellate;

sepals 3, ovate, erose; stamens usually 3; capsule smooth, 6 mm. long and 10 mm.

broad or smaller; seeds globose, 3 mm. in diameter.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 163

Called "white poison-wood" in British Honduras; "chitze'n"

(Tabasco). In this genus the heartwood is olive and variegated,

the sap wood white; not very attractive, of medium density, fine-

textured, easy to work, not highly durable.

Sebastiania longicuspis Standl. Field Mus. Bot. 11: 134. 1932.

S. Standleyana Lundell, Lloydia 2: 97. 1939 (type collected near

Vaca, British Honduras, P. H. Gentle 2544). Chechen (Izabal);

Chechem bianco, Icicheh (Pete*n, Maya).

Wet or dry forest, on plains or hillsides, often in second growth,

generally on limestone, 350 meters or less; Pete'n; Alta Verapaz;

Izabal. British Honduras.

A shrub or a tree, sometimes 40 meters high with a trunk 45 cm. in diameter,

glabrous, the trunk straight, the bark smooth, gray; leaves lustrous, membrana-

ceous or chartaceous, on slender petioles 7-1 5 mm. long, narrowly oblong to oblanceo-

late-oblong, mostly 7-14 cm. long and 2.5-4 cm. wide, abruptly long-caudate,

obtuse or subacute at the base, finely and irregularly serrulate, with glandular

teeth near the base, the slender veins prominulous on both surfaces; flower spikes

terminal on short leafy branchlets, mostly bisexual, with 1-2 pistillate flowers at

the base; staminate flowers mostly 6-9 in each bract, the bracts sessile, lunate,

erose, 2-glandular; calyx cupular, laciniate-dentate; stamens 2-6; capsule smooth,

8-11 mm. long, 10-13 mm. wide; seeds subglobose, 4 mm. long.

Known in British Honduras by the names "reventadillo,"

"white poison," "poison-wood," and "ridge white poison-wood."

The specific name was discarded by Lundell (loc. cit.) but reduced

to synonymy under S. Standleyana, because the type specimen

consisted in part of undetermined detached fruits. The specific

name, however, clearly pertains to the ample leaf material, whose

identity is not questionable. There is no need for the superfluous

second name for the tree, since there is no confusion as to its applica-

tion.

STILLINGIA Garden

Reference: F. Pax, Pflanzenreich IV. 147, v: 180-199. 1912.

Shrubs or perennial herbs, glabrous; leaves alternate or opposite, short-

petiolate, glandular-serrate, often 2-glandular at the apex of the petiole, mem-

branaceous to coriaceous, 2-stipulate; flowers monoecious, apetalous, in terminal

or rarely axillary spikes, simple, the bracts small and broad, 2-glandular at the

base; staminate flowers several or solitary within the bract, subsessile, the pistil-

late flowers solitary in the lowest bracts, or the spikes often wholly staminate;

disk none; staminate calyx small, shallowly and broadly 2-lobate; stamens generally

2, the filaments free, exserted, the anther cells ovoid-globose, longitudinally

dehiscent; pistillate calyx 3-parted or rudimentary or none; ovary 2-3-celled, the

164 FIELDIANA: BOTANY, VOLUME 24

style short-connate, undivided; ovules solitary in each cell; capsule usually tridy-

mous, separating into 2-valvate cocci, the base of the pericarp persistent between

the cocci, leaving a 3-radiate receptacle, the columella more or less winged, persis-

tent after dehiscence of the capsule, sometimes rudimentary or none; seeds sub-

globose, carunculate, the testa crustaceous; endosperm carnose, the cotyledons

broad, flat.

About 25 species, mostly in tropical America, a few in the Old

World and in temperate North America. One other species has been

described from Panama.

Leaves ovate, elliptic-ovate, or oblong-ovate, rounded at the base; large shrub or

small tree S. cruenta.

Leaves mostly lanceolate or oblong-lanceolate, generally acute at the base; low

shrubs, mostly 1-1.5 meters high.

Leaves small, mostly 3 cm. long or less S. sanguinolenta.

Leaves larger, mostly 5-10 cm. long.

Staminate bracts 1-flowered, the spikes very slender S. acutifolia.

Staminate bracts several-flowered, the spikes stout and thick. .S. zelayensis.

Stillingia acutifolia Benth. ex Hemsl. Biol. Centr. Amer. Bot.

3: 135. 1883. Sapium acutifolium Benth. PI. Hartweg. 90. 1842.

Stillingia propria Brandeg. Univ. Calif. Publ. Bot. 6: 185. 1915 (type

from Cerro del Boqueron, Chiapas). Hierba mala.

Moist or usually dry thickets or forest, often in rocky places or

in pine-oak forest, 900-3,000 meters; Chiquimula; Jalapa; Santa

Rosa; Guatemala; Sacatepe"quez; Chimaltenango; Solola (type from

Hacienda de Argueta, Hartweg 614); Totonicapan; Quezaltenango;

Huehuetenango; San Marcos. Chiapas.

A glabrous shrub, usually 1-3 meters high, rarely a tree of 6 meters, sparsely

or densely branched, the older branches brown or fuscous, terete; leaves alternate,

on stout petioles 3-5 mm. long, ovate-lanceolate to narrowly lanceolate or oblong-

lanceolate, mostly 5-14 cm. long and 1-4 cm. wide, narrowly long-acuminate,

acute or attenuate at the base, closely and acutely serrate, thick-membranaceous,

eglandular at the base, somewhat paler beneath; flower spikes terminal, about 4

cm. long, slender, the bracts broadly triangular, cuspidate-acuminate, spreading

in age, 1-flowered; Staminate flowers short-pedicellate; Staminate calyx 2-lobate,

the pistillate calyx obsolete; stamens 2; capsule scarcely 5 mm. long, the receptacle

3 mm. broad, the columella persistent; seeds not carunculate, 3.5 mm. long, gray.

Called "pavil" in Chiapas. The shrub has been reported from

Guatemala as S. aquatica Chapm. It is abundant in many localities

in the central and western highlands, often forming thickets. The

plants apparently are not browsed by sheep or goats, and people

leave them alone because, as the local name indicates, the copious

milky sap is believed to be irritating and poisonous to the skin.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 165

Stillingia cruenta Standl. & Steyerm. Field Mus. Bot. 23: 125.

1944.

Known only from the vicinity of the type locality, Santa Rosa,

Baja Verapaz, 1,500 meters, on dry brushy rocky hillsides (type,

Standley 91207).

A glabrous shrub or tree 2-8 meters high, densely branched, the branches

grayish brown, densely lenticellate; leaves on slender petioles 7-14 mm. long,

coriaceous or chartaceous, elliptic-ovate or oblong-ovate, 4-8 cm. long, 2-3.5 cm.

wide, long-acuminate, rounded at the base, lustrous, closely appressed-serrulate,

the margins with 2 large glands on each side at the base, paler beneath, the lateral

nerves about 6 pairs, prominulous, arcuate; perfect inflorescences not seen, terminal,

the rachis thick, the bracts in age spreading and indurate; capsule globose, smooth,

scarcely sulcate, about 7 mm. high and broad.

The tree is abundant at the only known locality, and during the

dry season is conspicuous because of the brilliant red coloring of the

leaves. Among Guatemalan species this is recognizable by the

relatively short and broad leaves and by its large size, the other

species being normally low shrubs.

Mr. J. D. Rogers, who has just completed a revision of the genus

in North and Central America, considers this species more properly

placed under Sapium.

Stillingia sanguinolenta Muell. Arg. Linnaea 32: 88. 1863.

At 800-1,200 meters, sometimes on limestone; Quiche"; Huehue-

tenango. Mexico; Honduras.

A stiff shrub, usually about a meter high, often densely branched, the branch-

lets sometimes reddish; leaves opposite and subalternate, on petioles 1-3 mm.

long, oblong-lanceolate or linear-lanceolate, 2-5 cm. long (very small in Guate-

malan material), 5-15 mm. wide, acuminate, acute at the base, serrate, mem-

branaceous or chartaceous; spikes 5-6 cm. long or often much shorter, straight

and stiff, the bracts broadly ovate, short-acute, the staminate 6-10-flowered, the

flowers subsessile; staminate calyx shallowly 2-lobate; pistillate sepals 3, broadly

ovate, denticulate; stamens 2; ovary 6-carinate at the apex, the styles connate

only at the base; capsule obovoid, subtruncate at the apex, apiculate, subacute

at the base, smooth, the receptacle 5-7 mm. broad; seeds 5-6 mm. long, grayish

white.

In Guatemalan specimens the leaves are sometimes less than

2 cm. long and the flower spikes very short and dense.

Mr. J. D. Rogers has placed the Guatemalan material as a new

species to be described in his forthcoming publication.

Stillingia zelayensis (HBK.) Muell. Arg. Linnaea 32: 87.

1863. Sapium zelayense HBK. Nov. Gen. & Sp. 2: 51. 1817. Stil-

166 FIELDIANA: BOTANY, VOLUME 24

lingia microsperma Pax & Hoffm. Pflanzenreich IY. 147, v: 187.

1912 (type from Santa Rosa, Heyde & Lux 4265). Pimientillo

(Jalapa).

Chiefly on rather dry, brushy, rocky hillsides, sometimes in oak

forest, 1,200-2,000 meters; Guatemala; Quiche". Mexico; British

Honduras (San Agustin); Panama.

A shrub of 1-2.5 meters, much-branched or almost simple, densely leafy,

often umbellately branched above, glabrous; leaves alternate or often pseudo-

verticillate above at the base of the branches, on petioles 2-6 mm. long, ovate-

lanceolate to lanceolate or oblong, mostly 4-10 cm. long and 1.5-2.5 cm. wide,

acuminate or long-acuminate, acute or attenuate at the base, thick-membrana-

ceous, reticulate- veined, somewhat paler beneath; spikes terminal, 8-12 cm. long,

stout, straight, the bracts very broadly triangular, subulate-acuminate, the

staminate 7-11-flowered; staminate flowers on very short pedicels, the pistillate

sessile; staminate calyx 2-lobate, the 3 pistillate sepals ovate, acute; stamens 2,

exserted; ovary subcarinate, the styles very shortly connate; capsule 10-12 mm.

long, 15 mm. broad, the receptacle 11 mm. broad; seeds 5-7 mm. long and broad,

whitish, smooth.

This has been reported from Guatemala as S. sylvatica Muell.

Arg. S. microsperma was separated on the basis of size of the

seeds, those of S. microsperma being 5 mm. long, those of S. zelayen-

sis 6-7 mm. long, scarcely an important difference even if it existed

constantly.

TETRORCHIDIUM Poeppig

Reference: F. Pax, Pflanzenreich IV. 147, iv: 29-32. 1912.

Trees, the pubescence of simple or malpighiaceous (appressed and attached

by the middle) hairs, usually soon glabrate; leaves alternate, membranaceous,

petiolate, stipulate, large, penninerved, entire or dentate, the petiole with 2 con-

spicuous glands at the apex; flowers small, dioecious or monoecious, apetalous, the

racemes axillary, slender, the staminate elongate, simple or branched, the flowers

subsessile, glomerate; pistillate and androgynous racemes shorter, simple, some-

times reduced to a single flower; staminate calyx small, 3-parted, the sepals

broad, slightly imbricate; disk none; stamens 3, episepalous, the filaments very

short, in bud connate into a subglobose mass; anthers broad, introrsely and

peltately 4-celled; ovary rudiment none or clavate, equaling the stamens; pistillate

calyx like that of the staminate flower, the disk cyathiform or of 3 petaloid scales;

ovary 2-3-celled, the style short, flat, 2-fid; ovules solitary in each cell; capsule

2-3-dymous, separating into 2-valvate cocci, the endocarp thin-crustaceous; seeds

globose, coarsely foveolate, the testa crustaceous; cotyledons broad, flat.

About 12 species, in tropical America. One other is known in

southern Central America.

Petioles about 1 cm. long; leaf blades 9-12 cm. long, abruptly short-acuminate;

rachis of the staminate inflorescence glabrous T. brevifolium.

Petioles 2-4 cm. long; leaf blades mostly 10-25 cm. long, rounded or very obtuse

at the apex; rachis of the staminate inflorescence pubescent. . .T. rotundatum.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 167

Tetrorchidium brevifolium Standl. & Steyerm. Field Mus.

Bot. 23: 126. 1944.

Known only from the type, Alta Verapaz, in virgin forest,

Rubelpec, C. L. Wilson 188.

A small tree, glabrous throughout, the branchlets subterete, ochraceous, the

young ones scarcely more than 2 mm. thick; leaves on slender petioles 4-10 mm.

long, membranaceous, oblong-obovate or obovate-elliptic, 9-12 cm. long, 4-5.5

cm. wide, abruptly short-acuminate, with an obtuse or subacute tip, cuneate-

attenuate at the base, entire or nearly so, almost concolorous, the lateral nerves

about 6 pairs, very slender and inconspicuous, arcuate; staminate spikes very

slender, flexuous, short-pedunculate, 3-7 cm. long, lax and interrupted, the

flowers sessile, solitary or few together, green, 3 mm. broad; sepals very broadly

ovate, rounded or very obtuse at the apex, ciliolate; stamens 3, the anthers sub-

sessile.

Tetrorchidium rotundatum Standl. Trop. Woods 16: 44.

1928. Canjura negro, (fide Aguilar).

Moist or wet, mixed forest, 300-1,500 meters; Guatemala;

Retalhuleu; Quezaltenango. Veracruz and Chiapas; Atlantic coast

of Honduras and Nicaragua and perhaps farther southward.

A rather small to large tree, sometimes 30 meters high with a trunk 50-75

cm. in diameter, the crown spreading, the trunk often short, the whole plant gla-

brous except in the inflorescence, the young branches stout, mostly 5-10 mm.

thick, terete, rather densely leafy; bark pitted and greenish gray below, smoother

above; petioles slender, 2-4 cm. long, bearing above the middle remote from the

blade 2 small, sessile, unequally inserted, orbicular glands; leaf blades obovate-

oblong, mostly 10-25 cm. long and 3-10 cm. wide, rounded or very obtuse at the

apex, gradually long-attenuate to the base, thick-membranaceous, bright green

above, somewhat paler beneath, entire, the lateral nerves about 7 pairs; staminate

spikes short-pedunculate, 7-15 cm. long, the rachis densely puberulent, the flowers

green, sessile, aggregate in dense, remote or crowded clusters; sepals broadly

triangular, 2 mm. long, densely puberulent; capsules mostly 2-celled, didymous,

8 mm. broad.

Called "manteca" in Honduras. The trunk is often supported

by high buttresses. The wood is almost white, light in weight,

soft, woolly, and perishable. Called "amate bianco" in Chiapas.

TRAGIA L.

Perennial herbs or rarely woody plants, erect or more often scandent, usually

hispid with more or less stinging hairs; leaves alternate, petiolate, mostly palmate-

nerved, 2-stipulate; flowers generally monoecious, apetalous, the inflorescences

terminal or opposite the leaves, rarely axillary, androgynous, the lower flowers

pistillate, the upper staminate, the flowers solitary or rarely cymulose in the

axils of the bracts; staminate calyx closed in bud, globose or obovoid, in anthesis

168 FIELDIANA: BOTANY, VOLUME 24

valvately 3-5-parted; glands of the disk developed between the outer stamens,

free or somewhat connate with the filaments, usually absent; stamens mostly 3,

alternate with the sepals, or by abortion 2-1, sometimes more numerous, the

filaments commonly short, more or less connate at the base; anthers oblong,

extrorse or introrse, longitudinally dehiscent; pistillate sepals 6, rarely 3, entire,

persistent and little accrescent in age, sometimes pinnatifid and accrescent; disk

none; ovary 3-celled, the styles connate into a column, simple; ovules solitary in

each cell; capsule 3-coccous, depressed, separating into 2-valvate cocci, the

endocarp crustaceous; seeds not carunculate, globose; endosperm carnose, the

cotyledons broad, flat.

About 125 species, in the tropics of both hemispheres, mostly in

tropical America, a few in the Old World tropics and in temperate

North America. No others are known in Central America.

Stamens about 40; leaves very large, mostly 8-25 cm. wide T. Bailloniana.

Stamens 3; leaves small, mostly 1.5-6 cm. wide.

Anthers extrorse.

Stems scandent and twining T. volubilis.

Stems erect or nearly so, not twining T. nepetifolia.

Anthers introrse.

Inflorescence hispidulous, without gland-tipped hairs T. yucatanensis.

Inflorescence bearing numerous short gland-tipped hairs, also pilosulous.

T. mexicana.

Tragia Bailloniana Muell. Arg. Linnaea 34: 178. 1865. Zucker-

tia cordata Baill. Etud. Euphorb. 496. pi. 4- 1858, not Tragia cordata

Michx. 1803.

Moist or wet, mixed forest or in thickets, 1,300 meters or less;

Huehuetenango; Quezaltenango; to be expected in Izabal. Tabasco;

Atlantic lowlands of Honduras.

A small or large, herbaceous or woody vine, hirsute almost throughout with

very slender, stiff, spreading hairs; leaves on very long and slender petioles, ovate

to rounded-ovate, as much as 25 cm. long and wide, thin-membranaceous, shallowly

3-lobate or undivided, the blades or their lobes undulate-dentate, deeply cordate

at the base and 5-nerved, thinly hirsute on both surfaces; stipules herbaceous,

triangular-ovate, acuminate, dentate; racemes opposite the leaves, as much as

25 cm. long and many-flowered, bifurcate, one branch staminate, the other pistil-

late, the bracts lanceolate, acuminate, reflexed, the pistillate ones 1-flowered, the

staminate 1-3-flowered, 6 mm. long, hispidulous, the staminate pedicels 9-12 mm.

long, the pistillate shorter and stouter; staminate buds pyriform, acute, the sepals

acuminate, pubescent; capsule about 8 mm. long, tridymous, densely hirsute.

Because of its very large, broad leaves this plant is very unlike

all other North American species of Tragia. In Honduras it is some-

times called "chichicaste."

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 169

Tragia mexicana Muell. Arg. Linnaea 34: 181. 1865.

Moist or wet forest or thickets, often on limestone, 350-1,400

meters; Alta Verapaz; Jutiapa; Guatemala. Southern Mexico.

Plants wholly or chiefly herbaceous, twining and scandent, bearing a few

stinging hairs, the stems puberulent or incurved-pilosulous; leaves on petioles

1.5-4.5 cm. long, membranaceous, oblong-ovate, 4.5-15 cm. long, 1.5-5 cm. wide,

acute or acuminate, shallowly cordate or truncate-cordate at the base, subentire

or conspicuous-dentate, 5-nerved at the base, thinly hispidulous or glabrate above,

densely puberulent beneath; stipules 5 mm. long or less, lanceolate, acuminate;

racemes terminal or opposite the leaves, simple or rarely branched from the base,

8 cm. long or usually much shorter, naked below, bearing at the base 1 or rarely

2 pistillate flowers, the upper flowers staminate, the rachis pubescent and stipitate-

glandular; bracts linear-lanceolate, the staminate about 3 mm. long, the pistillate

5 mm. long, the pedicels elongate, the pistillate sometimes 15 mm. long; staminate

flowers 4 mm. broad, the 3 sepals ovate, acute; stamens 3, the filaments short,

the minute anthers introrse; pistillate sepals 6, ovate, acuminate, in age 5-6 mm.

long, reflexed, pubescent and sparsely glandular; ovary densely hispid, the styles

connate into a column 4 mm. long, free and reflexed above; capsule 12 mm. broad

and 7 mm. high, densely hispid; seeds globose, 5 mm. in diameter, brownish

yellow mottled with fuscous.

Tragia nepetifolia Cav. Icon. PI. 6: 37. pi. 557, f. 1. 1801.

Grassy or brushy slopes, 900-1,500 meters; Chiquimula; Huehue-

tenango. Southwestern United States; Mexico.

Plants perennial from a ligneous root, branched from the base, erect or nearly

so, 15-40 cm. high, the stems slender, pubescent and hispidulous with somewhat

stinging hairs; leaves on petioles 3-15 mm. long, triangular or triangular-ovate

to triangular-linear, 1.5-5 cm. long, 4-15 mm. wide, acute or acuminate, cordate

or truncate at the base, coarsely serrate, rather densely hispidulous; stipules 2 mm.

long, triangular; inflorescences opposite the leaves, 7-20 mm. long, hispidulous,

eglandular, bearing 1 pistillate flower at the base and a few staminate ones above,

the staminate bracts small, lanceolate or linear-lanceolate, the pistillate bracts

ovate, the pedicels 1 mm. long in anthesis; staminate flowers 1.5-2 mm. broad,

the 3-4 sepals ovate, acute; stamens 3, the anthers extrorse, shorter than the

filaments; pistillate sepals 6, ovate or lanceolate, unequal, 2 mm. long; ovary

strigose-hirsute; capsule 6-7 mm. broad, hispid; seeds globose, 2-3 mm. in diameter,

mottled.

This species has been reported from Izabal (Livingston, Tuerck-

heim 8751), but probably incorrectly so.

Tragia volubilis L. Sp. PI. 980. 1753. T. guatemalensis Lotsy

in Bonn. Smith, Bot. Gaz. 20: 354. 1895 (type from Chicacao, Alta

Verapaz, J. D. Smith 1763). Chichicaste de raton (Huehuetenango).

Open or brushy slopes and fields, 200-1,400 meters; Alta Verapaz;

Zacapa; Chiquimula; Santa Rosa; Huehuetenango. Southern

170 FIELDIANA: BOTANY, VOLUME 24

Mexico; Salvador; Honduras; Costa Rica; West Indies; South

America; Africa.

A perennial herbaceous vine, usually not more than 1-2 meters long, the stems

very slender, hirsute and often puberulent; leaves thin-membranaceous, on petioles

1-5 cm. long, triangular-ovate to lance-ovate, 3-15 cm. long, 1-5 cm. wide, acute

or acuminate, cordate or truncate-cordate at the base, serrate-dentate, hirsute

when young, in age glabrate except on the nerves, palmate-nerved; stipules ovate-

lanceolate, 3 mm. long; inflorescences lateral, sometimes fasciculate, 2-6 cm. long,

bearing 1-2 pistillate flowers at the base, the staminate bracts 1-2 mm. long,

lance-ovate, acuminate, 1-flowered, the pedicels 2-3 mm. long; pistillate pedicels