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FLORA OF GUATEMALA
PAUL C. STANDLEY
AND
JULIAN A. STEYERMARK
FIELDIANA: BOTANY
VOLUME 24, PART IV
Published by
CHICAGO NATURAL HISTORY MUSEUM
APRIL 11, 1946
FLORA OF GUATEMALA
PART IV
FLORA OF GUATEMALA
PAUL C. STANDLEY
Curator of the Herbarium
AND
JULIAN A. STEYERMARK
Assistant Curator of the Herbarium
FIELDIANA: BOTANY
VOLUME 24, PART IV
Published by
CHICAGO NATURAL HISTORY MUSEUM
APRIL 11, 1946
lit* UtfNAKY OF THE
MAY 9 1946
PRINTED IN THE UNITED STATES OF AMERICA
BY THE CHICAGO NATURAL HISTORY MUSEUM PRESS
CONTENTS
Families Included in Part IV
Ulmaceae .
PAGE
1
Winteraceae ..................... 269
Moraceae ....................... 10 Annonaceae .......... . .......... 270
Proteaceae ...................... 58 Myristicaceae ................... 294
Loranthaceae ............... ..... 62 Monimiaceae .................... 299
Opiliaceae ...................... 86 Lauraceae ....................... 302
Olacaceae ....................... 88
Balanophoraceae ................. 92
Aristolochiaceae ................. 93
Hernandiaceae 344
Papaveraceae 347
Cruciferae 354
Rafflesiaceae 101 Tovariaceae . . 380
Polygonaceae . . 104 Capparidaceae 380
Resedaceae 397
Moringaceae 398
Droseraceae 399
Podostemonaceae . . .401
Chenopodiaceae 137
Amaranthaceae 143
Nyctaginaceae 174
Phytolaccaceae 192
Aizoaceae 203
Portulacaceae . . .207 Crassulaceae .... .404
Basellaceae 214 Saxif ragaceae . .416
Caryophyllaceae 217 Brunelliaceae .
Nymphaeaceae 239 Cunoniaceae
Ceratophyllaceae 242 Hamamelidaceae . ... 426
Ranunculaceae 243 Platanaceae. . .430
Berberidaceae 256 Rosaceae 432
Menispermaceae 258 Connaraceae 484
Magnoliaceae 266 Krameriaceae 488
INTRODUCTION
The Flora of Guatemala, of which this is the first part to be pub-
lished, has been in preparation for the past six years. It is based
upon published records of Guatemalan plants and upon the earlier
collections now available in the Herbarium of Chicago Natural
History Museum. Principally, however, it records new informa-
tion obtained by the authors during four botanical expeditions of
the Museum. These expeditions were extended to all the twenty-
two departments of Guatemala and to almost all corners of the
country. This intensive exploration was possible because of the
admirable network of Guatemalan roads, which enable one to reach
by automobile almost every village except in a few sparsely settled
areas. These the junior author has explored on foot or on horseback.
Almost all the manuscript of the Flora has been written, at least
in provisional form, and it was planned to publish it in systematic
order. Because of conditions imposed by the war, this has been
found impractical. Part I will include an account of the general
features of Guatemalan vegetation, a re'sume' of the history of its
exploration, and other pertinent matter.
It is believed that the form in which the data are presented on
the following pages will be found sufficiently obvious, but an explana-
tion of some of the details will be included in the introductory
chapters. The flora of Guatemala, as here considered, includes
that of British Honduras, which is continuous with that of the
departments of Pete"n and Izabal. There is no reason to suppose
that in British Honduras there exists more than a handful of species
that will not be found eventually in Guatemala.
ULMAGEAE. Elm Family
Trees or shrubs with watery sap; leaves alternate, petiolate, simple, entire to
serrate or crenate, stipulate, the stipules usually small and fugacious, free or united;
flowers small and usually green or yellowish, monoecious, dioecious, perfect, or
polygamous, mostly in small cymes or racemes, or the pistillate often solitary in
the leaf axils; perianth normally 4-5-parted or of 4-5 distinct sepals; petals none;
2 FIELDIANA: BOTANY, VOLUME 24
stamens as many as the perianth segments and opposite them, the filaments
straight or nearly so; anthers erect in bud, 2-celled, longitudinally dehiscent;
ovary 1-celled, the ovule solitary, pendulous from the apex of the cell, anatropous
or amphitropous; styles or stigmas 2; fruit a samara, nut, or drupe; endosperm
scant or none; embryo straight or curved, the cotyledons usually flat.
About a dozen genera, in tropical and temperate regions of
both hemispheres. No other genera occur in Central America.
Leaves opposite Lozanella.
Leaves alternate.
Fruit dry, sometimes winged.
Fruit winged, not ciliate Phyllostylon.
Fruit not winged, long-ciliate Chaetoptelea.
Fruit drupaceous.
Cotyledons narrow; fruit scarcely more than 2 mm. long, juicy; leaves dentate
and 3-nerved (in Guatemalan species) Trema.
Cotyledons broad; fruit usually 7-15 mm. long and not red.
Stamens as many as the perianth segments; leaves entire or serrate; plants
sometimes armed with spines Celtis.
Stamens twice as many as the perianth segments or more numerous; leaves
entire, 3-nerved; plants unarmed Ampelocera.
AMPELOCERA Klotzsch
Unarmed trees; leaves alternate, very shortly petiolate, membranaceous or
coriaceous, remotely serrate or entire, penninerved or obscurely 3-nerved ; stipules
lateral, free; flowers small, polygamous, perfect or by the abortion of the ovary
staminate, fasciculate or racemose in the leaf axils; perianth cuplike, 5-lobate, the
lobes ovate, imbricate; stamens 10, the filaments filiform; exserted; ovary ovoid,
the style 2-parted, the branches subulate, divaricate; ovule pendulous from the
apex of the cell; fruit small, drupaceous.
Two other species are known, in Cuba and Peru.
Ampelocera Hottlei Standl. Trop. Woods 51: 11. 1937. Celtis
Hottlei Standl. Trop. Woods 20: 20. 1929. Lain (Pete'n).
Wet or swampy forest, at or little above sea level; Pete'n; Alta
Verapaz; Izabal; Retalhuleu; Huehuetenango. Oaxaca; British
Honduras; Honduras; Panama; Colombia.
Sometimes only a large shrub but usually a large or medium-sized tree,
sometimes 30 meters high with a trunk 50 cm. or more in diameter; branchlets
grayish-puberulent, often glaucescent; petioles stout, mostly 7-12 mm. long, the
blades coriaceous, oblong or elliptic-oblong, mostly 8-16 cm. long and 3-7.5 cm.
wide, somewhat lustrous, abruptly acuminate, subacute to almost rounded at
the base and often somewhat oblique, obviously 3-nerved from the base but
essentially penninerved, glabrous; flowers densely congested and sessile in the
leaf axils or often in small cymes, the inflorescences scarcely longer than the
petioles, densely puberulent and sometimes glaucescent; drupes oval-globose,
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 3
about 13 mm. long, densely scabrous-puberulent, the persistent style branches
about 3 mm. long.
The name "chaperno" has been reported from Guatemala,
probably in error. In British Honduras the tree is called "bullhoof" ;
in Honduras "manteca"; in Oaxaca "cautivo" and "frijolillo."
The trunk is sometimes buttressed; the bark is grayish brown and
fairly smooth, about 1 cm. thick, the inner bark yellowish or pale
brown. Sapwood creamy yellow to light brown; heartwood dark
with almost black streaks; fresh heartwood with a faint fragrance;
wood not difficult to cut, splits rather easily. In Oaxaca the wood
is utilized for railroad ties and house construction.
CELTIS L.
Trees or shrubs, sometimes armed with spines; leaves alternate, deciduous or
persistent, serrate or entire, penninerved and often also 3-nerved, frequently
oblique at the base; stipules lateral, free; flowers polygamous, small, greenish, in
small cymes, the cymes staminate or androgynous, lax and open or sometimes
small and congested, the fertile flowers usually long-pedicellate; perianth shallowly
or deeply 5-lobate, the segments imbricate; stamens normally 5, the filaments
erect or nearly so, finally exserted, the anthers ovate; torus usually densely pilose;
ovary sessile, the style central, 2-parted, the branches plumose-stigmatose, diver-
gent, simple or bifid; ovule pendulous from the apex of the cell, anatropous; fruit
drupaceous, usually with scant flesh, ovoid or globose, sometimes 2-carinate, the
endocarp osseous; testa of the seed membranaceous, the endosperm scant or none;
embryo curved, the cotyledons broad, incurved-replicate, sometimes corrugate.
About 75 species, in temperate and tropical regions of both
hemispheres. No other species are native in Central America.
Branches armed with recurved spines C. iguanaea.
Branches unarmed.
Leaves not at all 3-nerved, with numerous pairs of lateral nerves . . . . C. monoica.
Leaves conspicuously 3-nerved, the lateral nerves usually 4 or fewer pairs.
Leaves conspicuously and evenly serrate or crenate C. trinervia.
Leaves, at least most of them, entire or nearly so.
Leaves glabrous C. Schippii.
Leaves densely and softly pubescent beneath C. caudata.
Celtis caudata Planch. Ann. Sci. Nat. III. 10: 294. 1848.
At 1,200-1,300 meters; Huehuetenango (along Rio Cuilco
between Cuilco and San Juan, Steyermark 50906). Southern Mexico.
A tree about 12 meters high, the young branchlets densely and softly pubes-
cent; leaves subcoriaceous, short-petiolate, ovate, asymmetric, mostly 4-6.5 cm.
long, acuminate or narrowly long-acuminate, rounded at the base, entire or some-
times dentate near the base, scabrous on the upper surface and very rough to the
touch, densely and softly pubescent beneath, 3-nerved from the base; fruiting
4 FIELDI AN A: BOTANY, VOLUME 24
pedicels 1-1.5 cm. long or longer; fruit subglobose, about 8 mm. long, probably
black at maturity.
Celtis iguanaea (Jacq.) Sarg. Silva N. Amer. 7: 64. 1895.
Rhamnus iguanaeus Jacq. Enum. PI. Carib. 16. 1760. C. aculeata
Swartz, Prodr. Veg. Ind. Occ. 53. 1788. C. anfractuosa Liebm.
Dansk. Vid. Selsk. Skrivt. V. 2: 338. 1851. Cagalero; Rompa-caite ;
Piscucuy (Zacapa) ; Clavo verde (Huehuetenango).
Dry or wet thickets of plains and hillsides, mostly at 1,000
meters or less; Zacapa; Chiquimula; El Progreso; Jalapa; Jutiapa;
Santa Rosa; Escuintla; Guatemala; Chimaltenango; Retalhuleu;
Suchitepe"quez; Huehuetenango. Florida and western Texas;
Mexico to British Honduras and Panama; West Indies; South
America.
A shrub or small tree, often with long, recurved or somewhat scandent
branches, the trunk sometimes as much as 30 cm. in diameter, often branched
from the base, the branches armed with stout, short, usually recurved spines, the
ultimate branches often compressed; leaves short-petiolate, ovate to oval, mostly
5-13 cm. long, abruptly acuminate, rounded or subcordate at the base, conspicu-
ously 3-nerved at the base, serrate except near the base, sparsely pubescent or
almost glabrous; flowers greenish yellow, in small, lax or dense cymes about as
long as the petioles; fruit ovoid, orange, yellow, or red, 8-12 mm. long.
Called "ufia de gato" and "cagalero bianco" in Salvador; in
Yucatan "muc" or "zitsmuc" (Maya); in Oaxaca "palo de arco."
The fruit is reported to be edible but the flesh is scant and its flavor
not appetizing. Birds are said to be fond of it and children some-
times gather it. The shrub is a common one in the dry Motagua
Valley, and in thickets of the dry Pacific plains sometimes is domi-
nant. The bark is brown, smooth or slightly fissured; sap wood
white, the heartwood scant, dark brown or almost black. The
name "palo de arco" used in Oaxaca refers to the fact that the main
branches often are recurved, like a bow. The specific name used by
Jacquin (iguanaea) is said to relate to the fact that iguanas eat the
fruit.
Celtis monoica Hemsl. Biol. Centr. Amer. Bot. 3: 139. pi. 77.
1883. Capulin macho (fide Aguilar); Mescal.
Moist or wet, mixed, mountain forest, 1,000-1,600 meters;
Guatemala; Chimaltenango; Huehuetenango. San Luis Potosi,
Veracruz, and Oaxaca; Honduras; Salvador.
A tall tree, sometimes 25 meters high, with slender branchlets; leaves on
short slender petioles, lance-oblong or oblong-ovate, mostly 7-11 cm. long and
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 5
2-4 cm. wide, rather thin or coriaceous, sometimes lustrous on the upper surface,
long-attenuate or caudate-acuminate, acute or obtuse at the base, rather coarsely
serrate almost to the base, sparsely strigose beneath with straight, closely appressed
hairs; fruit subglobose, covered with sharp tubercles.
Called "duraznillo" in Salvador and "yaya" in Honduras.
Celtis Schippii Standl. Field Mus. Bot. 12: 409. 1936. Bullhoof.
British Honduras, and probably extending into Pete"n or Izabal;
type from Temash River, Schipp 1322; collected also at Middlesex
and in Silk Grass Forest Reserve.
A glabrous tree 15 meters tall, the trunk 25 cm. in diameter, with slender
branchlets; leaves short-petiolate, subcoriaceous, the petioles 5-8 mm. long, the
blades oblong or elliptic-oblong, mostly 8-11 cm. long and 3.5-4.5 cm. wide,
short-acuminate, obtuse or subacute at the base and more or less oblique, entire,
lustrous above, 3-nerved at the base; pistillate flowers mostly solitary, axillary,
the pedicels as much as 7 mm. long, the sepals persistent, rounded, 1 mm. long,
ciliate; drupes ellipsoid, glabrous, 1.5 cm. long and 1 cm. broad, narrowed at the
base.
Celtis trinervia Lam. Encycl. 4: 140. 1797. C. petenensis
Lundell, Bull. Torrey Club 69: 387. 1942 (type from Lake Yaxha,
C. L. Lundell 4306).
At 200 meters or less; Pete"n (Lake Yaxha; Uaxactun). Greater
Antilles.
A slender tree, sometimes 18 meters high, the bark smooth and gray; leaves
short-petiolate, membranaceous, ovate or ovate-lanceolate, 4-13 cm. long, 2-7.5
cm. wide, long-acuminate, at the base usually rounded on one side and acute on
the other, closely and regularly serrate or crenate, bright green, short-pilose,
especially beneath, or glabrate; staminate flowers in lax axillary cymes, the pistil-
late flowers often solitary; fruit purple-black, 7-8 mm. long, about equaling the
pedicel or longer, the stone subglobose, rugose.
CHAETOPTELEA Liebmann
Large trees, unarmed; leaves alternate, somewhat distichous, penninerved,
serrate, deciduous or often persistent; stipules lateral, scarious, caducous; fascicles
of flowers borne at the nodes, solitary and sessile, at first covered with imbricate
scales; flowers numerous in the clusters, polygamous or most of them perfect;
perianth campanulate, 4-8-1 obate, the lobes imbricate; stamens usually 5, the
filaments erect, finally exserted, the anthers glabrous; ovary stipitate, compressed,
the style short, 2-fid, the branches introrsely stigmatose; ovule pendulous from
the apex of the cell; fruit dry, compressed, elliptic, very thin, not winged, densely
long-ciliate; seed compressed; endosperm none, the embryo straight, the coty-
ledons plane.
The genus consists of a single species. Some authors have united
it with Ulmus, but in that the fruit is conspicuously winged.
6 FIELDIANA: BOTANY, VOLUME 24
Chaetoptelea mexicana Liebm. Nat. For. Kjoebenhavn Vid.
Medd. 1850: 76. 1851. Ulmus mexicana Planch, in DC. Prodr. 17:
156. 1873. Duraznillo; Mescal; Muyaul (San Marcos).
Moist mountain forest, 900-2,700 meters; Alta Verapaz; Zacapa;
Sacatepe"quez; Chimaltenango; Solola; Retalhuleu; Huehuetenango ;
Quezaltenango; San Marcos. Southern Mexico; Honduras; Costa
Rica; Panama.
A large tree, usually 10-25 meters tall or even more, with a rather open crown,
the trunk often 75 cm. in diameter, the bark gray, somewhat scaly; leaves decidu-
ous, short-petiolate, lance-oblong to oblong-ovate, mostly 5-9 cm. long, sometimes
larger, especially on young shoots, acuminate or long-acuminate, obtuse to sub-
cordate at the base, unequally and often coarsely serrate, scabrous, especially
beneath and usually very rough to the touch, sometimes smooth on the upper
surface, the lateral nerves numerous, prominent beneath; flowers yellowish, the
pistillate or perfect ones in lax racemes; fruit slender-stipitate, about 5 mm. long,
pale green, the margins densely beset with long soft hairs, bidentate at the apex.
Called "membrillo" in Honduras. The wood is rather hard,
heavy, tough and strong, in structure like that of Ulmus (elm) ; the
heartwood is deep reddish brown, often with darker streaks, while
the thick sapwood is light brownish gray. No data are available
regarding any use of the wood in Guatemala but in Salvador it is
employed for railroad ties, cart axles, beds, and cart wheels. The
tree is a common one of the central mountains of Guatemala, and
some individuals must be fully 30 meters tall, with very massive
trunks. It is plentiful also on hillsides about Fuentes Georginas in
Quezaltenango. Small trees and seedlings seem to be scarce in these
same regions.
LOZANELLA Greenman
Reference: E. P. Killip & C. V. Morton, The genus Lozanella,
Journ. Wash. Acad. Sci. 21: 336-339. 1931.
Shrubs or small trees, the branches opposite; leaves opposite, slender-petiolate,
membranaceous, serrate, 3-nerved; stipules united; flowers dioecious, small, green,
in rather lax, small, axillary cymes; pistillate perianth 5-6-parted, the segments
imbricate; ovary sessile, the single ovule pendulous; style 2-parted to the base,
the branches papillose; fruit a small drupe, ovoid, somewhat compressed, with
juicy exocarp and osseous endocarp; endosperm fleshy; embryo curved, the coty-
ledons broad, equal.
Two species are known, the other in Peru and Bolivia.
Lozanella enantiophylla (Donn. Smith) Killip & Morton,
Journ. Wash. Acad. Sci. 21: 339. 1931. Trema enantiophylla Donn.
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 7
Smith, Bot. Gaz. 33: 259. 1902. L. trematoides Greenm. Proc. Amer.
Acad. 41:236. 1905.
Damp mixed upland forest, 1,400-3,000 meters; El Progreso;
Zacapa (Sierra de las Minas); Guatemala; Chimaltenango; Quiche";
Quezaltenango; San Marcos. Southern Mexico; Costa Rica;
Colombia; Peru.
A tree 6-9 meters high, the trunk as much as 25 cm. in diameter, the young
branches densely villous-pilose, the older branches brown; leaves slender-petiolate,
broadly ovate to lance-ovate, 9-16 cm. long, 5-9 cm. wide, usually acuminate or
long-acuminate, obtuse or acute at the base, rather evenly and closely crenate,
bright green, very rough above, rather densely and shortly harsh-pilose beneath,
conspicuously 3-nerved; inflorescences usually about equaling the petioles; fruits
subglobose, very juicy, scarcely more than 2 mm. in diameter, bright orange.
PHYLLOSTYLON Capanema
Unarmed trees with rough pubescence; leaves alternate, short-petiolate,
deciduous, penninerved, crenate or serrate; stipules small, lateral, distinct, cordate-
lanceolate; flowers polygamous, the clusters fasciculate on leafless branches, sessile,
subtended by a few imbricate scales; lower flowers of the fascicle staminate, the
uppermost fertile; perianth 5-8-parted, the segments narrow, thin, slightly imbri-
cate; stamens usually fewer than the perianth segments, unequal, the filaments
short, erect, the anthers glabrous; ovary sessile, compressed; style continuous
with the ovary, plane, broadly falcate, or usually unequally and divaricately
bilobate, the upper margin stigmatose; ovule pendulous from the apex of the cell;
fruit dry, samaroid, compressed, terminated by a large membranaceous unequal
falciform wing, with another small wing at the base; seed subcordiform, with a
thin testa; endosperm none, the embryo straight.
One or perhaps two other species are known, in South America.
Phyllostylon rhamnoides (Poisson) Taubert, Oesterr. Bot.
Zeit. 40: 409. 1890. Samaroceltis rhamnoides Poisson, Journ. Bot. 1:
256. 1887.
Dry brushy hillsides of the Oriente, 300-700 meters; Zacapa;
Chiquimula. Southern Mexico; Cuba and Haiti; Colombia; Vene-
zuela; Argentina.
A tree, in some parts of its range 15 meters high with a trunk 70 cm. in diame-
ter, but in Guatemala usually much smaller, with stiff irregular branches, the
young branchlets as well as the leaves scabrous or scabrous-puberulent; leaves on
very short petioles, the blades broadly ovate to oval, mostly 2-4.5 cm. long and
1.5-2.5 cm. wide, on young branches often larger, obtuse or subacute, broadly
rounded to shallowly cordate at the base, pale green when dried, very rough to
the touch; flowers usually produced when the tree is leafless, small, greenish,
inconspicuous; fruits mostly 2.5-3 cm. long, resembling the samaras of Acer, the
seed-bearing portion short and hard, densely short-pilose, the thin wing 6-8 mm.
wide, thickened along one edge.
8 FIELDIANA: BOTANY, VOLUME 24
The Maya name used in Yucatan is "canche"; in Mexico the tree
is called "ceron." The wood is clear deep yellow to very pale brown,
with a thin layer of white or colorless sap wood, heavy, hard, and
compact, with very fine and uniform texture, the grain straight or
nearly so, easy to carve and turn, and taking a high polish. From
the Dominican Republic it has been exported to the United States
under the trade name of "San Domingan boxwood." It is employed
for weaver's shuttles, rulers, and piano keys. So far as known, the
wood is used in Guatemala only for firewood.
TREMA Loureiro
Trees or shrubs, unarmed, usually with rough pubescence; leaves alternate,
commonly distichous, short-petiolate, serrate or entire, penninerved and 3-nerved
at the base; stipules lateral, free, small, caducous; flowers small, monoecious,
dioecious, or polygamous, sessile in the leaf axils or in axillary cymes; staminate
perianth usually 5-lobate, the segments induplicate-valvate; stamens normally
5, the filaments short, erect; segments of the pistillate calyx slightly imbricate;
ovary sessile, the style central, divided, often to the base, the branches stigmatose,
linear, the ovule pendulous; torus of the flower usually pilose; fruit a small drupe,
ovoid or subglobose, usually terminated by the persistent style branches, the
exocarp succulent and juicy, the endocarp hard; testa of the seed membranaceous,
the endosperm fleshy; embryo curved or almost involute, the cotyledons narrow,
the radicle incumbent, ascending.
Perhaps 20 species, widely dispersed in tropics of both hemi-
spheres. One other Central American species occurs in Panama.
It seems that in all regions where the trees grow their characters are
variable and the species separable only with difficulty. Guatemalan
material of the genus is separable into three groups that usually can
be distinguished, at least when ample material is available for com-
parison. These may represent distinct species but it appears more
satisfactory to treat them all as forms of the widespread T. micrantha.
Trema micrantha (L.) Blume, Mus. Bot. Lugd. Bat. 2: 58.
1853. Rhamnus micranthus L. Syst. Nat. ed. 10. 2: 937. 1759.
Sponia micrantha Decaisne, Nouv. Ann. Mus. Paris 3: 498. 1834.
Capulin; Kib (Quecchi).
Chiefly in dry thickets, along streams, or often on plains, ascend-
ing from sea level to about 2,000 meters; Alta Verapaz; Zacapa;
Baja Verapaz; El Progreso; Chiquimula; Santa Rosa; Escuintla;
Guatemala; Sacatepe"quez ; Chimaltenango; Solola; Quiche"; Que-
zaltenango; Suchitepe"quez; probably in all or most of the other
departments except Totonicapan. Mexico to British Honduras
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 9
and Panama; West Indies; through most of South America except
the highlands.
A shrub or a tree, in Guatemala sometimes 15 meters high or more, the bark
thin, brown, shallowly fissured; leaves short-petiolate, oblong-ovate to oblong-
lanceolate, mostly 6-15 cm. long, acute or acuminate, rounded to subcordate at
the base, finely serrate, very rough to the touch, beneath usually densely pilose
with short spreading hairs, or the pubescence sometimes sparse; flowers very small,
green or yellowish, the cymes small, dense or lax, little exceeding the petioles;
fruit about 2 mm. long, bright red or orange-red.
Known in British Honduras by the names "white capulin,"
"wild bay cedar," and "bastard bay cedar"; in Salvador called
"capulin macho," "capulin months," "capulincillo," and "chu-
rrusco"; "capulin negro" (Honduras). The bark contains a strong,
tough fiber that is used as cordage in Guatemala and throughout
Central America. Along the North Coast the tree is sometimes
found in such places as Manicaria swamps, but dry situations are
more usual. This is one of the most common and characteristic
species of second-growth thickets.
Trema micrantha var. floridana (Britton) Standl. & Steyerm.
Field Mus. Bot. 23: 40. 1944. T. floridana Britton ex Small, Fl.
Southeast. U. S. 366: 1329. 1903. Capulin cimarrdn (Pete'n).
Occasional in thickets, 1,600 meters or less; Pete'n; Jalapa;
Huehuetenango. Southern Florida; Yucatan Peninsula of Mexico;
British Honduras.
A large shrub or small tree, similar to the species, but the leaves mostly ovate
or even broadly ovate, short-acuminate, rather deeply cordate at the base, densely
and rather softly short-pilose beneath.
Usually this can be recognized by the relatively broad and con-
spicuously cordate leaves but some intermediate forms are found
in Central America. This seems to be the only form of the species
in Yucatan.
Trema micrantha var. strigillosa (Lundell) Standl. & Steyerm.
Field Mus. Bot. 23: 40. 1944. T. strigillosa Lundell, Phytologia 1:
337. 1939 (type from Middlesex, British Honduras, W. A. Schipp
439). Capulin; Kiim (Coban, Quecchi).
Moist thickets or forest, sometimes in pine forest, chiefly at 800-
1,400 meters; Alta Verapaz; Chimaltenango; Quezaltenango.
Southern Mexico; British Honduras.
Sometimes a tree of 15 meters with a trunk 40 cm. in diameter; leaves lance-
oblong, green, less rough than in other forms, long-acuminate, rounded at the base,
10 FIELDIANA: BOTANY, VOLUME 24
almost glabrous beneath, the hairs, if any, confined to the veins and closely
appressed.
A form very similar to this and perhaps identical is found in the
West Indies, and there may well be an earlier name for the variety.
MORACEAE. Mulberry Family
Trees or shrubs, usually with milky sap, rarely herbs, sometimes epiphytic;
leaves alternate, stipulate, entire, dentate, or variously lobate; flowers small and
commonly green or greenish, monoecious or dioecious, in ament-like spikes,
capitate, on a flat, entire or lobate receptacle, or sometimes on the inner surface
of a closed receptacle; staminate perianth 2-4-1 obate or 2-4-parted, or the perianth
tubular, sometimes none; stamens usually as many as the perianth lobes and
opposite them, rarely only 1; petals none; pistillate flower with a 3-5-parted
perianth, or sometimes tubular with a small aperture at the apex; ovary superior
or partly inferior, 1-2-celled; styles or stigmas 1-2; ovule solitary, pendulous,
anatropous, or erect and orthotropous; fruit a syncarp of numerous small fruits
upon a usually fleshy receptacle, or the fruits separate and enclosed in the more
or less enlarged and fleshy perianth ; seeds small or large, the endosperm scant or
none; embryo straight or curved, the cotyledons often unequal, usually thick.
About 50 genera, in both hemispheres, most of the species tropi-
cal. Other genera represented in Central America, chiefly in Costa
Rica and Panama, are Olmedia, Perebea, Helicostylis, and Ogcodeia,
the last of which may well extend into Guatemala.
Plants herbaceous Dorstenia.
Plants trees or shrubs.
Flowers borne upon the inner surface of a more or less globose, hollow receptacle,
this having at the apex a small opening closed by scales Ficus.
Flowers variously arranged but never upon the inner surface of a closed recep-
tacle.
Leaves palmately or pinnately lobate or parted, rarely entire (cultivated
tree) but the fruit then a very large syncarp 15-30 cm. long.
Leaves pinnately lobate or rarely entire Artocarpus.
Leaves palmately lobate or parted; fruits small, the individual ones less
than 2 cm. long.
Flowers in dense spikes, these clustered at the end of a peduncle; leaves
peltate Cecropia.
Flowers in cymes; leaves not peltate Pourouma.
Leaves, at least those of adult branches, entire or dentate, the leaves of young
shoots rarely lobate.
Flowers of one or both sexes in ament-like spikes or racemes.
Trees, often armed with spines; pistillate flowers in globose heads, the
staminate in ament-like spikes; leaves dentate Chlorophora.
Trees or shrubs, unarmed; both pistillate and staminate flowers in ament-
like spikes; leaves dentate or entire.
Pistillate perianth of distinct segments, not enclosing the fruit; seeds
minute Morus.
Pistillate perianth tubular, enclosing the fruit; seeds large.
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 11
Staminate perianth none Clarisia.
Staminate perianth present.
Stamens not inflexed in bud Sorocea.
Stamens inflexed in bud Trophis.
Flowers never in ament-like spikes or racemes, arranged in heads or upon
flattened receptacles, or sometimes solitary.
Stipules and usually the leaves armed with small prickles .... Poulsenia.
Stipules and leaves unarmed, the branches rarely armed with spines.
Branches often armed with spines; stamens inflexed in bud; staminate
flowers in ament-like spikes; leaves dentate Chlorophora.
Branches unarmed; stamens not inflexed in bud; staminate flowers
not in ament-like spikes; leaves dentate or entire.
Ovule erect; staminate peduncles bearing several flower heads;
plants often epiphytic; leaves entire Coussapoa.
Ovule pendulous; staminate peduncles bearing a single flower head;
plants not epiphytic; leaves entire or dentate.
Leaves cordate at the base, membranaceous, densely pilose on
both surfaces Castillo,.
Leaves not cordate at the base, usually coriaceous, glabrous or
practically so, at least on the upper surface.
Staminate and pistillate flowers borne upon the same recep-
tacle Brosimum.
Staminate and pistillate flowers borne upon separate recep-
tacles. . . .Pseudolmedia.
ARTOCARPUS Forster
Trees with milky sap; leaves large, alternate, coriaceous, penninerved, per-
sistent, entire or pinnate-lobate, the stipules lateral; flowers monoecious, in
unisexual, globose or oblong, very dense heads, the peduncles axillary, solitary,
the individual flowers very small and numerous, the receptacle becoming fleshy;
staminate perianth with 2-4 lobes or segments, these concave at the apex, slightly
imbricate; stamen 1, the filament erect and usually complanate, the anther short-
exserted; pistillate perianth mostly tubular or obovoid, immersed in the receptacle,
its apex free; ovary straight, included, buried in the receptacle but free from it;
style central or somewhat lateral, the stigmatose apex exserted, linear-spatulate
or rarely subpeltate; ovule affixed near the apex, pendulous; fruiting perianths
numerous, forming with the receptacle a fleshy syncarp; achenes included in the
syncarp, the pericarp membranaceous or coriaceous; endosperm none, the embryo
straight or incurved, the cotyledons fleshy, equal or very unequal.
About 40 species, in tropical Asia, Malaysia, and the Pacific
Islands, one of them now cultivated in all tropical regions.
Artocarpus altilis (Parkinson) Fosberg, Journ. Wash. Acad.
Sci. 31: 95. 1941. Sitodium altile Parkinson, Journ. Voy. Endeavour
45. 1773. A. communis Forst. Char. Gen. 102. 1776. A. incisa L. f.
Suppl. PL 411. 1781. Palo de pan; Arbol de pan; Mazapan; Fruta
de pan; Pan de fruta; Castana (Pete"n, presumably the name applied
to the seeds).
12 FIELDIANA: BOTANY, VOLUME 24
Planted abundantly in the North Coast, in the Pacific plains
and bocacosta, and in the lowlands of Alta Verapaz. Native of the
Pacific islands but now grown in all tropical regions.
A medium-sized or large tree, in Guatemala often 25 meters tall, with thick
trunk and smooth gray bark, the crown very dense; leaves stout-petiolate, 30-80
cm. long, 25-40 cm. wide, dark green above and often sparsely hairy, paler beneath,
scabrous, often pubescent, cuneate and entire at the base, deeply pinnate-lobate,
the lobes acuminate; staminate spikes dense and clublike, 25-40 cm. long; pistillate
inflorescence subglobose, long-pedunculate; fruits large and fleshy, subglobose or
oval, often 30 cm. long, smooth or spiny, with or without seeds.
The name "mazapan" is said to be restricted to the seedless fruits,
and it is these that are most used as food where the trees are grown
for the purpose. Breadfruit is an important food in the Pacific
islands and Malaysia, but in Central America little use is made of it
except in areas where there are people of African origin, as in the
banana regions of the Atlantic coast. They consume the young
fruits in large quantities, usually sliced and fried. The fruits are
said to be eaten at times along the Pacific bocacosta, where there
are many hundreds of giant trees, the largest we have observed in
Central America. The Indian and ladino people, however, have
little taste for breadfruit, and we have not seen it upon the table in
Pacific Guatemala. Both the smooth and spiny forms are planted,
as well as seedless and seed-bearing trees. It is stated with some
authority that all the trees of the Pacific slope have fruits with seeds.
The tree does not grow well except in rather hot regions and it is
rarely if ever seen in the central regions as high as 1,500 meters.
At Coban, for instance, the tree is rarely if ever planted, but fruits
are sometimes brought to the market from the Rio Polochic. The
fruits are used in the Pacific lowlands for fattening pigs, and there
are large quantities of them available about many of the fincas.
The seeds, after having been cooked, also are sometimes eaten by
people. The story of the introduction of breadfruit into the West
Indies, to which it was introduced in order to provide food for the
suffering population, is a long and romantic one, too long to be
repeated here. A good re'sume' of it may be found in Curtis' s Botani-
cal Magazine, under plates 2869-2871, published in 1828. The
story of the expedition of the Bounty under Captain Bligh, to obtain
the plants, has been the subject of many books, short articles, and
moving picture films. The plants were introduced into the West
Indies on the island of St. Vincent in January, 1793. No data are
at hand as to the date at which the breadfruit reached Central
America and Mexico, but it seems reasonable to suppose that it
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 13
may have reached Panama and Mexico one or two centuries earlier,
by the ships that plied every year between those coasts and the
Philippines.
Artocarpus integrifolia L., the jack-fruit, with similar fruits
but entire, mostly ovate leaves, is planted in Guatemala City and
elsewhere. It is a native of the East Indies.
BROSIMUM Swartz
Trees with milky latex; stipules small, lateral, caducous; leaves short-petiolate,
entire, usually coriaceous, penninerved; flowers monoecious, affixed to a globose
receptacle, the receptacles axillary, geminate; bracts closely appressed to the
receptacle, or sometimes none, the bractlets numerous among the staminate flowers,
usually peltate and short-stipitate, before anthesis covering the whole surface,
usually persistent in fruit; staminate flowers numerous, the perianth short-cupu-
late or scarcely distinguishable; stamen 1, the filament short, erect, the anther
small, ovate; staminate flowers 1 or 2 in the center of the receptacle, more or less
immersed, their perianth none or concrete with the receptacle; ovary adnate to the
receptacle, attenuate above to a short style, the stigma branches exserted, thick,
spreading; fruit globose, more or less surrounded by the fleshy or rather dry
receptacle; seed subglobose, the testa membranaceous; endosperm none, the
cotyledons thick, fleshy, subequal, the radicle small, superior.
A group of perhaps 25 species, in tropical America. Three other
species have been reported from southern Central America.
Leaves pale and glaucescent on the lower surface, also minutely and sparsely
sericeous, usually abruptly short-acuminate, commonly lustrous on the upper
surface; pistillate flowers 2 in each receptacle B. panamense.
Leaves almost concolorous, not glaucescent beneath and usually glabrous, not or
scarcely lustrous on the upper surface; pistillate flowers 1 or rarely 2.
Leaves abruptly caudate-acuminate, with a long linear tip .... B. costaricanum.
Leaves merely acute or abruptly short-acuminate, never with a long linear tip.
Leaves brownish or rufescent beneath when dried, the ultimate veins elevated
and prominently reticulate B. terrabanum.
Leaves pale greenish beneath when dried, not at all brownish or rufescent, the
veins not prominent or conspicuously reticulate B. Alicastrum.
Brosimum Alicastrum Swartz, Prodr. Veg. Ind. Occ. 12. 1788.
Ujushte; Ujushte bianco; Masico; Ox (Maya); Ramon; Ramon
bianco; Capomo (British Honduras).
Moist or wet forest, ascending to about 1,000 meters but mostly
at 300 meters or less; Pete"n; Alta Verapaz; Izabal; Escuintla;
Guatemala (valley of Rio Motagua); Retalhuleu; Quiche"; Hue-
huetenango; Baja Verapaz. Southern Mexico and British Honduras;
Salvador; West Indies.
A medium-sized or large tree, sometimes 30 meters tall with a trunk a meter
in diameter, the crown broad and dense, the bark gray; leaves short-petiolate,
14 FIELDIANA: BOTANY, VOLUME 24
coriaceous, bright green when fresh, glabrous, entire, mostly oblong-elliptic to
elliptic, chiefly 7-14 cm. long and 3-5.5 cm. wide, acuminate or abruptly short-
acuminate, sometimes merely acute, obtuse or acute at the base, the lateral nerves
about 14 pairs; flower heads about 1 cm. in diameter, short-pedunculate; fruit
yellow or orange, about 1.5 cm. in diameter, containing a single seed 12 mm. in
diameter.
Called "breadnut" in British Honduras; "ajah," "tsotz ax,"
"ax," "mo," "muju," "talcoite" (Chiapas). Wherever it grows
in quantity, this tree, like some other species of the genus, is much
used as food for stock, especially during the dry season when other
forage is scarce. In Guatemala this is chiefly in Pete*n, but the tree
is still more important for the purpose in British Honduras and
Yucatan. In the latter region it is often the principal food for stock
during the drier months. The branches, of course, must be cut,
and this is done by men who climb the tree with machetes, and cut
down limbs for the stock to browse upon. Mr. J. B. Kinloch states
that the men who do this are more expert tree climbers even than
the chicleros, who are noted in this respect. According to Lundell,
in Pete"n the tree is most abundant on the sites of old Maya villages,
where it forms groves called ramonales. The pulp of the fruit is
edible, and the seeds when boiled are nutritious, with a flavor some-
what like that of potatoes. They are eaten alone or with plantains,
maize, or honey. Sometimes they are roasted and eaten. Also,
they are dried and ground to form a meal, from which a kind of cake
(probably a tortilla) is made, and sometimes are boiled in sirup to
make a sweetmeat. In southern Mexico the roasted seeds are said
to be used sometimes as a coffee substitute. The milky latex, which
flows freely when the trunk is cut, resembles cream and when diluted
with water is said to afford a substitute for cow's milk. There is a
belief in Yucatan that if the seeds are eaten by nursing women the
flow of milk is increased. The wood is described as white or some-
times grayish or tinged with pink, compact, hard, and fine-grained.
It is used at times for construction and other purposes in the Yucatan
Peninsula.
Brosimum costaricanum Liebm. Dansk. Vid. Selsk. Skrivt.
V. 2: 334. 1851. Ojushte; Ajuste; U juste; Albaricoce (Solola); Ramon
Colorado (Alta Verapaz).
Moist or wet, mixed forest, ascending from sea level to about
2,200 meters, but chiefly at low elevations; Alta Verapaz; Izabal;
Santa Rosa; Escuintla; Retalhuleu; Solola; Quezaltenango; San
Marcos. Honduras; Costa Rica; Panama.
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 15
A small, medium-sized, or often very tall tree, sometimes 30 meters high or
more, the trunk 30 cm. or more in diameter, the crown spreading, the bark light
brown, rather smooth, the sap with whitish or yellowish latex, the branchlets
often short-pilose; leaves short-petiolate, oblong-elliptic to lance-oblong, chiefly
8-15 cm. long and 3.5-6.5 cm. wide, caudate-acuminate, with a long linear tip,
acute or obtuse at the base, glabrous above, beneath sparsely strigose or almost
wholly glabrous, the venation elevated and reticulate beneath; flower heads
grayish white, hemispheric or subglobose, about 8 mm. in diameter, the peduncles
stout and usually very short; fruit 1-1.5 cm. in diameter.
Called "masicaran" and "masicaron" in Honduras. The
inflorescences often fall from the trees in great numbers and carpet
the ground. In this state they are sometimes cooked and eaten in
Costa Rica. The tree is one of the most abundant species of the
forest on the plains near Retalhuleu and extends upward on the
hills above the cabecera. It is said that here the seeds are an impor-
tant food during seasons when there is a failure of the maize crop,
being cooked in various ways and eaten. The young inflorescences,
too, are eaten on the Pacific plains of Guatemala. The trees there
are said often to attain a height of 27 meters. They are good for
shade and often are left for this purpose in pastures. The leaves
and flower heads are much eaten by stock as well as by deer and other
wild animals, and by some birds, and the foliage is used by chicleros
as food for stock. The word "ojushte" is of Nahuatl derivation,
probably signifying "trail flower," in allusion to the fallen flowers
seen strewing the trails at some seasons of the year. Two caserios
of Guatemala in the departments of Jutiapa and Santa Rosa are
called Ujuxte" and Ujuxtales, their names derived from that of
this tree. Leaves of seedlings that are assumed to belong to this
species often have undulate, shallowly lobate, or somewhat dentate
or serrate margins. It may be remarked here that, because of lack
of abundant fertile specimens of the genus from Guatemala and
from other parts of Central America the species of Brosimum still
are imperfectly known and their classification is not altogether
satisfactory.
Brosimum panamense (Pittier) Standl. & Steyerm. Field
Mus. Bot. 23: 40. 1944. Piratinera panamensis Pittier, Contr.
U. S. Nat. Herb. 20: 100. pi. 7. 1918.
Wet mixed forest, eastern border of Pete"n, on the boundary of
British Honduras, probably extending into Izabal. Oaxaca; Panama.
A small to large tree, sometimes 25 meters high with a trunk 60 cm. in diame-
ter, the bark grayish, smooth, the sap with latex, the crown narrow or irregular
and depressed; leaves distichous, short-petiolate, oblong or elliptic-oblong, some-
16 FIELDIANA: BOTANY, VOLUME 24
times obovate-oblong, mostly 5-10 cm. long and 2.5-3.5 cm. wide, abruptly short-
acuminate to merely acute or subobtuse, entire, usually very lustrous above,
glaucescent beneath and minutely and sparsely sericeous, the lateral nerves about
10 pairs, divergent at a wide angle; stipules caducous, 3 mm. long; receptacles
solitary, pedunculate, irregularly obconic or in age subglobose, 1-1.5 cm. broad,
the whole surface covered with orbicular peltate bracts; staminate flowers yellow;
pistillate flowers 2; fruit containing 1 or 2 seeds.
Although it has been maintained by some recent authors, Pira-
tinera seems not essentially different from Brosimum, with which
it often has been merged. In typical Brosimum there is only one
pistillate flower, in Piratinera two; but with ordinary herbarium
specimens it is difficult to find even one pistillate flower, to say
nothing of discovering two, and the difference is at least not a
practical one. The tree is called "asta" in Oaxaca. There the inner
bark is described as yellowish brown and yielding a fair quantity
of turbid latex; fruiting in May; sap wood cream-colored but turning
pale brown on exposure to air; heartwood grayish brown or vermilion,
thin; used for railroad ties, boards, and ax handles.
Brosimum terrabanum Pittier, Contr. U. S. Nat. Herb. 18:
69. /. 76. 1914. Piratinera terrabana Lundell, Carnegie Inst. Wash.
Publ. 478: 208. 1937. Masicaran (British Honduras).
British Honduras and Chiapas (near Palenque), and doubtless
extending into Pete*n, Alta Verapaz, or Izabal. Veracruz and
Oaxaca; Honduras; Salvador(?); Nicaragua; Costa Rica; Panama.
A medium-sized or large tree, sometimes 27 meters tall with a trunk 50 cm. or
more in diameter, the crown narrow, irregular, the trunk round and slightly fluted,
sometimes with buttresses a meter high, the bark light gray, fairly smooth; stipules
small, caducous; leaves glabrous, coriaceous, short-petiolate, elliptic-oblong to
narrowly elliptic, mostly 8-14 cm. long and 3-6 cm. wide, acuminate or abruptly
short-acuminate, acute to usually obtuse or narrowly rounded at the base, darken-
ing when dried, often slightly lustrous above, the lateral nerves about 15 pairs, the
margins entire; receptacles in flower 1 cm. or slightly less in diameter, subglobose,
short-pedunculate, with 1 or 2 pistillate flowers, in fruit slightly larger; fruit with
1 or 2 seeds.
Called "masica" and "pisma" on the Atlantic coast of Honduras,
where the wood is used for preparing charcoal and also is sometimes
sawed into lumber. About Tela the seeds are boiled and eaten or
made into a sort of tortilla. Parrots are said to be fond of the fruits.
In Oaxaca and Veracruz the tree is called "ojoche," "ojoche bianco,"
and "ojochillo." The inner bark is white to pinkish brown, and
yields a small amount of thick latex; the sap wood is white, the
heartwood not sharply defined, pale pink or sometimes darker brown.
The fruit is described as reddish pink.
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 17
Cannabis sativa L. Canamo; Marijuana; Mariguana.
This species (of which C. indica L. is a synonym), native of the
Old World but cultivated and introduced in many parts of the earth,
seems to be little known in Guatemala or in other parts of Central
America except Panama. In fact, when we have made inquiries
regarding marijuana (the name by which it is generally known in
Mexico), and explained what it was, the person questioned often has
remarked that he had always wondered what the word meant as it
is used in that most common of street and tavern songs, La Cucaracha.
In many parts of the world Cannabis is cultivated for its fiber, from
which hemp rope is made. It formerly was planted extensively in
the United States for fiber but has been abandoned, largely because
it exhausts the soil rapidly. At the present time it is more known in
the United States as the source of the narcotic hashish or marijuana,
whose production and sale are forbidden in all or most parts of this
country. When the dry plant is smoked, in the form of cigarettes,
mixed with tobacco, it produces hallucinations and often homicidal
mania. The plant is grown clandestinely in the United States and
the cigarettes are peddled in many places, especially to school
children. Its complete suppression is difficult, in part because the
plant is naturalized in many regions and often grows profusely along
roadsides and in waste ground of cities. The seeds are one of the
principal ingredients of bird seed, and it is perhaps on this account
that hemp has become naturalized in so many places.
CASTILLA Cervantes
References: Pittier, A preliminary treatment of the genus Castilla,
Contr. U. S. Nat. Herb. 13: 247-279. 1910; 0. F. Cook, The culture
of the Central American rubber tree, U. S. Dept. Agr. Bur. PI. Ind.
Bull. 49. 1903.
Trees with milky latex; leaves large, deciduous, alternate, short-petiolate,
distichous, entire or denticulate; stipules large, caducous; flowers monoecious,
inserted upon large flat receptacles, these unisexual, covered outside with imbri-
cate bracts; staminate receptacles of two kinds, the primary ones one to several
pairs or sometimes absent, flabellate or compressed, in the axils of leaves or at
defoliate nodes, the complementary ones smaller, clavate or flabellate, always
accompanying the pistillate inflorescences; perianth none; stamens numerous,
irregularly scattered among the bractlets; pistillate receptacles flattened or cup-
iike, the perianth urceolate, with 3-5 short lobes; fruit enclosed in the accrescent,
dry or fleshy perianth.
About 10 species, distributed from western Mexico to the Amazon
Valley of Brazil, Peru, and Bolivia. In Central America 5 species
18 FIELDIANA: BOTANY, VOLUME 24
are known, the others in Honduras, Nicaragua, Costa Rica, and
Panama. The number of species is somewhat indefinite because
the species still are imperfectly understood. The generic name has
sometimes been written Castilloa.
Castilla elastica Cervantes, Gaceta de Literatura de Mexico,
Suppl. July 2, 1794. Ficus gummifera Bertol. Fl. Guat. 40. pi. 9.
1859 (type from Escuintla, Velasquez}. C. lactiflua 0. F. Cook,
Science, n. ser. 18: 438. 1903. C. guatemalensis Pittier, Contr. U. S.
Nat. Herb. 13: 272. 1910 (type from Secanquim, Alta Verapaz, Cook
295). C. gummifera Pittier, Contr. U. S. Nat. Herb. 20: 34. 1917.
Ule; Rule; Cheel k'i'c (Poconchi); Kik (Lacandon); Kiikche
(Quecchi).
Common in dry or wet forest or thickets of the lowlands, some-
times planted in fincas at somewhat higher elevations, chiefly at
300 meters or less, most abundant on the Pacific plains and perhaps
also in Pete"n; Pete*n; Alta Verapaz; Izabal; Santa Rosa; Escuintla;
Suchitepe'quez; Retalhuleu; San Marcos; also growing in the Zona
Reina of Quiche" and Huehuetenango. Tepic to San Luis Potosi and
British Honduras; Salvador, Honduras, Nicaragua, and perhaps
even farther southward.
A medium-sized or sometimes large tree, the branchlets pilose with fulvous,
appressed or ascending hairs; petioles stout, mostly 1-2.5 cm. long; leaf blades
oblong or elliptic-oblong, chiefly 20-45 cm. long and 8-18 cm. wide, abruptly
acuminate, usually shallowly cordate at the base, with a narrow sinus, scabrous-
pilose and rough above, somewhat paler and hirtellous or velutinous-pilose beneath;
primary staminate receptacles mostly in clusters of 6, about 2 cm. broad, the com-
plementary receptacles geminate, pedunculate, claviform, 2-2.5 cm. long; pistillate
receptacles sessile or nearly so, in fruit often more than 5 cm. broad, red or orange-
red at maturity; fruits almost 2 cm. long, the seeds about 1 cm. long.
The trees of this genus are well known as a source of rubber and
are the only native Central American plants from which rubber has
been extracted commercially. Some of the species of southern
Central America, it may be noted, do not produce rubber, or only in
insignificant amounts. For export, Castilla rubber has never attained
great importance in Central America, although it long has been
exported on a small scale from various countries and is still being
exported. The quality and price obtained for it have not encouraged
its cultivation and development, for it is considered greatly inferior
to Hevea rubber. The greater part of the rubber produced in Central
America is used locally, and it is stated that probably no more than
200 tons of it ever were exported in a year from Guatemala at the
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 19
height of the industry. If its price were sufficiently high, no doubt
a larger quantity could be gathered, for the trees are numerous and
widely dispersed in the tierra caliente on both coasts. As a matter of
fact, it is unusual to find a tree that has not already been tapped;
the oblique slashes along the trunk leave huge scars.
In Guatemala, as elsewhere in Central America, the rubber is
much used for making rain capes or coats, which in these tropical
regions are much superior for shedding rain to anything brought
from Europe or North America, because the rubber withstands
combined heat and moisture. The white sap flows freely when any
t part of the tree is cut, and coagulates upon exposure to air. To
hasten its coagulation various substances often are added to it,
particularly the sap of Calonyction and other Convolvulaceae.
The extraction and uses of rubber were well known to the aborigi-
nal inhabitants of Central America and Mexico, who used it for
waterproofing articles of clothing, bottles, etc. They also made from
it large balls that were used in the game of pelota, played somewhat
like basketball. The ball was thrown through large stone rings
inserted high in the walls of courtyards, but the ball was manipu-
lated by catching it upon the hips and tossing it into the ring without
touching it with the hands. The stone rings may be found now in
some of the ruined Maya temples. A similar or the same game is
sometimes played at the present time, but in most places has been
forbidden by law because of the danger to the players. It was
through the use of the ball in such games that rubber first became
known to the Old World, for the games must have been observed
by the earliest Spaniards who visited Mexico.
The Guatemalan rubber trees are rather handsome, some of
them attaining a great height, especially in the North Coast. Even
from a distance they are easily recognizable because the very large,
soft leaves are 2-ranked and droop limply along each side of the
spreading or often pendent branches. The mature fruits are con-
spicuous because of their bright coloring. The trees usually lose
their leaves toward the end of the dry season (in the spring of the
North) and produce their flowers at the same time. One of the
rivers of Guatemala bears the name Ulapa, said to signify "river
of ule trees."
Tozzer states that the sap of the rubber trees was used as incense
among the Lacandon Indians but probably this is an error, for no
one who knows the odor of burning rubber would consider it a
pleasing offering to any god. The bolillos with which Guatemalan
20 FIELDIANA: BOTANY, VOLUME 24
marimbas are played are made from Castillo, rubber. The Maya
names reported from Yucatan are "yaxha" and "kiikche."
The crown of the rubber tree is either rounded or spreading or,
when the trees are crowded, tall and narrow; its trunk is sometimes
buttressed; the bark is light brown or light gray. The wood is
creamy white throughout, or the heartwood light brown, without
distinctive odor or taste; it is light and soft, although firm, with
fairly straight grain, rather coarse in texture, easy to cut; it is rather
tough and strong for its weight, but is not durable. Little or no use
is made of it in Central America. One peculiarity of the tree is its
extensive root system, the roots being near the surface of the ground,
where sometimes they may be traced for 30 meters.
GECROPIA L.
Trees or large shrubs with milky sap, the trunk simple or usually sparsely
branched, smooth, whitish, hollow, with cross partitions at the nodes; stipules
large, deciduous; leaves large, long-petiolate, peltate, palmately lobate or parted,
usually scabrous and rough on the upper surface and white-tomentose beneath,
the lobes mostly entire; flowers minute, dioecious, in very dense, cylindric, short
or elongate spikes, these few or numerous, sessile or pedicellate, digitate at the end
of a short or elongate, axillary peduncle; staminate perianth tubular or campanu-
late, entire or 2-cleft; stamens 2; pistillate perianth thin, with a small aperture at
the apex; ovary included, with very short style, the stigma exserted, penicillate;
ovule erect, orthotropous; fruit oblong, included in the very thin perianth, the
exocarp very thin or obsolete, the endocarp crustaceous or hard; seed with mem-
branaceous testa; endosperm none, the cotyledons oblong or ovate, equal, straight,
the radicle small, superior.
Probably 50 species or more, in tropical America. A few other
species are known from other parts of Central America. The
Central American species, like those of other regions, are imper-
fectly known because of lack of ample material for their study, and
it is not known just how many species really are represented in this
area.
All species of Cecropia are much alike' in general appearance, and
are distinguished by their smooth whitish trunks, few branches, and
large, deeply palmate-lobed leaves with often snowy white under
surfaces. No tree is more exotic in appearance to one coming from
the North, and not even palms are more important in giving to the
lowland vegetation of Central America its distinctive facies. The
hollow trunks and branches usually but not invariably are inhabited
by ants that bite severely when the tree is molested. Spruce states
that in the Amazon region the hollow trunks often are inhabited
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 21
by bees. The branches are said to have been used by some of the
American aborigines for making trumpets, hence the English name
"trumpet tree" often applied to the genus. The split trunks some-
times are employed as troughs or conduits for conducting water.
The bark contains a tough fiber utilized in some regions (not in
Central America, so far as known) for making cordage, mats, and a
kind of coarse cloth. The stems have been used in Brazil for making
paper. The sap contains a kind of rubber but in too small quantities
for commercial purposes. It is reported that some South American
Indians ate the pith of the branches. The trees have been much
used in domestic medicine but no definite properties seem to be
ascribed to them.
Guarumo (the usual name in all Central America) trees grow
rapidly, like weeds, and thrive best in cut-over or abandoned land.
In many regions they are almost if not quite confined to second-
growth thickets, but some species are seen in primeval forest. In
Guatemala this is true of C. sylvicola, which has not been noted out-
side the wet forest of the Tactic region.
Pistillate spikes long and slender, mostly 15-30 or even 40 cm. long. . C. obtusifolia.
Pistillate spikes short, mostly 3-5 cm. long.
Leaves white beneath, covered with a dense cobwebby tomentum, membrana-
ceous C. peltata.
Leaves green beneath, appearing glabrous but with a very minute, close, sparse
tomentum between the veinlets, coriaceous C. sylvicola.
Cecropia obtusifolia Bertoloni, Fl. Guat. 439. 1840. C.
mexicana Hemsl. Biol. Centr. Amer. Bot. 3: 151. pi. 80. 1883.
C. mexicana var. macrostachya Donn. Smith, Bot. Gaz. 27: 442. 1899
(type from Costa Rica). Guarumo (often modified, especially about
Coban, to Guarumbo) ; Pad, Choop (Coban, Quecchi) ; Xobin (Baja
Verapaz, fide Tejada).
Common through most of the lowlands, usually in wet or moist
thickets, sometimes in wet forest, even in Manicaria swamps,
frequent along borders of pastures or forest, ascending from sea
level to (in the Occidente) about 1,300 meters; Pete"n(?); Alta
Verapaz; Baja Verapaz; Izabal; Santa Rosa; Escuintla (type from
Escuintla, Velasquez); Suchitepe"quez; Retalhuleu; Sacatepe'quez ;
Chimaltenango; Huehuetenango; Quezaltenango; San Marcos.
Southern Mexico and British Honduras throughout the lowlands
of Central America to Panama.
A small to large tree, sometimes 22 meters tall but usually much lower, the
trunk seldom more than 30 cm. in diameter, the branchlets very stout and thick;
22 FIELDIANA: BOTANY, VOLUME 24
stipules large, whitish-pubescent or glabrate; leaves on very long, terete petioles,
suborbicular in outline, 30-50 cm. wide or larger, cleft about halfway to the base
into usually 10-13 lobes, green and scaberulous above, densely white-tomentulose
beneath or sometimes glabrate, the lobes entire, broad or narrow, rounded or
abruptly short-acuminate at the apex; spathe at the base of the inflorescence
white-tomentose or rarely glabrate, closed and pointed before an thesis; staminate
peduncles elongate, the spikes few, 3-4 mm. thick, long and slender; pistillate
spikes usually 2-4 or sometimes more, sessile or nearly so, mostly 20-40 cm. long
and 6-7 mm. thick, in fruit very fleshy.
This is an abundant and characteristic tree almost throughout
the Pacific plains, and almost equally so in the North Coast. It is
easily distinguished from C. peltata by the very long and pendent
flower spikes. The leaves, especially young ones, often are tinged
beneath with red or purple but this coloring is not very conspicuous.
Trees brought from the Pacific coast have been planted in Guate-
mala City, where they seem to grow well. This species sometimes
flowers when only a shrub of 4 meters. The leaves are eaten by
stock, and in Salvador the leaves are salted, after which cows are
said to eat .them in quantity. The wool separated from the stems
and leaves is said to be sometimes smoked by the Indians of Alta
Verapaz, like tobacco. Velasquez, in notes accompanying the origi-
nal specimens of C. obtusifolia, remarks that it is on this tree that the
bird called "ciacia" (chacha or chachalaca) builds its nests. The
local name, "guarumo," gives its name to a caserio of San Marcos,
called El Guarumo. In British Honduras the tree is called "trumpet."
The name "guarumo" is probably of West Indian origin. Oviedo
cites it as "yaruma," which probably is closest to the original form
of the word.
Gecropia peltata L. Syst. ed. 10. 1286. 1759. C. asperrima
Pittier, Contr. U. S. Nat. Herb. 19: 227. 1917. Guarumo; Igarata,
Ix-coch (Maya); Trumpet (British Honduras); Ixcochle (Pete"n).
Chiefly in pastures or second-growth, often in thickets or modified
forest, at 900 meters or less; Pete"n; Izabal; Santa Rosa. Yucatan and
British Honduras; Honduras; Nicaragua; Costa Rica; West Indies;
northern South America.
A small or medium-sized tree, attaining sometimes a height of 20 meters;
petioles often longer than the leaf blades, these suborbicular in outline, 30-50 cm.
wide or larger, mostly 7-9-lobate, shallowly or deeply lobate, dark green and sca-
brous above, rough to the touch, densely covered beneath with a white, often
snowy tomentum, or sometimes greenish and only sparsely tomentose; spathes
about 6 cm. long, cuspidate at the apex, caducous; staminate spikes numerous,
about 4 cm. long and 3 mm. thick, short-pedicellate; pistillate spikes usually 2-6,
sessile, yellowish at first, 3-6 cm. long, in fruit very thick and succulent.
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 23
The Central American tree has never, so far as we know, been
referred to the common West Indian C. peltata, but there are no
apparent characters by which two species may be distinguished in
the fairly ample material at hand. Specimens from Guatemala and
Yucatan have been referred in the past to C. obtusa Tre'cul and C.
Humboldtiana Klotzsch. The wood is whitish or light-colored, very
light and soft, with a specific gravity of about 0.45, with straight or
fairly straight grain, coarse- textured, easy to cut, tough and strong
for its weight, but perishable. So far as known, no use is made of it
in Central America. The Maya name reported from Yucatan is
"xco-che."
Cecropia sylvicola Standl. & Steyerm. Field Mus. Bot. 23: 153.
1944.
Known only from the type region, dense wet mixed forest,
mountains along the road between Tactic and the divide on the road
to Tamahu, about 1,500 meters; type, Standley 90762.
A tall tree, growing in primeval forest, abundantly branched above; leaves
large, coriaceous, long-petiolate, the petioles terete, multicostulate, as much as
50 cm. long or even longer, densely hirtellous; leaf blades suborbicular, about
9-lobate almost to the base, finely scabrous above, paler beneath, hirtellous or
hispidulous on the nerves and veins, in age glabrate but minutely tomentulose
between the veins, not whitened, the lobes oblong or obovate-oblong, as much as
40 cm. long and 11 cm. wide, very obtuse at the apex; pistillate inflorescence
borne on a stout peduncle 3.5-6 cm. long, the spikes numerous, crowded, on stout
pedicels almost 1 cm. long, the spikes 5-5.5 cm. long, 5-6 mm. thick, rounded at
the apex.
The tree is apparently rare and very local. It is rather plentiful
in the one locality where it has been found, but all the trees were
so tall that it was impossible to reach the branches by ordinary
means.
CHLOROPHORA Gaudichaud
Trees with white latex, often armed with spines; leaves alternate, petiolate,
entire or dentate, penninerved; stipules lateral, caducous; flowers dioecious, the
staminate in long slender dense ament-like spikes, the bracts small; pistillate
inflorescence capitate, globose or oblong; bracts similar to the perianth segments
and of equal length; staminate perianth 4-parted, the segments broad, obtuse,
slightly imbricate; stamens 4, the filaments inflexed in bud, porrect and exserted
in anthesis; pistillate perianth 4-fid or 4-parted, the segments concave and thick-
ened at the apex; ovary included, oblique, the style sublateral, filiform, usually
simple; ovule laterally affixed, descending; fruiting perianths fleshy, forming a
globose or oblong syncarp; achene equaling the perianth or somewhat exserted,
ovate, compressed, oblique at the apex, the pericarp coriaceous; seed with mem-
branaceous testa; endosperm none, the embryo incurved, the cotyledons ovate.
24 FIELDIANA: BOTANY, VOLUME 24
Probably three species, one in Africa, one in Mexico, and the
following:
Chlorophora tinctoria (L.) Gaud, in Freyc. Voy. Bot. 508. 1826.
Moras tinctoria L. Sp. PI. 986. 1753. Mora.
Moist or usually dry thickets or forest in the tierra caliente,
common in the plains and lowlands of the Oriente and the Pacific
coast, 1,200 meters or less; Pete'n; El Progreso; Chiquimula; Jutiapa;
Santa Rosa; Escuintla; Suchitep^quez; Retalhuleu; Huehuetenango;
doubtless in all the Pacific coast departments. Southern Mexico
to British Honduras and Panama; West Indies; South America.
Sometimes only a shrub but usually a tree, sometimes 20 meters tall with a
trunk 30-60 cm. or more in diameter, sometimes with buttresses, the bark light
brown with numerous lighter excrescences, the branches often armed with stout
sharp axillary spines; leaves deciduous, membranaceous, short-petiolate, oval to
ovate or ovate-oblong, 5-10 cm. long, usually cuspidate-acuminate, obtuse to
subcordate at the base, entire or serrate, often deeply lobate on young branches,
glabrous or nearly so; staminate spikes cylindric, 4-12 cm. long and about 4 mm.
thick, whitish or greenish, short-pedunculate; pistillate heads spheric, 6-10 mm.
in diameter; fruits 1-1.5 cm. in diameter, globose; styles very long and thread-
like.
Known in Tabasco as "lora de clavo," "mora lisa," and "palo
amarillo." The wood is of various shades of yellow, lustrous, becom-
ing reddish or brownish on exposure; sap wood white, sharply defined;
without distinctive odor or taste; hard and heavy, with fairly straight
or somewhat interwoven grain, medium to coarse in texture, not
very difficult to work, finishes smoothly, and is tough, strong, and
durable. It is sometimes used in regions where plentiful for interior
finish, cart wheels, and other purposes, but its principal value is as a
dyewood. It is the fustic of commerce, long an important export
from tropical America to the United States and Europe, the wood
being exported chiefly from the Antilles but also from Mexico,
Honduras, Nicaragua, Costa Rica, and Panama. The coloring
principle, maclurin, gives a yellowish brown or khaki color much
used for military uniforms. With other dyes it gives various colors
for cotton and silk materials, and also a permanent black. During
wars it usually is much in demand. In Salvador, and probably also
in Guatemala, the wood is utilized for railroad ties, posts, wheels,
wooden balls, and other articles. By the Indians of Guatemala it
is used to color wool yellow or olive-drab. The bark is bitter and
has a disagreeable odor.
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 25
CLARISIA Ruiz & Pavon
Reference: J. Lanjouw, Recueil Trav. Bot. Ne"erl. 33: 254-276.
1936.
Trees or shrubs with milky sap; leaves alternate, short-petiolate, entire or
dentate, membranaceous to coriaceous, penninerved; flowers dioecious; staminate
inflorescences spicate, pedunculate, axillary and simple or forming short raceme-
like inflorescences; staminate flowers consisting only of one stamen, more or less
arranged in rows, intermixed with bracts, these often peltate, the spike usually
having on one side a naked strip without flowers and bordered by two rows of
peltate bracts; filaments erect and straight in bud; pistillate flowers racemose or
capitate, the inflorescences axillary, sessile or pedunculate; pistillate perianth
ovoid, adnate to the ovary, with a small, often irregularly lobate apical opening;
ovary inferior or semi-inferior; style short, the two stigmas exserted, short or
elongate; fruit globose or ovoid, included in the enlarged perianth, the pericarp
membranaceous; seed subglobose, the testa membranaceous; endosperm none;
embryo erect, the cotyledons thick, fleshy, equal.
About eight species, distributed from Mexico to southern Brazil.
A single species is known from Central America and Mexico.
Clarisia mexicana (Liebm.) Lanjouw, Recueil Trav. Bot.
N4erl. 33: 270. /. 3, B. 1936. Sahagunia mexicana Liebm. Dansk.
Vid. Selsk. Skrivt. V. 2: 316. 1851.
Moist or wet, mountain forest, or in lowland forest, 2,450 meters
or less; Suchitepe"quez; Quezaltenango; San Marcos; Huehuetenango.
Veracruz.
A tree 30 meters tall, the trunk 60-100 cm. in diameter, the bark smooth,
brown, the young branchlets sparsely appressed-pilose; leaves membranaceous, on
petioles 5-10 mm. long, oblong or elliptic-oblong, 8-19 cm. long, 2.5-5 cm. wide,
short-acuminate, acute or obtuse at the base, entire, glabrous, with 12-14 pairs
of lateral nerves; staminate spikes arranged in a small panicle or raceme, 1.5-3
cm. long; bracts usually spatulate or obliquely peltate; flowers greenish white;
pistillate flowers usually two together in the leaf axils, the pedicels 2-3 mm. long;
perianth 3-4 mm. long, almost glabrous; styles 5-6 mm. long.
COUSSAPOA Aublet
Trees or shrubs with milky sap, usually epiphytic when young, in age often
standing alone, sometimes somewhat scandent; leaves mostly large and coriaceous,
alternate, petiolate, entire or undulate, the stipules small or large, caducous;
flowers dioecious, globose-capitate, the heads sessile or pedunculate, the bracts
filiform below, spatulate or peltate at the apex; staminate perianth tubular or
subclavate, 3-dentate or 3-parted, the lobes imbricate; stamens 1-2, the filaments
connate into a column, the anthers ovate; pistillate perianth tubular or clavate,
with a minute aperture at the apex or 3-dentate; ovary and style included, only
the stigma exserted, it subpeltate, penicillate-capitate; ovule erect, orthotropous;
perianths slightly accrescent in fruit; fruit included in the perianth, the pericarp
26 FIELDIANA: BOTANY, VOLUME 24
fleshy or succulent, the endocarp crustaceous or harder, separable into two valves;
seed with a membranaceous testa, the cotyledons ovate or oblong, the radicle
short, superior.
About 30 species, chiefly in South America. A few besides those
listed here occur in other parts of Central America. In habit the
trees are like Ficus, but they seem to be less aggressive and are
usually much rarer than trees of that genus.
Leaves glabrous beneath or essentially so C. Purpusii.
Leaves sparsely or usually densely tomentose beneath.
Leaves narrowly oblong to elliptic-oblong, mostly 3-6 cm. wide . . C. oligocephala.
Leaves chiefly ovate or broadly ovate, commonly more than 10 cm. wide.
C. panamensis.
Coussapoa oligocephala Bonn. Smith, Bot. Gaz. 40: 11. 1905.
Cop 6 zotz (Pete"n, fide Lundell).
Moist or wet forest, 350 meters or less; Pete"n; Alta Verapaz
(type from Cubilgiiitz, Tuerckheim 8659); Izabal. Tabasco; British
Honduras.
A large epiphytic shrub or tree, or often an independent tree (probably after
death of the host) sometimes 30 meters high with a trunk 60 cm. or more in diame-
ter, when cut exuding a yellow latex; leaves on petioles 1.5-3.5 cm. long, the blades
narrowly oblong to elliptic-oblong, mostly 10-15 cm. long and 2-6 cm. wide, some-
times larger, obtuse or subacute, rounded or emarginate at the base, glabrous
above, whitish or grayish beneath and arachnoid-tomentose, the lateral nerves
9-11 pairs, ascending at a very narrow angle; staminate peduncles bearing 3-5 or
more heads, these pedunculate, 4-5 mm. broad, yellow or whitish; pistillate
peduncles mostly 2.5-3.5 cm. long, bearing a single globose head about 1 cm. broad.
Coussapoa panamensis Pittier, Contr. U. S. Nat. Herb. 18:
226. 1917.
Wet forest, sometimes in wooded swamps, at or little above sea
level; Izabal. Honduras; Costa Rica; Panama.
An epiphytic or terrestrial tree, sometimes 30 meters tall but usually lower,
the branchlets glabrous or nearly so; stipules caducous, 2-4 cm. long or larger,
acuminate, tomentose or sericeous; leaves long-petiolate, coriaceous, the blades
broadly ovate or elliptic-ovate, chiefly 10-30 cm. long and 7-15 cm. wide, obtuse or
rounded at the apex and apiculate, rounded to subcordate at the broad base,
glabrous above or nearly so, whitish or grayish beneath and closely tomentulose,
the lateral nerves about 15 pairs, almost straight, ascending at a rather wide angle;
staminate peduncles about equaling the petioles, cymosely branched and bearing
several globose heads 5 mm. in diameter; pistillate peduncles 4-6 cm. long, bearing
a single globose head 1.5-2.5 cm. broad.
Called "matapalo" in Honduras. The heartwood is pinkish
gray or oatmeal-colored, becoming somewhat yellowish upon
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 27
exposure, the sapwood not distinguishable from the heart, without
distinctive odor or taste, of medium density and hardness, with
straight or somewhat irregular grain, rather coarse- textured, fairly
easy to work, finishes smoothly, is not durable.
Coussapoa Purpusii Standl. Field Mus. Bot. 8: 6. 1930.
Matapalo.
Moist mixed mountain forest of the Occidente, 900-1,800 meters;
Quezaltenango; San Marcos. Jalisco to Veracruz and Chiapas.
An epiphytic or terrestrial tree, sometimes 18 meters high, with a trunk 15 cm.
or more in diameter, the branchlets fuscous-ferruginous, glabrous; stipules acumi-
nate, glabrous or minutely puberulent, about 2 cm. long; leaves on long slender
petioles, the blades broadly elliptic or oval to ovate-elliptic or lance-elliptic, mostly
9-14 cm. long and 4-7 cm. wide, acuminate at the apex or rounded and abruptly
short-pointed, rounded at the base or subemarginate, green above and lustrous,
paler beneath, glabrous, 5-nerved at the base, the lateral nerves about 5 pairs;
staminate peduncles equaling or longer than the leaves, branched and bearing
mostly 4-5 globose pedunculate heads 5-6 mm. broad; pistillate peduncles 2-4 cm.
long, slender, stiff, glabrous, each bearing a single globose many-flowered head
1 cm. in diameter.
DORSTENIA L.
Perennial herbs with milky sap, with rhizomes, acaulescent or with somewhat
elongate stems; leaves very variable, usually long-petiolate, mostly membrana-
ceous, entire, dentate, angulate, or pinnate-lobate; flowers minute, monoecious,
densely crowded on a usually large, explanate, commonly saucer-like, entire or
angulate or lobate receptacle, the receptacles axillary, long-pedunculate, the flowers
of both sexes numerous and intermixed, the pistillate flowers usually surrounded
each by 3-4 staminate ones, the bracts minute and inconspicuous; perianths com-
monly connate with the receptacle, their margins sometimes obscurely bilobate or
bidentate; stamens 2, rarely 1 or 3, the filaments at first inflexed, finally porrect
and exserted; ovary included, the style excentric or almost lateral, exserted, 2-fid,
the short branches subulate; fruits very small, finally protruded from the pits of
the receptacle, the exocarp fleshy, the endocarp crustaceous; testa of the seed
thin-membranaceous; endosperm none; cotyledons subequal, embracing the ascend-
ing radicle.
About 50 species, mostly in tropical America and Africa, one or
more in eastern Asia. No other species known in Central America.
Plants with elongate, erect or ascending, herbaceous stems.
Receptacles hispidulous on the lower surface; leaves mostly obtuse, not lobate.
D. Lindeniana.
Receptacles glabrous on the lower surface; leaves acute or acuminate, often
pinnate-lobate D. choconiana.
Plants acaulescent or practically so, never with elongate stems.
Receptacles orbicular or oval, entire D. Drakena.
Receptacles more or less quadrate, often deeply lobate D. Contrajerva.
28 FIELDIANA: BOTANY, VOLUME 24
Dorstenia choconiana Wats. Proc. Amer. Acad. 22: 477. 1887.
Usually in dense wet forest at or near sea level; type collected
"in dry stream beds in the forest at the head of Black River, a
branch of the (Rio) Chocon," Sereno Watson; Izabal. Atlantic
lowlands of Costa Rica.
Plants with elongate rhizomes, the stems erect, 15-30 cm. tall, stout, the whole
plant glabrous or nearly so; leaves long-petiolate, 10-15 cm. long, shallowly or
deeply pinnate-lobate, often blotched with silver on the upper surface, the lobes
usually 7 or 9, acute or acuminate, entire, spreading or ascending; receptacles pale
green, long-pedunculate, glabrous, turbinate, 1.5-3 cm. broad, entire.
Dorstenia choconiana var. integrifolia Donn. Smith, Bot.
Gaz. 13: 76. 1888.
Dense wet mixed forest, often on steep stream banks, ascending
from sea level to about 1,600 meters; Alta Verapaz (type from
Pansamala, Tuerckheim 751); Baja Verapaz; Izabal. Honduras;
Costa Rica.
Similar to the species and apparently much more common; foliage very dif-
ferent from that of the typical form, the blades lanceolate to lance-oblong or
oblong, 7-20 cm. long, usually long-acuminate, truncate or rounded at the base,
entire or nearly so.
Dorstenia Contrajerva L. Sp. PI. 121. 1753. Contrahierba;
Mano de leon (Quezaltenango) ; Hierba de sapo (Pete"n); Cambahan
(Pete"n, Maya); Contaul (Chimaltenango, fide Tejada).
Moist forest or thickets, ascending from sea level to about 1,800
meters; Pete"n; Alta Verapaz; Chiquimula; Jalapa; Santa Rosa;
Escuintla; Guatemala; Sacatepe'quez; Chimaltenango; Retalhuleu;
Quezaltenango; Huehuetenango. Mexico and British Honduras to
Panama; West Indies; South America.
Plants acaulescent or nearly so, the stems, if any, very short; leaves often
very numerous and crowded, long-petiolate, deeply and pinnately or almost
palmately lobate, sparsely scabrous or puberulent, usually somewhat rough to the
touch, the lobes acute to acuminate, narrow or broad; receptacles on long slender
peduncles, quadrangular or deeply and irregularly lobate, accrescent in age and
2-5 cm. wide, scaberulous beneath.
Maya names reported from Yucatan are "xcambalhan" and
"cabalhau." The plant is well known in Central America because
of its use in domestic medicine. It is a common household remedy
for dysentery and is also employed in treating bites of poisonous
animals of all kinds. The name "contrahierba," employed by
Linnaeus as the specific name of this widespread species, usually
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 29
is used in Spanish to designate plants of supposed outstanding value
as counteragents for poisons. The aromatic rootstocks are much
used in Salvador, and probably also in Guatemala, for flavoring
cigarette tobacco.
Dorstenia Contrajerva var. Houston! (L.) Bureau in DC.
Prodr. 17: 259. 1873. D. Houstoni L. Sp. PI. ed. 2. 176. 1762.
Contrahierba; Hierba de loro (fide Aguilar).
Moist forest or thickets, often a weed in cafetales, ascending from
sea level to about 1,100 meters; Pete"n; Alta Verapaz; Izabal; Santa
Rosa; Sacatepe"quez; Chimaltenango; Quiche; Quezaltenango.
Widely distributed, like the typical form of the species.
Like the species except in leaf form, the blades large or small, usually ovate-
cordate or triangular-cordate, acute to long-acuminate, subentire or undulate or
crenate, often somewhat hastate-angulate.
This scarcely deserves varietal designation, being nothing more
than a leaf form and often growing with lobate-leaved plants, quite
possibly even from the same root.
Dorstenia Contrajerva var. tenuiloba (Blake) Standl. &
Steyerm. Field Mus. Bot. 23: 40. 1944. D. Contrajerva subsp.
tenuiloba Blake, Contr. U. S. Nat. Herb. 24: 2. pi. 1. 1922.
Contrahierba.
Type collected in damp forest along trail from Los Amates to
Izabal, Izabal, lower slopes of Sierra de las Minas, Blake 7803;
collected also in Chimaltenango, Huehuetenango, and San Marcos,
600-1,000 meters.
Differing from the species in having the leaves very deeply lobate, with
exceptionally long and narrow lobes.
This is a feebler variety even than the preceding and less worthy
of special designation.
Dorstenia Drakena L. Sp. PI. ed. 2. 176. 1762. Contrahierba.
Moist or wet, open or forested banks or slopes, mostly at 200-
900 meters; Alta Verapaz; Izabal; Zacapa; Chiquimula; Jutiapa.
Mexico, the type from Veracruz; Nicaragua; Costa Rica; South
America.
Plants acaulescent, with short thick rhizomes; petioles long and slender,
sometimes 20-25 cm. long; leaves membranaceous, sparsely puberulent, often
rough to the touch, very variable in form, often cordate-ovate and long-acuminate,
frequently pinnate-lobate or angulate, mostly 10-25 cm. long, the lobes often
30 FIELDIANA: BOTANY, VOLUME 24
sinuate or dentate; receptacles long-pedunculate, orbicular or oval, 2-4 cm. broad,
puberulent beneath, rather thin and almost flat, excentrically peltate.
The species is used like D. Contrajerva, which it much resembles
except in the form of the receptacles. The two species can not be
separated by foliage alone. Although in this species there are found
the same leaf variations as in D. Contrajerva, apparently they have
not been designated by name.
Dorstenia Lindeniana Bureau in DC. Prodr. 17: 269. 1873.
Contrahierba.
Wet mixed forest, 1,100 meters or less; Pete"n; Alta Verapaz;
Izabal. Tabasco, the type from Teapa; British Honduras.
Plants with slender rhizomes, the stems erect or decumbent, 10-20 cm. long,
hispidulous; petioles about half as long as the blades, these obovate or oblong-
obovate, mostly 7-10 cm. long, obtuse or rounded at the apex or sometimes sub-
acute, narrowed to the cordate base and with a deep narrow sinus, sinuate-denticu-
late, sparsely hispidulous; peduncles about equaling the petioles, hispidulous;
receptacle rounded, cyathiform, dentate, 7-10 mm. broad.
The leaves often are blotched with silver on the upper surface.
FICUS L. Fig
Reference: Standley, The Mexican and Central American species
of Ficus, Contr. U. S. Nat. Herb. 20: 1-35. 1917.
Trees or shrubs with milky sap, often epiphytic or scandent; leaves alternate,
petiolate, entire in American species; stipules mostly caducous; flowers monoecious,
inserted on the inner surface of a usually globose, fleshy receptacle, this with a
small opening (ostiole) at the apex, the opening closed by several small scales;
receptacle subtended at the base by a lobate involucre; staminate perianth of 2-6
small segments; stamens usually 1-2; fruit of numerous small achenes crowded
over the inner surface of the usually succulent and juicy receptacle.
Probably 600 species or more, widely distributed in tropical
regions. Several additional ones are known from other parts of
Central America. It is probable that some further species remain
to be listed for Guatemala since a few sterile specimens may repre-
sent species not found in the following list.
Most or all the native Central American figs have a characteristic
habit of growth, although the same habit is found also in other
tropical groups, such as Clusia and Araliaceae. They are hemi-
parasites; that is, they often or usually begin growth upon other
trees, frequently upon palms, germinating and developing a stem
from which aerial roots descend to the ground and take root. Young
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 31
plants are consequently often vine-like. With age the aerial roots
and the stems increase in size, ultimately forming a shell-like trunk
that envelops the host plant. The stems at first are flat, broad, and
thin, and as they increase in size several will unite, assuming fan-
tastic and often serpent-like forms. Finally the host plant dies,
but it often survives for a long time and one may see the top of a
palm or some other tree rising above the crown of a large fig. Trees
of this type are known usually in Central America by the term
"matapalo." Large fig trees often send down from their branches
cord-like aerial roots that may take root in the soil and develop into
secondary trunks, thus forming trees, often of enormous size, of the
banyan type, best developed, apparently, in India. Banyan trees
are scarcely if at all known in Central America but in Mexico some
species occasionally develop thus.
Because of their broad dense crowns and handsome foliage, many
of the native figs make fine shade trees and they often are planted for
this purpose in Central America. A few exotic species also are
planted for shade or ornament.
The fruits of all native species are edible, but generally they are
so small and dry that they are not very palatable. Birds and domes-
tic animals are fond of them, and they are much sought by such
birds as toucans and parrots and by monkeys. The latex, often
copious, contains a kind of rubber that may some day be utilized.
From the bark, pounded into thin sheets, the ancient Mexicans
and probably also the Guatemalans obtained a kind of paper
upon which their manuscripts were written. In many parts of
Guatemala, as about Antigua, leafy branches are cut and given
as fodder to cattle when pasture is poor. The wood is light, soft, and
of little value even for fuel, but canoes are said to be made sometimes
from the trunks. The small seeds (achenes) are spread widely by
birds, and it is doubtless thus that the epiphytic habit of most of
the species has been established. It is worthy of note that terrestrial
seedlings of the white figs (subgenus Pharmacosyce) are abundant
in the forests, the seedlings developing rapidly and never becoming
epiphytes. Fig trees are little infested by epiphytes, possibly because
their bark is ordinarily smooth and does not afford a good lodging
place for seeds.
The names "higuero" and "higueron" as well as "matapalo"
(tree-killer) are given commonly to the wild figs, but in mere general
use in Central America is the term "amate," from the Nahuatl
amatl, signifying "paper." The Nahuatl term appears naturally in
32 FIELDIANA: BOTANY, VOLUME 24
many place names, notably Amatitlan and its lake in the Depart-
ment of Guatemala, and Los Amates in Izabal. Amatepeque
(fig mountain) is an aldea of Jutiapa, while the diminutive Ama-
tillo also is used as a place name. The caserio Cuxapa of Jalapa
derives its name from the Nahuatl cux, a kind of fig, and apan, "in"
or "on the water." In Salama the Pipil name for the genus Ficus
is dmat, a modification of the Nahuatl term. Fig trees figure com-
monly in poetry and romance of Central America, and the trees,
occurring as they do about many dwellings, become intimately
associated with daily life and often are regarded with affection.
The amate is called the national tree in Salvador. Village markets
of ten are held in the ample shade of some giant fig tree, although
the larger ceiba is preferred.
Wisdom reports the following curious belief among the Chorti
Indians of the Jocotan region of Chiquimula: "The flower of the
amate tree is a talisman and assures its owner of lifelong happiness,
good health, success in love- and money-making, and safety from
the harm of sorcerers and evil spirits. He will also possess bravery
and boldness, will be invulnerable to all harm, and will be able to
dominate all wild animals, even poisonous snakes. The tree is said
not to possess visible flowers, being reproduced by spores, but the
curers insist that it produces a single flower each year. It becomes
visible on a Friday at midnight, at which time an evil spirit, usually
the Devil, suddenly appears and seizes it for himself. The tree
from which it is taken must be deep in the forest, far from any
habitation, and it can be obtained only when it falls to the ground.
It is said that many men have tried to get one of these flowers but
have failed, owing to their being stricken with terror upon seeing
the evil spirit. They immediately sickened from fright, and some
are believed to have died."
Leaves deeply lobate; cultivated species F. Carica.
Leaves entire.
Cultivated species; plants scandent, or receptacles oblong, or the leaves broadly
obovate and deeply cordate at the base.
Plants creeping and scandent F. pumila.
Plants trees.
Leaves acuminate, not cordate at the base . . . F. elastica.
Leaves broadly rounded at the apex, deeply cordate at the base.
F. pandurata.
Native species, never with any of the three characters mentioned above.
Receptacles solitary; involucre 3-lobate; stamens 2; leaves often scabrous.
Subgenus Pharmacosyce.
Leaves hirtellous or short-pilose beneath; receptacles pilose or hirtellous.
F. glaucescens.
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 33
Leaves glabrous or merely scabrous beneath ; receptacles glabrous or nearly
so, at least in age, sometimes scabrous.
Leaves gradually and evenly acute to long-acuminate F. glabrata.
Leaves rounded or very obtuse at the apex, often abruptly apiculate.
Leaf blades broadly ovate or rounded-oval, broadly rounded at the
apex, less than twice as long as wide F. guajavoides.
Leaf blades short-acute or obtuse and apiculate at the apex, oblong-
elliptic to oval-elliptic, sometimes oblong, much more than twice
as long as wide.
Stipules 1-1.5 cm. long; epidermis of the petioles exfoliating; leaves
usually rough to the touch F. radula.
Stipules 4-6 cm. long; epidermis of the petioles not exfoliating;
leaves smooth to the touch F. crassiuscula.
Receptacles geminate; involucre bilobate; stamen 1. Subgenus Urostigma.
Involucre very asymmetric, adherent to the receptacle over a large portion
of its surface, the receptacle attached excentrically to the peduncle
and its main axis thus parallel to that of the supporting branch.
Receptacles small, 5-10 mm. in diameter; leaves glabrous.
Receptacles pedunculate F. tecolutensis.
Receptacles sessile.
Leaves broadly rounded at the apex F. Jimenezii.
Leaves acute or short-acuminate, sometimes obtuse or very obtuse
but not rounded.
Petioles short, mostly 5-17 mm. long; leaf blades mostly 5-8 cm.
long F. Lundellii.
Petioles elongate, mostly 2-5 cm. long; leaf blades mostly 8-15 cm.
long.
Leaf blades mostly 4-8 cm. wide, the lateral nerves prominent
beneath, 7-9 on each side F. Tuerckheimii.
Leaf blades mostly 2-4.5 cm. wide, the lateral nerves very slender,
scarcely prominent, inconspicuous, 8-12 on each side.
F. eugeniaefolia.
Involucre symmetric, free from the receptacle or nearly so, the receptacle
attached centrally to the peduncle or branch, its main axis thus form-
ing an angle with that of the supporting branch.
Receptacles sessile or, in one species, some sessile and others pedunculate.
Receptacles partly sessile and partly pedunculate upon the same
branch F. Cookii.
Receptacles all sessile.
Leaves cuspidate-acuminate, with a long acute acumen. Leaves
glabrous F. panamensis.
Leaves not cuspidate, sometimes short-acuminate but with an
obtuse acumen.
Leaves conspicuously and often densely pilose or pubescent on
the upper surface, usually rough to the touch.
Receptacles globose F. inamoena.
Receptacles oval, conspicuously longer than broad.
F. Popenoei.
Leaves glabrous on the upper surface or nearly so, not rough.
34 FIELDIANA: BOTANY, VOLUME 24
Receptacles 5-6.5 mm. in diameter; leaves abruptly acute or
short-acuminate, with only 2-4 pairs of lateral nerves.
F. Colubrinae.
Receptacles 6-12 mm. in diameter; leaves broadly rounded to
obtuse at the apex, sometimes short-apiculate, with usually
5-7 or more pairs of nerves.
Leaves conspicuously cordate at the base, mostly 7.5-14 cm.
wide F. cabusana.
Leaves obtuse to subcordate at the base, mostly 4-7 cm. wide.
Involucre small, about 5 mm. in greatest diameter, incon-
spicuous; receptacles usually quite glabrous.
F. costaricana.
Involucre large, conspicuous, enclosing the receptacle for
half to two-thirds its length; receptacles finely pubes-
cent or in age glabrate F. cotinifolia.
Receptacles all pedunculate, the peduncles sometimes short but usually
elongate and conspicuous.
Leaf blades pilose or puberulent beneath, sometimes glabrate in age.
Receptacles 8-9 mm. in diameter, minutely puberulent or glabrate;
leaves 3-5 times as long as broad F. Donnell-Smithii.
Receptacles 13-22 mm. in diameter, usually conspicuously pubes-
cent or pilose; leaves less than 2.5 times as long as broad.
Young branches densely ferruginous- villous; peduncles 2-3 mm.
long, the receptacles pilose F. velutina.
Young branches glabrate; peduncles 4-9 mm. long, the receptacles
finely puberulent F. lapathifolia.
Leaf blades glabrous beneath or practically so.
Receptacles 15-25 mm. in diameter.
Leaves cuneate-obovate, rounded at the apex, long-tapering to
the base F. involuta.
Leaves oblong to ovate, broadest at or below the middle, not
cuneate at the base F. Goldmanii.
Receptacles 4-12 mm. in diameter.
Leaves rounded or very obtuse at the apex F. ovalis.
Leaves acute or acuminate.
Leaf blades oblong-oblanceolate, broadest above the middle;
receptacles only 5-6 mm. in diameter F. Oerstediana.
Leaf blades broadest at or below the middle; receptacles
usually larger.
Leaves mostly 1.5-3 cm. wide; ostiole depressed.
F. padifolia.
Leaves mostly 4.5-8.5 cm. wide; ostiole convex.
Receptacles 8-10 mm. in diameter, long-pedunculate.
F. Hemsleyana.
Receptacles 4-5 mm. in diameter, short-pedunculate.
F. Schippii.
Ficus cabusana Standl. & Steyerm. Field Mus. Bot. 22: 226.
1940. Matapalo.
In quebradas or thickets, 500-1,300 meters; Escuintla; Sacate-
pe"quez (above Barranco Hondo); San Marcos (type from Potrero
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 35
Matasan, along Rio Cabus, Volcan de Tajumulco, Steyermark
37583); sterile specimens from Izabal, near sea level, possibly are
referable here.
A small to large tree, sometimes 30 meters high, glabrous almost throughout;
stipules caducous, 2 cm. long, long-acuminate; leaves large, long-petiolate, the
petioles 2.5-6.5 cm. long; leaf blades oval or broadly oval-ovate, 12-21 cm. long,
7.5-14 cm. wide, very obtuse or rounded at the apex, not apiculate, broad and
rounded at the base and shallowly and narrowly cordate, the lateral nerves about
10 pairs, slender and prominent beneath; receptacles sessile, 12 mm. long, often
crowded and obtusely angulate, glabrous, almost wholly included in the large
thin involucre, rounded at the apex, the ostiole small, prominent; involucre
bilobate, brownish, glabrous or sparsely and minutely puberulent.
Ficus Carica L. Sp. PI. 1059. 1753. Higo; Higuero (the plant).
Native of Asia but cultivated for its fruit in all warmer regions
of the earth, where the climate is not too unfavorable. Planted
sporadically in the mountains of Guatemala and sometimes even
at low elevations, but never, so far as we know, in quantity; Usually
one to a dozen bushes or small trees are found about a dwelling here
and there through the country.
A coarse shrub or a tree 9 meters high or less, usually or often branching from
the ground, scabrous throughout; leaves long-petiolate, palmately 3-5-1 obate,
the lobes obtuse, undulate or often again lobate, cordate at the base; receptacles
solitary, pyriform.
We have noted trees in cultivation in Alta Verapaz, Zacapa,
Santa Rosa, Guatemala, Sacatepe"quez, Chimaltenango, Solola,
Huehuetenango, Totonicapan, Retalhuleu, Quezaltenango, and
San Marcos; doubtless a few are to be found in every department.
The trees often bear well, especially in drier regions or during the
dry months, and the fruit is of reasonably good quality. It some-
times is offered for sale in the markets, where imported dried foreign
figs also are obtainable. The fig was introduced into the North
American continent at an early date, and has thrived in many
regions, particularly southwestern United States and northern
Mexico. In Central America, however, it is little grown, and
scarcely ever on a large scale. The largest plantings we have seen
were in the mountains of Honduras and the Pacific lowlands of
Costa Rica.
Ficus Colubrinae Standl. Contr. U. S. Nat. Herb. 20: 16. 1917.
In forest or pastures, 450 meters or less; Alta Verapaz (type from
Cubilguitz, Tuerckheim 11.156); Izabal. British Honduras; Hon-
duras; Costa Rica; Panama.
36 FIELDIANA: BOTANY, VOLUME 24
A tree 9-15 meters high, the trunk sometimes 45 cm. in diameter, often epiphy-
tic, the young branchlets densely appressed-pilose with long sordid hairs; stipules
5-8 mm. long, appressed-pilose outside, long-acuminate; petioles 8-24 mm. long,
appressed-pilose; leaf blades oval to obovate-oval or oval-oblong, 5-9 cm. long, 2-5
cm. wide, rounded or obtuse at the apex and abruptly contracted into a triangular
acumen, rounded or very obtuse at the base, 5-nerved, glabrous above, appressed-
pilose beneath along the nerves and veins, the lateral nerves 2-4 pairs; receptacles
sessile, subglobose, 5-6.5 mm. in diameter, glabrous, green or yellow, sometimes
streaked with red, the ostiole not prominent; involucre very small, bilobate, the
lobes rounded, hirsute at the base.
Schipp reports the tree as epiphytic in British Honduras upon
Orbignya. The species has been reported from Guatemala as F.
Hartwegii Miq.
Ficus Cookii Standl. Contr. U. S. Nat. Herb. 20: 15. 1917.
Amate.
Along streams or on forested hillsides, 350-2,000 meters; El
Progreso; Huehuetenango. Chiapas; several times collected, the
type from San Vicente.
A small or large tree, commonly 9-15 meters high, the branchlets puberulent
or glabrate; stipules about 1.5 cm. long, glabrous or nearly so, caducous, acumi-
nate; petioles stout, 2.5-7.5 cm. long; leaf blades broadly oval to rounded-oval or
orbicular-ovate, 6-11 cm. long, 4.5-8.5 cm. wide, broadly rounded at the apex,
sometimes apiculate, shallowly cordate or broadly rounded at the base, 5-7-nerved,
coriaceous, the lateral nerves 8-10 pairs; receptacles geminate, subglobose, about
1 cm. in diameter, red or pinkish, glabrous, the ostiole slightly elevated; involucre
two-thirds as long as the receptacle and closely investing it, bilobate, the lobes
broadly rounded, rigid, finely puberulent; receptacles partly sessile and partly
pedunculate, the peduncles equaling or shorter than the receptacles.
Ficus costaricana (Liebm.) Miq. Ann. Mus. Bot. Lugd. Bat.
3: 298. 1867. Urostigma costaricanum Liebm. Dansk. Vid. Selsk.
Skrivt. V. 2: 322. 1851. F. Kellermanii Standl. Contr. U. S. Nat.
Herb. 20: 18. 1917 (type from El Rancho, El Progreso, Kellerman
5595). Amate; Higo; Matapalo; Cuxamate (fide Aguilar).
Dry or moist hillsides, in forest or open places, often along road-
sides, frequently planted as a shade tree, ascending from sea level
to about 2,000 meters, most common below 1,000 meters; Izabal;
Zacapa; Chiquimula; Jalapa; Santa Rosa; Escuintla; Guatemala;
Sacatepe"quez; Retalhuleu; Solola; San Marcos. British Honduras;
Honduras; Salvador; Costa Rica; Panama.
A small to large tree, often epiphytic, the trunk usually low, often a meter
thick, the crown broad and spreading, dense; branchlets glabrous or when young
sometimes sparsely hirsute; stipules 1-2.5 cm. long, often persisting for a long
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 37
time, brown; petioles 1-3.5 cm. long, glabrous or sometimes sparsely hirsute;
leaf blades narrowly obovate-oblong to obovate, oblong, or elliptic-oblong, mostly
6-15 cm. long and 2.5-6.5 cm. wide, rounded or obtuse at the apex or sometimes
subacute, rounded and emarginate or subcordate at the base, usually 5-nerved,
the lateral nerves 5-7 pairs, often coriaceous, sometimes lustrous on the upper
surface; receptacles geminate, sessile, depressed-globose, 8-12 mm. in diameter,
glabrous, green to red or sometimes whitish, the ostiole not prominent; involucre
bilobate, about 1 cm. long, the lobes rounded, somewhat strigose outside.
Called "higuero" and "higuillo" in Honduras. This is one of
the most common Ficus species of the central region and of the
Pacific slope, especially in Santa Rosa and Escuintla. It is probably
the species most seen as a shade tree about houses or planted along
roads and streets, as about Antigua and Amatitlan (whence proba-
bly its name, signifying "place of fig trees"). Like other species,
it can be reproduced quickly from branches set in the ground, which
take root and grow rapidly. Tourists are recommended to see the
avenues of this species about Antigua, which probably are the finest
in all Guatemala. The trees lose their leaves toward the end of the
dry season but do not remain naked for long. The Guatemalan
material referred here is variable, and it is possible that more ample
collections will make possible its division into two or more species.
It may be that ultimately F. Kellermanii may be maintained as a
distinct species, but at present it is not obvious on what characters
it can be separated.
Ficus cotinifolia HBK. Nov. Gen. & Sp. 2: 49. 1817.
Chiquimula (Quebrada Shusho, above Chiquimula, 480 meters,
in arenal}. Mexico; Costa Rica.
Often a large tree with broad spreading crown and low trunk, the young
branchlets tomentulose or glabrate; stipules 5-13 mm. long, sericeous; petioles
1-7 cm. long; leaf blades broadly oblong to suborbicular, usually broadest slightly
above the middle, 5-14 cm. long, 2.5-10 cm. wide, usually broadly rounded at the
apex, sometimes only obtuse, rounded or subcordate at the base, commonly grayish
green when dried, glabrous or tomentulose above, tomentulose or short-villous
beneath or in age glabrate, with 5-7 pairs of lateral nerves; receptacles globose or
slightly depressed, 6-11 mm. in diameter, pale green, often spotted with red or
dark green, finely sericeous or in age glabrate, the ostiole not prominent; involucre
bilobate, half as long as the receptacle or more, densely white-sericeous on both
surfaces.
Known in Yucatan by the names "alamo" and "copo"; called
"congo" in Oaxaca. The roots, as in other species, are often exposed
above the ground. The bark is dark brown or grayish; wood white
throughout. The leaves and branches are much used in the Yucatan
38 FIELDIANA: BOTANY, VOLUME 24
Peninsula as fodder for horses and mules, and the sap is reported to
be used as an adulterant of chicle.
Ficus crassiuscula Warb. ex Standl. Contr. U. S. Nat. Herb.
20: 12. 1917. Amate.
Wet or moist forest, sometimes in cafetales, occurring at sea
level or as high as 1,100 meters; Pete"n; Izabal; Alta Verapaz;
Escuintla; Chimaltenango; Quezaltenango; San Marcos. Hon-
duras; Costa Rica; Panama.
A large tree, sometimes 30 meters high with a trunk 2 meters in diameter,
often buttressed, the crown rounded or spreading, the bark brown, slightly rough,
the branchlets glabrous or nearly so; stipules 4-6 cm. long, caducous, glabrous or
nearly so; petioles 2.5-4 cm. long; leaf blades pale green when dried, thick, gla-
brous, broadly oblong to narrowly oval or oblong-obovate, 10-23 cm. long, 5-11
cm. wide, obtuse at the apex and abruptly contracted into a short acumen, obtuse
or rounded at the base, the lateral nerves 14-22 on each side; peduncles solitary,
2 cm. long, the involucre small and inconspicuous; receptacle obovoid-globose,
2 cm. in diameter or larger, green, soft and succulent, sometimes pink at maturity.
Ficus Donnell-Smithii Standl. Contr. U. S. Nat. Herb. 20:
21. 1917.
Alta Verapaz, 350 meters (type from Cubilgiiitz, Tuerckheim
11.597; J. D. Smith 8289). British Honduras.
A small tree of 4-5 meters, or sometimes 15 meters tall, with a trunk 20 cm.
in diameter, the branchlets puberulent or short-hirtellous, tardily glabrate;
stipules 5-7 mm. long, puberulent; petioles 7-18 mm. long; leaf blades oblong or
narrowly oblong, sometimes oblanceolate-oblong, 7-15 cm. long, 2-2.7 cm. wide,
acuminate to rounded at the apex, rounded at the base, scaberulous or glabrate
above, short-pilose or glabrate beneath, the lateral nerves 7-8 pairs; peduncles
geminate, 6-7 mm. long, the involucre 3-4 mm. broad; receptacles subglobose,
8-9 mm. in diameter, the ostiole not prominent.
The species has been reported from Guatemala as F. lancifolia
Hook. & Arn.
Ficus elastica Roxb. Hort. Beng. 65. 1814, nomen nudum; Fl.
Ind. ed. 2. 3: 541. 1832.
Native of India. Planted occasionally in parks and gardens of
Guatemala City and elsewhere for ornament.
Becoming a large tree, glabrous; leaves very thick, oblong to elliptic, 10-30
cm. long, cuspidate-acuminate, obtuse at the base, the lateral nerves very numer-
ous; receptacles axillary, sessile, geminate, oval or oblong, about 12 mm. long,
greenish yellow.
This species is little planted in Guatemala but in some other
regions of Central America it is more plentiful. It is the India
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 39
rubber plant of the United States, where it is much grown in pots
as a house plant, since it withstands neglect and especially the dry
air of steam-heated dwellings. As a shade tree it is not to be recom-
mended since the large heavy limbs are easily broken by wind.
Var. variegata L. H. Bailey is rarely planted in Guatemala City.
Its leaves have creamy white or yellow margins. The stipules in
this species are extraordinarily large and enclose the young leaves
like a sheath, which is rose-colored or purplish.
Ficus eugeniaefolia (Liebm.) Hemsl. Biol. Centr. Amer. Bot.
3: 144. 1883. Urostigma eugeniaefolium Liebm. Dansk. Vid. Selsk.
Skrivt. V. 2: 329. 1851. Amate.
Moist or wet forest or open fields, Alta Verapaz (vicinity of
Coban, 1,250-1,400 meters). Salvador; Costa Rica.
A small or large tree, glabrous throughout or nearly so; stipules 1-2.5 cm.
long, long-acuminate, puberulent outside or glabrate; petioles 1.5-3 cm. long;
leaf blades obovate or elliptic-obovate, mostly 5-12 cm. long and 3-6.5 cm. wide,
obtuse or acute and apiculate, obtuse at the base or on sterile branches sometimes
shallowly cordate, glabrous, the lateral nerves 8-12 pairs, very slender and not
prominent beneath; involucre asymmetric, large, thin, at first completely enclosing
the receptacle, at maturity about two-thirds as long; receptacles globose or some-
what depressed, 1 cm. in diameter, the ostiole large, slightly elevated.
Ficus glabrata HBK. Nov. Gen. & Sp. 2: 47. 1817. F. anthel-
mintica Mart. Syst. Mat. Med. Bras. 88. 1843, not F. anthelmintica
Raeuschel, 1797. Pharmacosycea angustifolia Liebm. Dansk. Vid.
Selsk. Skrivt. V. 3: 333. 1851. F. segoviae Miq. Ann. Mus. Bot.
Lugd. Bat. 3: 300. 1867. Amate; Higueron; Matapalo.
Forest or open fields or hillsides, often along roadsides, frequently
growing about habitations, ascending from sea level to about
1,400 meters, but chiefly at low elevations; Pete"n; Alta Verapaz;
Baja Verapaz; Izabal; Zacapa; El Rancho; Jutiapa; Santa Rosa;
Escuintla; Guatemala; Sacatepe"quez; Suchitepe'quez; Retalhuleu;
Quezaltenango; San Marcos. Southern Mexico; British Honduras
to Salvador and Panama; Colombia to Brazil and Peru.
Usually a large tree, commonly 12-40 meters high, with pale, almost smooth
bark and often with low buttresses, the trunk usually low and thick and the crown
spreading, the branchlets glabrous; stipules caducous, pale green, long and narrow,
sometimes 6 cm. long, glabrous; leaves slender-petiolate, mostly elliptic-oblong to
elliptic-oval, 12-23 cm. long, 5-10 cm. wide, often lance-elliptic or narrowly
oblong-lanceolate, acute or acuminate to long-attenuate, obtuse or acute at the
base, glabrous, green or pale green when dried, the lateral nerves conspicuous,
14-21 pairs; peduncles solitary, thick, 7-15 mm. long, the involucre very small;
receptacles subglobose, 1.5-4 cm. in diameter or even larger, glabrous or obscurely
40 FIELDIANA: BOTANY, VOLUME 24
scaberulous, usually mottled with light and dark green, very soft and juicy at
maturity.
Called "higuero" in Honduras, and in Salvador sometimes
"chilamate" and "chilamaton." This is one of the common large
trees of the Pacific plains and the lower Motagua Valley, where
there are some huge examples that almost rival the ceibas in size.
The fruit is larger than that of most other Central American species
and more like that of the cultivated fig. It is of mediocre flavor and
is little eaten by man although much sought by many birds and
mammals. In its native regions it has long been known that the
copious white latex that issues from the trunk or branches when
cut has anthelmintic properties, and in recent years the latex has
attracted the attention of local and foreign physicians. It is said
that some fresh latex is now being exported to the United States
for hospital use, and in some of the hospitals of Panama and the
Atlantic coast it is the practice to give a dose of it to all or most
patients entering for hospitalization, on the well-grounded assump-
tion that they need a vermifuge. Ficus segoviae has often been
maintained as a distinct species, but it now appears certain that the
specimens referred to it are merely juvenile shoots or seedlings,
which usually have narrow and greatly elongate leaves, much nar-
rower than those of normal adult branches. In Alta Verapaz the
young hard fruits are used for making designs on hats and probably
other articles. If a cross section of the fruit is pressed against the
surface, a blackish circular figure of more or less permanence is left
upon it.
Ficus glaucescens (Liebm.) Miq. Ann. Mus. Bot. Lugd. Bat.
3: 300. 1867. Pharmacosycea glaucescens Liebm. Dansk. Vid. Selsk.
Skrivt. V. 2: 332. 1851. Amate.
Forest or thickets, often along streams, ascending from sea level
to about 1,600 meters, but chiefly at 900 meters or less; Alta Verapaz;
Baja Verapaz; Izabal; Zacapa; Santa Rosa; Escuintla; Guatemala;
Suchitepe"quez; Retalhuleu; Quezaltenango; San Marcos. Southern
Mexico; Salvador; Nicaragua; Panama.
A medium-sized or often very large tree with pale, almost smooth bark and
usually a low spreading crown, the trunk usually low and thick, the branchlets at
first hispidulous or puberulent; stipules 1-2 cm. long, scabrous or glabrate; petioles
1-4 cm. long, with exfoliating epidermis; leaf blades oval-oblong to obovate-oval,
8-23 cm. long, 4-11 cm. wide, rather thick, usually pale grayish green when dried,
rounded or very obtuse at the apex and usually abruptly short-pointed, rounded
or obtuse at the base, scaberulous on the upper surface and rough to the touch,
beneath usually densely hirtellous, or sometimes glabrate except on the nerves,
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 41
the lateral nerves 7-12 pairs, stout, conspicuous; peduncles solitary, 5-20 mm.
long, the involucre very small; receptacles subglobose, 1.5-2.5 cm. in diameter,
usually mottled with light and dark green, commonly densely pilose or hirtellous.
This species is noteworthy for its very rough leaves, suggestive
of sandpaper, a feature characteristic also of F. radula. These two
species are not always sharply separable.
Ficus Goldmanii Standl. Contr. U. S. Nat. Herb. 20: 32.
1917. Amate.
Baja Verapaz(?); Jutiapa (in finca near Jutiapa). Western and
southern Mexico; British Honduras; Salvador.
Usually a medium-sized tree with low trunk and spreading crown, the branch-
lets glabrous or nearly so; stipules short, sericeous or puberulent outside; petioles
2-3.5 cm. long; leaf blades oblong to elliptic-oval, 7-18 cm. long, 4-10 cm. wide,
rounded or very obtuse at the apex, obtuse at the base or narrowly rounded,
glabrous, usually coriaceous, the lateral nerves 5-13 pairs; receptacles short-
pedunculate, globose, 1.5-2.5 cm. in diameter, puberulent or glabrate, the ostiole
not prominent.
Called "matapalo" in British Honduras. The species is common
in Salvador and should occur more plentifully in the Oriente of
Guatemala, where probably we have overlooked it.
Ficus guajavoides Lundell, Bull. Torrey Club 64: 547. 1937.
Type collected near Valentin, El Cayo District, British Hon-
duras, Lundell 6295, in high, advanced forest; to be expected in
Pete"n.
A tree 45 meters tall, the low trunk 75 cm. in diameter, with thin buttresses,
glabrous throughout; stipules 3-4.5 cm. long, attenuate, caducous; petioles stout,
2.5-6 cm. long, with exfoliating epidermis; leaf blades broadly oval or rounded-
oval, 10-20 cm. long, 8-14.5 cm. wide, broadly rounded at the apex, rounded or
very obtuse at the base, thick, paler beneath, smooth to the touch, the lateral
nerves 11-17 pairs, divergent at right angles; peduncles solitary, 2-3.5 cm. long;
receptacles globose or obovoid-globose, 2-3 cm. in diameter.
Further material is necessary to determine whether this is a
species with constant characters or only a leaf form of F. Tonduzii
Standl.
Ficus Hemsleyana Standl. Contr. U. S. Nat. Herb. 20: 29.
1917. Urostigma verrucosum Liebm. Dansk. Vid. Selsk. Skrivt.
V. 2: 321. 1851. F. verrucosa Hemsl. Biol. Centr. Amer. Bot. 3:
148. 1883, not F. verrucosa Miq. 1867. Amate; Matapalo.
Wet or dry forest or thickets, often by roadsides, ascending to
1,200 meters, but mostly in the lowlands; Izabal; Escuintla; Suchi-
42 FIELDIANA: BOTANY, VOLUME 24
tep^quez; Retalhuleu; Quezaltenango; Quiche*. British Honduras;
Honduras; Salvador; Nicaragua; Costa Rica; Panama.
A medium-sized or large tree, often epiphytic, the branchlets glabrous or
obscurely puberulent; stipules 1-2 cm. long, glabrous or puberulent; petioles 2-6
cm. long, slender; leaf blades oblong or elliptic-oblong, 10-22 cm. long, 4-8 cm.
wide, abruptly acuminate or caudate-acuminate, rounded or subcordate at the
base, rather thin, glabrous, slightly paler beneath, with 7-13 pairs of lateral nerves;
peduncles slender, mostly shorter than the receptacles, the involucre 4 mm. broad;
receptacles globose, green, about 1 cm. in diameter, minutely puberulent or
glabrate, the ostiole slightly prominent.
This has been reported from British Honduras as F. laevigata
Vahl.
Ficus inamoena Standl. Contr. U. S. Nat. Herb. 20: 16. 1917.
Amate; Cushamate (Jutiapa); Cuxche (fide Aguilar).
Moist or rather dry forest, often in pine forest, frequently in
brushy quebradas or along streams, sometimes in fields or by road-
sides, ascending from sea level to about 1,600 meters; El Progreso;
Jalapa; Jutiapa; Santa Rosa; Escuintla; Guatemala; Sacatepe 1 -
quez; Chimaltenango; Quiche" (type from Joyabaj, 0. F. Cook 22).
Honduras.
A small to large tree, often 12-15 meters high or more, with low thick trunk
and dense spreading crown; branchlets mostly whitish-pilose or hirtellous; stipules
commonly 5-12 mm. long, strigose dorsally; petioles 1.5 cm. long or less, stout;
leaf blades rounded-oval to oblong or obovate-oblong, 6-14 cm. long, 4.5-6.5 cm.
wide, broadly rounded or obtuse at the apex, usually conspicuously cordate at
the base, with a shallow narrow sinus, rather softly pilose on both surfaces, some-
times glabrate above, the lateral nerves prominent, 5-8 pairs; involucre bilobate,
strigose; receptacles sessile, depressed-globose, 1 cm. in diameter, glabrous or
nearly so, green, the ostiole not elevated.
Called "higuero" in Honduras.
Ficus involuta (Liebm.) Miq. Ann. Mus. Bot. Lugd. Bat. 3:
298. 1867. F. obtusifolia HBK. Nov. Gen. & Sp. 2: 49. 1817, not
F. obtusifolia Roxb. 1814. Urostigma involutum Liebm. Dansk.
Vid. Selsk. Skrivt. V. 2: 320. 1851. U. Bonplandianum Liebm. op.
cit. 323. 1851. F. Bonplandiana Miq. loc. cit. Amate; Matapalo;
Copo zotz (Pete*n); Cux (fide Aguilar).
Open forest, wet or rather dry regions, often in fields, frequently
by roadsides, ascending from sea level to 1,500 meters; Pete'n; Izabal;
Zacapa; Jutiapa; Santa Rosa; Escuintla; Guatemala; Sacatepe'quez ;
Retalhuleu; Huehuetenango. Southern Mexico; British Honduras
to Panama.
STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 43
A medium-sized or large tree, usually with short thick trunk and broad
spreading crown, the thick branchlets sparsely puberulent; stipules 1.5-3 cm. long,
glabrous; leaves on petioles 1-2 cm. long, oblanceolate-oblong or cuneate-oblong,
sometimes cuneate-obovate, rounded or very obtuse at the apex, gradually long-
cuneate to the base, glabrous, thick, the lateral nerves 6-8 pairs; peduncles
geminate, 2-3 mm. long, the involucre large, often covering almost half the
receptacle; receptacles globose, often appearing sessile, 1.5-2 cm. in diameter,
finely sericeous, the ostiole prominent.
Sometimes called "capulamate" in Salvador. Well distinguished
by the narrowly wedge-shaped leaves, unlike those of any other
Central American species. The tree is abundant in many regions,
especially along the hills of the lower Pacific slope.
Ficus Jimenezii Standl. Contr. U. S. Nat. Herb. 20: 14. 1917.
Forest or open hillsides, about 300 meters; Escuintla. Salvador;
Costa Rica.
A large tree, sometimes epiphytic, the branchlets glabrous; stipules 1-1.5 cm.
long, puberulent, caducous; petioles 2-3 cm. long; leaf blades obovate-oval, oval,
or obovate-oblong, mostly 5-11 cm. long and 3.5-6 cm. wide, rounded at the apex,
rounded or obtuse at the base, thick, usually blackening when dried, glabrous, with
6-9 pairs of lateral nerves, these slender and inconspicuous; involucre asymmetric,
large and conspicuous; receptacles sessile, geminate, depressed-globose, 5-8 mm.
in diameter, glabrous or minutely puberulent, green spotted with red or brown.
The species was named for Oton Jime'nez Luthmer of Costa
Rica, enthusiastic student of the rich flora of Costa Rica, and
esteemed friend of all botanists visiting that country. The latex
of this species is said to be employed in Salvador as a medicament
for expelling intestinal parasites.
Ficus lapathifolia (Liebm.) Miq. Ann. Mus. Bot. Lugd. Bat.
3: 297. 1867. Urostigma lapathifolium Liebm. Dansk. Vid. Selsk.
Skrivt. V. 2: 319. 1851. Urostigma guatemalanum Miq. Versl. Med.
Kon. Akad. Amsterdam 13: 411. 1862 (described from plants grown
at Berlin from seed said to have been collected in Guatemala by
Warscewicz). F. guatemalana Miq. Ann. Mus. Bot. Lugd. Bat. 3:
298. 1867. Amate; Amate cusho (Oriente).
Moist thickets or forest, often on open hillsides or along streams,
ascending from sea level to about 1,200 meters; Pete"n; Zacapa;
Jalapa; Jutiapa; Guatemala; Escuintla. Southern Mexico; British
Honduras.
A medium-sized or large tree, often epiphytic, the branchlets puberulent and
hirtellous; stipules 1.5-2 cm. long, sericeous; petioles 1-3 cm. long; leaf blades
oval to broadly oblong, mostly 10-25 cm. long and 5-15 cm. wide, rounded or
44 FIELDIANA: BOTANY, VOLUME 24
obtuse at the apex and often short-apiculate, rounded to subcordate at the base,
pubescent or glabrate above, densely velutinous-pilose beneath or in age glabrate,
the lateral nerves prominent, 7-13 pairs; peduncles short, geminate, the involucre
about 8 mm. broad, bilobate, sericeous; receptacles globose, 1.5-2 cm. in diameter,
minutely sericeous, green, the ostiole not prominent.
Called "alamo" and "higo" in Campeche, the fruit said to be
eaten there.
Ficus Lundellii Standl. Carnegie Inst. Wash. Publ. 461: 54.
1935. Amate.
Pete"n, the type from La Libertad, Lundell 3406; known only
from the region of the type locality.
Branchlets puberulent or almost glabrous; stipules caducous, 1.5-2.5 cm.
long, minutely puberulent; petioles 5-17 mm. long; leaf blades elliptic-oblong,
broadest near the middle, 4.5-9 cm. long, 2-4.5 cm. wide, very obtuse or rounded
at the apex, obtuse at the base, glabrous, the lateral nerves about 8 pairs; recep-
tacles sessile, geminate, 7-8 mm. in diameter, minutely puberulent or almost
glabrous, green spotted with dark purple or red, the ostiole prominent; involucre
asymmetric, glabrous, deeply bilobate, shorter th
