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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
tpart 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; Sacatepe1-
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 than the receptacle.

Ficus Oerstediana Miq. Ann. Mus. Bot. Lugd. Bat. 3: 299.
1867. Urostigma Oerstedianum Miq. in Seem. Bot. Voy. Herald
196. pi. 36. 1854. Matapalo.

Moist or wet forest or thickets, sometimes in Manicaria swamps,
at or little above sea level (360 meters or less); Pete"n; Alta Verapaz;
Izabal. Chiapas; British Honduras; Honduras; Costa Rica; Panama.

A small or medium-sized tree, mostly 15 meters high or less, the trunk some-
times 60 cm. in diameter, often epiphytic, the branchlets puberulent or glabrate;
stipules 5-15 mm. long; leaf blades coriaceous, mostly obovate to oblanceolate-
oblong, 4-11 cm. long, 1-4.5 cm. wide, acute or obtuse, acute or cuneate-attenuate
at the base, sometimes obtuse, glabrous, the lateral nerves 9-15 pairs; peduncles
geminate, 3-7 mm. long, the involucre small and inconspicuous; receptacles
globose, glabrous, green or reddish, 5-6 mm. in diameter, the ostiole plane or
slightly elevated.

Called "higuillo" in Honduras. This has the smallest fruits of
all Central American species of Ficus.

Ficus ovalis (Liebm.) Miq. Ann. Mus. Bot. Lugd. Bat. 3: 298.
1867. Urostigma ovale Liebm. Dansk. Vid. Selsk. Skrivt. V. 2:
324. 1851.

Guatemala (near Fiscal, 1,100 meters, dry rocky hillsides);
probably in Pete*n. Southern Mexico; British Honduras; Salvador;
Nicaragua; Costa Rica (type from Guanacaste).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 45

A large tree with spreading crown, the trunk low, often fluted, the bark yellow-
ish or brownish, the branchlets glabrous; stipules 1-1.5 cm. long, glabrous; petioles
1-3.5 cm. long; leaf blades oval to oblong-obovate, 7-11 cm. long, 4-6 cm. wide, or
sometimes larger, rounded at the apex, rounded and emarginate at the base,
glabrous, the lateral nerves mostly 4-6 pairs, sometimes more numerous; peduncles
geminate, 3-6 mm. long, the involucre 5 mm. broad; receptacles globose, glabrous,
green or red, 6-8 mm. in diameter or slightly larger.

Called "matapalo" in British Honduras.

Ficus padifolia HBK. Nov. Gen. & Sp. 2: 47. 1817. F. lanci-
folia Hook. & Arn. Bot. Beechey Voy. 310. 1839. Urostigma
sapidum Liebm. Dansk. Vid. Selsk. Skrivt. V. 2: 327. 1851. F.
sapida Miq. Ann. Mus. Bot. Lugd. Bat. 3: 298. 1867. Amate;
Cush; Matapalo; Cushamate; Higo; Capulamate; Amatillo; Gus
(fide Aguilar); Moco; Capulin (Huehuetenango).

Moist or rather dry forest or thickets, often in second growth,
frequent along streams and in hedgerows, often growing about
dwellings, ascending from sea level to 1,700 meters (in Huehue-
tenango), most common at 900 meters or less; Alta Verapaz; Izabal;
Zacapa; Baja Verapaz; El Progreso; Chiquimula; Jutiapa; Santa
Rosa; Escuintla; Guatemala; Suchitepe"quez; Retalhuleu; Que-
zaltenango; San Marcos; Huehuetenango. Mexico; British Hon-
duras to Panama; Colombia.

A large or small tree, often epiphytic, the bark whitish or pale yellowish, the
branchlets glabrous or minutely puberulent; stipules 5-15 mm. long, glabrous or
minutely puberulent; petioles 0.5-3 cm. long, slender; leaf blades mostly narrowly
lance-oblong or elliptic-lanceolate, 4-12 cm. long, 1.5-4.5 cm. wide, acute to long-
attenuate, rounded or obtuse at the base, glabrous, usually green when dried, the
lateral nerves 5-12 pairs; peduncles geminate, mostly shorter than the receptacle,
the involucre 3-4 mm. broad; receptacles subglobose, 9-12 mm. in diameter, green,
usually spotted with red or purple, glabrous or minutely puberulent, the ostiole
rather large, depressed.

Called "higuillo" in Honduras and "chilamate" in Salvador.
The species has been reported from Guatemala as F. ligustrina
Kunth & Bouche". This is perhaps the commonest Ficus species
of all Central America, abundant in many regions. Unlike other
local species, this often has abundant aerial roots dangling from the
high branches and in Mexico, at least, it often becomes a tree of the
banyan type. It is frequently planted in Guatemala for living fence-
posts. The leaves are said to furnish excellent forage for cattle
along the Pacific lowlands.

Ficus panamensis Standl. Contr. U. S. Nat. Herb. 20: 15. 1917.
Wet forest, sometimes in Manicaria swamps, at sea level; Izabal.
Tabasco; British Honduras; Costa Rica; Panama; Colombia.

46 FIELDIANA: BOTANY, VOLUME 24

A large or medium-sized tree, often epiphytic, the branchlets puberulent or
glabrous; stipules 2 cm. long, caducous, puberulent; petioles 1-3.5 cm. long; leaf
blades oblong or narrowly obovate-oblong, 9-17 cm. long, 4-6 cm. wide, abruptly
short-acuminate or cuspidate-acuminate, obtuse to emarginate at the base, gla-
brous, the lateral nerves about 16 pairs, slender; receptacles geminate, sessile,
subglobose, 1 cm. in diameter, green, glabrous, the ostiole prominent.

Called "amatillo" in Tabasco.

Ficus pandurata Hort.

Planted for ornament or as a shade tree in Guatemala City and
along the Pacific slope, probably also elsewhere. Native country
unknown, but long established in cultivation in various parts of the
tropics, and often grown in greenhouses of the United States and
Europe.

Becoming a large tree, glabrous or nearly so; leaves sessile or subsessile, broadly
obovate, often panduriform, broadly rounded at the apex, deeply and narrowly
cordate at the base, often 30 cm. long or more, coriaceous, dark lustrous green,
the very prominent, coarse nerves whitish.

The tree is planted rather frequently in parks and gardens of
Central America. The large thick leaves, of unusual form, are very
handsome.

Ficus Popenoei Standl. Field Mus. Bot. 4: 301. 1929.

Wet forest, sometimes in Manicaria swamps, at or little above
sea level; Pete"n(?) ; Izabal. British Honduras; Honduras (type from
Lancetilla Valley near Tela, Atlantic coast).

A small or medium-sized tree, the trunk seldom more than 15 cm. in diameter,
often epiphytic, the thick branchlets densely hirsute with brownish or ferruginous
hairs; stipules 2 cm. long or less, deciduous, appressed-hirsute; petioles 1-2.5 cm.
long; leaf blades rather thin, oval or oval-obovate, mostly 8-20 cm. long and 4-9.5
cm. wide, broadly rounded to obtuse at the apex, somewhat narrowed toward the
cordate or broadly rounded base, densely hispidulous or glabrate above, usually
rough to the touch, paler beneath, densely velutinous-pilose with short yellowish
hairs, the lateral nerves about 12 pairs; peduncles geminate, about 4 mm. long, the
involucre bilobate, appressed-pilose outside; receptacles oblong-obovoid, 1.5-2
cm. long, 1 cm. broad, fulvous-hirsute, the ostiole minute, slightly elevated.

Ficus pumila L. Sp. PI. 1060. 1753. Una de gato.

Native of Japan and China, but grown commonly for ornament
in many tropical and warm regions. Planted frequently for orna-
ment in central Guatemala, usually running over plaster or brick
walls or tree trunks.

Stems often greatly elongate, woody, creeping closely against walls, coarse,
pilose; leaves 2-ranked, on very short petioles, oblong or ovate, commonly 3-7

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 47

cm. long, very obtuse, rounded to shallowly cordate at the base, hirtellous or gla-
brate beneath, the veins very prominent and closely reticulate; receptacles solitary,
pedunculate, pear-shaped, 5-7 cm. long, dark blue or red-purple.

Called "hiedra" in Costa Rica. The plant grows rapidly with
little care, and spreads widely on walls and similar places.

Ficus radula Willd. Sp. PL 4: 1144. 1806. Amate; Chimdn
(Pet&i).

Moist or wet forest or fields, often in pastures, by roadsides, or
along stream beds, 750 meters or less; Pete"n; Izabal; Zacapa;
Escuintla; Retalhuleu; Huehuetenango. Southern Mexico; British
Honduras to Panama; Colombia and Venezuela.

Often a large tree with low trunk and widely spreading crown, the branchlets
puberulent or glabrate; stipules 1-1.5 cm. long; petioles 1-3 cm. long, with exfoliat-
ing epidermis, becoming ferruginous; leaf blades oblong to obovate or oval, 8-16
cm. long, 3.5-8 cm. wide, rounded or obtuse at the apex and abruptly apiculate,
rounded or obtuse at the base, thick, usually scabrous or scaberulous and rough
to the touch, the lateral nerves 7-12 pairs, coarse and prominent beneath; pedun-
cles solitary, about 5 mm. long, the involucre very small; receptacles subglobose,
1.5-3 cm. in diameter, green, scabrous, becoming soft and pulpy at maturity.

Called "higo" and "higuero" in Honduras, and sometimes
"salamate" in Salvador. The specific name signifies "scraper," in
allusion to the rough leaves. Leaves of vigorous seedlings are
sometimes as much as 30 cm. long and 19 cm. wide.

Ficus Schippii Standl. Field Mus. Bot. 8: 7. 1930.

Known only from the type, Middlesex, British Honduras, 60
meters, Schipp 334.

An epiphytic tree of 15 meters, the trunk 10-12 cm. in diameter, the branchlets
glabrous; stipules 15-18 mm. long, long-attenuate, caducous, glabrous; petioles
slender, 1.5-5.5. cm. long; leaf blades oblong or oval-oblong, 8-14 cm. long, 4.5-5.5
cm. wide, obtuse or rounded at the apex and abruptly caudate or acuminate,
glabrous, the lateral nerves about 13 pairs; receptacles on very short peduncles or
almost sessile, subglobose, 5 mm. in diameter, glabrous, the ostiole plane; involucre
appressed, bilobate, the lobes 1 mm. long, rounded.

In the original description this was compared with F. Colubrinae,
with which it certainly has little relationship. It is questionable
whether the type is more than a specimen of F. Hemsleyana with
very young fruit.

Ficus tecolutensis (Liebm.) Miq. Ann. Mus. Bot. Lugd. Bat.
3: 299. 1867. Urostigma tecolutense Liebm. Dansk. Vid. Selsk.
Skrivt. V. 2: 324. 1851.

48 FIELDIANA: BOTANY, VOLUME 24

i

Sacatepe*quez (Alotenango, 1,500 meters). Southern Mexico;
British Honduras.

A tree, sometimes 35 meters high with a trunk a meter in diameter, the branch-
lets glabrous; stipules 1-1.5 cm. long, glabrous; petioles slender, 1-2 cm. long;
leaf blades oblong to elliptic-oblong or obovate-oblong, 6-10 cm. long, 2-4 cm.
wide, rounded to subacute at the apex, obtuse at the base and sometimes emargi-
nate, glabrous, the lateral nerves 7-9 pairs; peduncles geminate, short, the invo-
lucre asymmetric, 5-6 mm. long; receptacles subglobose, 5-8 mm. in diameter,
glabrous.

Apparently this species is one of the rarest of all the Mexican
and Central American ones and only a few collections of it are known.

Ficus Tuerckheimii Standl. Contr. U. S. Nat. Herb. 20: 13.
1917. Amate.

Moist or wet forest or thickets, often on open dry rocky hill-
sides, ascending from sea level to about 1,500 meters; Pete"n; Alta
Verapaz; Baja Verapaz; Izabal; Huehuetenango. Tabasco; British
Honduras; Costa Rica (type from Volcan de Irazu).

A small to large tree, glabrous almost throughout; stipules 1.5-4.5 cm. long,
glabrous or minutely puberulent; petioles 1.5-5.5 cm. long; leaf blades oval or
oblong-oval, 9-17 cm. long, 4-9.5 cm. wide, usually rounded and short-apiculate
at the apex, obtuse or rounded at the base, coriaceous, the lateral nerves 7-9
pairs, often conspicuous; involucre at first enclosing the receptacle, at maturity
about two-thirds its length, very asymmetric; receptacles depressed-globose,
8-10 mm. in diameter, glabrous or sparsely puberulent.

Ficus velutina Willd. Sp. PI. 4: 1141. 1806. Amate; Matapalo.

Roadsides or open fields, sometimes on dry rocky hillsides,
1,250-1,800 meters; Alta Verapaz; Baja Verapaz; Zacapa; Huehue-
tenango. British Honduras to Panama; Colombia and Venezuela.

A small or often large tree, the branchlets thick, brown-pilose; stipules 1.5-2
cm. long, ferruginous-sericeous; petioles 1.5-3 cm. long, thick; leaf blades oval,
ovate-oval, or obovate, 9-25 cm. long, 5.5-11.5 cm. wide, rounded or obtuse at
the apex and usually apiculate, rounded or subcordate at the base, coriaceous,
scaberulous or puberulent above, beneath tomentose or short-pilose or finally
glabrate, the lateral nerves 7-12 pairs; peduncles geminate, usually only 2-3 mm.
long, the involucre 1 cm. long or less; receptacles 1.5-2 cm. in diameter or even
larger, globose, sericeous or glabrate.

MORUS L. Mulberry

Trees or shrubs with milky sap; leaves alternate, dentate, entire, or 3-lobate,
3-nerved at the base; stipules lateral, small, caducous; flowers monoecious or
dioecious, those of each sex in ament-like spikes, these axillary, solitary, short-
pedunculate, the staminate spikes elongate, the pistillate long or short; staminate

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 49

perianth 4-parted, the segments ovate, imbricate; stamens 4, the filaments inflexed
in bud, in an thesis porrect and exserted; segments of the pistillate flower 4, ovate,
decussate-imbricate, succulent in fruit; ovary included, ovoid or subglobose, the
style central, 2-parted almost to the base, the branches linear, equal; ovule pendu-
lous from the apex of the cell; fruit included in the accrescent juicy perianth, the
exocarp more or less succulent, thin or very thick, the endocarp crustaceous; seed
subglobose, with membranaceous testa; endosperm fleshy, abundant, the embryo
incurved, the cotyledons oblong, equal.

About a dozen species, in temperate and tropical regions of both
hemispheres. Several are cultivated for their edible fruits. Only
the following ones are native in Central America, but one other
occurs in Mexico, and it and another are native in the eastern half
of the United States.

Cultivated tree; leaves glabrate, often cordate at the base M. alba,

Native trees; leaves usually or often copiously pubescent beneath, not cordate at

the base.
Flower spikes mostly 6-10 cm. long or longer; leaves 6-10 cm. wide or larger,

with about 8 pairs of lateral nerves M. insignis.

Flower spikes 1-2 cm. long; leaves mostly 3-5 cm. wide, with usually 5 or fewer
pairs of nerves above the basal ones M. celtidifolia.

Morus alba L. Sp. PI. 986. 1753. White mulberry.

Native of China. Naturalized or cultivated in many parts of
the earth. Represented in cultivation in Central America by the
following variety:

Morus alba var. multicaulis (Perrotet) Loudon, Arboret.
Brit. 3: 1348. /. 1223. 1838. M. multicaulis Perrotet, Ann. Soc. Linn.
Bot. Paris 3: 129. 1825.

Planted in many parts of Guatemala, although rather sparingly,
most of the trees small but large ones seen occasionally; noted in
Guatemala, Alta Verapaz, Huehuetenango, Quiche", and Quezalte-
nango, and doubtless planted elsewhere. Native of Asia, but grown
in many parts of the earth.

A large shrub or small, densely branched tree with rough, pale gray bark;
leaves ovate or broadly ovate, mostly large and 8-15 cm. long, almost glabrous
but sometimes slightly rough, frequently cordate at the base, long-petiolate, the
teeth mostly large and rounded; flower spikes about 2.5 cm. long; fruit white or
pinkish at maturity.

This tree is in cultivation in many parts of Central America, to
which it has been introduced largely by perhaps not too scrupulous
foreign promoters of an evasive silk industry. The promoters,
whose primary interest was sale of trees, perhaps have been success-

50 FIELDIANA: BOTANY, VOLUME 24

ful, but so far the silk industry has not been developed and is not
likely to be. It is said that silk was produced in Guatemala and other
Central American countries during early colonial days but to what
extent we do not know. It is claimed that mulberry trees thrive in
Guatemala when properly tended, but most of the scattered ones
we have seen did not appear thrifty. The black mulberry, Morus
nigra L., of eastern Asia, may be planted in some parts of Guatemala
for its fruit, but if so, it is rare. Var. multicaulis is said to be the
mulberry grown in China and Japan as food for silkworms.

Morus celtidifolia HBK. Nov. Gen. & Sp. 2: 33. 1817. M.
mexicana Benth. PI. Hartweg. 71. 1840. Mora.

Rocky stream banks or in moist forest, 250-2,500 meters;
Zacapa; Guatemala; Sacatepe"quez; Chimaltenango; Huehuetenango;
Quezaltenango. Mexico; Colombia to Bolivia.

Usually a small tree, 5-12 meters high; leaves on rather short petioles, broadly
ovate to lance-ovate, mostly 6-15 cm. long, acuminate or attenuate-acuminate,
truncate or rounded at the base, serrate or crenate-serrate, at least at first usually
abundantly pilose beneath but in age often almost glabrous; flower spikes mostly
1-2 cm. long; fruit of few or numerous, very juicy, red or almost black, small
drupes.

The tree seems to be of scattered occurrence and nowhere in
Guatemala has been observed as common. The fruit is edible.
This species, like the following one, has a remarkably wide distri-
bution, and it is barely possible that characters may be found for
separating the North and South American trees. The few available
specimens of the latter do not show any obvious differences. The
type was collected in Ecuador.

Morus insignis Bureau in DC. Prodr. 17: 247. 1873.

Moist mixed forest, 2,000-3,000 meters; El Progreso; Quezalte-
nango; San Marcos. Costa Rica; Colombia to Ecuador and perhaps
Peru.

A large shrub or a tree, sometimes fruiting when only 4.5 meters tall, but
sometimes 18 meters high, the young branches densely whitish-tomentose; leaves
large and thin, the stout petioles tomentose, short, the blades broadly ovate to
oblong-elliptic, mostly 14-25 cm. long, shortly caudate-acuminate, at the base
usually rounded and more or less oblique, finely and closely serrate, usually finely
bullate, bright green above, slightly roughened, beneath usually pale and densely
soft-pubescent; flower spikes short-pedunculate, very slender and usually lax;
fruiting spikes mostly 5-10 cm. long or even longer, the drupes mostly few and
remote, red.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 51

In Costa Rica the trunk is reported to reach sometimes a diame-
ter of 2 meters; the bark is pale brown and somewhat roughened.
To one familiar with the United States and Old World species of
Moras this is a most remarkable tree because of its fantastically
elongate fruits. If only these were of quality proportional to their
size, the tree would be a most desirable one for cultivation, but
unfortunately the drupes are so few that the fruit is quite worthless,
except as food for birds. M. insignis is doubtless the most distinct
and best-marked species of the genus. The Guatemalan form has
the leaves densely soft-pubescent beneath, as in South American
material, but the Costa Rican form differs, perhaps varietally, in
having glabrate leaves. The tree is plentiful at some places in the
barranco of the Rio Samala in the region of Santa Maria de Jesus.

POULSENIA Eggers

Reference: Standley, Trop. Woods 33: 4-5. 1933.

A large or medium-sized tree, the stipules and branchlets armed with prickles;
stipules large and clasping, deciduous; leaves large, somewhat distichous, entire,
often coriaceous, penninerved, variable in size and shape; flowers monoecious,
axillary, the staminate in pedunculate globose heads, the perianth of 4 segments,
the 2 inner ones imbricate; stamens 4, 2 of them longer than the others; pistillate
inflorescences small and few-flowered, the receptacle obscurely bracteate, or the
bracts coalescent, the perianth tubular, 4-dentate; ovary semi-inferior, the ovule
pendulous from the apex of the cell, anatropous; style short, thick, included, the
2 stigmas short, narrow, acute; fruit a somewhat fleshy and juicy syncarp, the
individual fruits covered by the accrescent and somewhat coriaceous perianth;
seed oval, the embryo straight, the cotyledons inrolled.

The genus consists of only a single species, of unusually wide
distribution.

Poulsenia armata (Miq.) Standl. Trop. Woods 33: 4. 1933.
Olmedia armata Miq. in Seem. Bot. Voy. Herald 196. 1854. P.
aculeata Eggers, Bot. Centralbl. 73: 66 (err. typ. "50"). 1898.
Inophloeum armatum Pittier, Journ. Wash. Acad. Sci. 6: 114. 1916.
Coussapoa Rekoi Standl. Contr. U. S. Nat. Herb. 20: 211. 1919.

Wet forest, at or little above sea level; Pete"n; Izabal. Oaxaca
and Veracruz to Chiapas and British Honduras; Honduras; Costa
Rica; Panama; Colombia; Ecuador.

A large tree, sometimes 25 meters high, with a dense, irregular or rounded
crown, the trunk straight, rounded or somewhat compressed, 30-60 cm. or more
in diameter, often with small narrow buttresses; bark brownish, when cut exuding
abundant cream-colored latex; branchlets, petioles, leaves, and stipules armed with

52 FIELDIANA: BOTANY, VOLUME 24

short stout prickles; leaves large, on petioles 1.5-2.5 cm. long, the blades rounded-
ovate to elliptic, mostly 14-40 cm. long and 11-25 cm. wide, very oblique, at least
the larger ones, rounded or obtuse at the apex and apiculate or short-acuminate,
rounded at the base, glabrous; stipules 2-2.5 cm. long or larger; staminate inflores-
cences globose, about 12 mm. in diameter, on peduncles of about the same length,
many-flowered; pistillate receptacles sessile or nearly so, mostly 3-7-flowered, the
perianth about 6 mm. long; fruit 1.5-2.5 cm. in diameter.

Local names reported are "chirimoya" and "carnero" (Oaxaca);
"abababite," "huichilama" (Veracruz); "mastate" (Panama). The
ripe fruits are edible and are sometimes sold in the markets of Vera-
cruz. They somewhat resemble small chirimoyas (Annona Cheri-
mola), hence the name "chirimoya." The Indians of Panama soak
the bark in water and beat it out into a coarse fabric that they
employ for hammocks, blankets, and women's clothes. The inner
bark is very thick and composed of numerous layers of strong crossed
fibers. Similar use of bark of Moraceae is made in many parts of
the earth by primitive people, who sometimes, as in the Pacific
islands, have made really handsome fabrics from it. It is quite
probable that bark of some of the Guatemalan Moraceae may have
been used in this manner by the Mayas or other Indians of northern
Guatemala. The tree is easy of recognition because of the prickly
branches and stipules. Its wood is yellowish brown.

POUROUMA Aublet

Trees; leaves alternate, usually long-petiolate, entire or more often palmately
parted or lobate, the lobes or the blade entire, conspicuously parallel-veined,
coriaceous or often membranaceous, usually tomentose beneath; stipules large,
connate and spathe-like, caducous; peduncles axillary, solitary or geminate,
cymose-branched; flowers dioecious, the staminate glomerulate or capitate, the
pistillate cymose, sessile and often crowded; staminate perianth ovoid or globose,
with 3-4 teeth or lobes, the lobes subvalvate; stamens 3-4, the filaments erect,
free or connate at the base, the anthers ovate; pistillate perianth tubular, with a
small aperture at the apex; ovary included, the style short, the stigma exserted,
peltate-discoid, densely papillose; ovule affixed laterally above the base, ascending,
shortly amphitropous, the micropyle terminal; fruit ovoid, relatively large,
included in the accrescent, fleshy or juicy perianth, the pericarp crustaceous or
hard; seed laterally affixed above the base, the funicle erect from the base of the
cell, the testa membranaceous; embryo straight, the cotyledons thick, oblong,
the very small radicle superior.

About 25 species, all except the following South American.

Pourouma aspera Tre"cul, Ann. Sci. Nat. III. 8: 102. 1847.
Guarumo de montana; Trumpet (British Honduras).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 53

Wet forest of the North Coast, at or little above sea level;
Izabal. British Honduras; Honduras; Nicaragua; Costa Rica;
Panama; Venezuela and the Guianas.

A large tree, often 15 meters tall with a trunk 30 cm. or more in diameter,
with a tall naked trunk and rounded crown; leaves on very long, terete petioles,
the blades 20-30 cm. long or larger, cordate at the base, deeply 3-5-lobate, in
young leaves often divided almost to the base, the lobes oblong to broadly elliptic,
short-acuminate, often abruptly so, entire, appressed-pilose along the nerves,
pale beneath or even whitish, covered with a minute close tomentum, the veins
very prominent and numerous; stipules large, caducous; inflorescences long-
pedunculate, cymose-paniculate, usually about equaling the petioles; fruits ovoid,
1.5 cm. long, minutely and very densely scaberulous, purplish black and juicy at
maturity.

The bark is smooth, mottled in various shades of brown, mauve,
and gray. When freshly cut, the stump exudes a quantity of watery
sap. In Guatemala macaws seem to be fond of the ripe fruits, and
it is stated that the Indians of Costa Rica and other regions also eat
them.

PSEUDOLMEDIA Tre^cul
Reference: H. Pittier, Contr. U. S. Nat. Herb. 13: 432-433. 1912.

Shrubs or often tall trees; stipules small, caducous; leaves alternate, short-
petiolate, entire, coriaceous; flowers dioecious, the staminate in sessile heads, the
pistillate solitary, sessile, surrounded by numerous imbricate bracts; perianth
none in the staminate flowers, the stamens irregularly scattered over the surface
of the receptacle, the filaments short, erect, the anthers oblong; perianth of the
pistillate flower ovoid or tubular, with a small opening at the apex; ovary included,
adnate on one side at the base, the style filiform, the branches exserted, subequal;
ovule pendulous from the apex of the cell; fruit ovoid, included in the enlarged
and fleshy perianth, the bracts unchanged in fruit; pericarp crustaceous; testa of
the seed membranaceous, the endosperm scant or none, the cotyledons thick-
fleshy, very unequal, the radicle small, superior.

About 20 species are known in tropical America. The only
other Central American one, P. mollis Standl., with softly pilose
leaves, occurs in Salvador (type from Comasagua; local name
"tepeujushte") and is to be expected in the Oriente of Guatemala.

Leaves densely long-pilose on the lower surface; branchlets densely hirsute with
long spreading hairs P. simiarum.

Leaves and branchlets glabrous or nearly so.

Lateral nerves of the leaves 15-20 pairs; bracts of the inflorescence densely
sericeous P. oxyphyllaria,

Lateral nerves of the leaves 10-12 pairs; bracts of the inflorescence glabrate.

P. spuria.

54 FIELDIANA: BOTANY, VOLUME 24

Pseudolmedia oxyphyllaria Bonn. Smith, Bot. Gaz. 20: 294.
1895. Manax (Pete'n, Maya).

Moist forest, ranging from sea level to about 1,800 meters;
Pete'n; Izabal; Santa Rosa (type from Volcan de Tecuamburro,
Heyde & Lux 4429) ; Quezaltenango (Chiquihuite) ; Huehuetenango.
Veracruz and Oaxaca; British Honduras; Costa Rica.

A tree of 6-9 meters, or sometimes as much as 30 meters high; stipules narrow,
2 cm. long or less, sparsely sericeous outside; leaves on very short petioles, lance-
oblong or narrowly oblong, mostly 10-19 cm. long and 3-6.5 cm. wide, acuminate,
usually abruptly so, rounded to subacute at the base, glabrous or nearly so, some-
what paler beneath; staminate heads solitary or glomerate, about 5 mm. broad,
the bracts obtuse, fulvous-sericeous; fruit oval or ellipsoid, 1-2 cm. long.

Called "cherry" in British Honduras.

Pseudolmedia simiarum Standl. & Steyerm. Field Mus. Bot.
23: 154. 1944. Durazno de mono; Durazno de monte.

Dense wet mixed forest, 1,500-1,600 meters; endemic; Huehue-
tenango (type from Maxbal, about 17 miles north of Barillas,
Steyermark 48741; collected also between Maxbal and Xoxlac).

A tall tree as much as 30 meters high, the trunk sometimes 60 cm. in diameter,
the branchlets stout, somewhat flexuous, densely hirsute with long spreading
fulvescent soft hairs; stipules caducous, as much as 2.5 cm. long, hirsute; leaves
on short stout petioles 7-10 mm. long, oblong-elliptic, 18-27 cm. long, 7.5-13 cm.
wide, abruptly caudate-acuminate, obliquely rounded at the base, green and almost
glabrous above, puberulent or hirtellous on the costa and nerves, paler and brown-
ish beneath, rather densely pilose with long slender spreading soft hairs, the
lateral nerves about 17 pairs, arcuate, prominent, the veins prominent and laxly
reticulate; pistillate inflorescences axillary, apparently sessile; immature fruit
globose or oval-globose, 2-2.5 cm. long, rounded at the base and apex, very densely
and softly pilose with long yellowish hairs; bracts persistent, rounded-ovate,
obtuse, 5-6 mm. long, densely sericeous-pilose on both surfaces.

The bark exudes a cream-colored milk-like sap when cut. The
local names refer to the fact that the fruits resemble young and
immature peaches.

Pseudolmedia spuria (Swartz) Griseb. Fl. Brit. W. Ind. 152.
1859. Brosimum spurium Swartz, Prodr. Veg. Ind. Occ. 12. 1788.
P. havanensis Tre"cul, Ann. Sci. Nat. III. 8: 130. 1847. Manax
(Pete'n, Maya).

Common in climax forest, northern Pete'n; Izabal, wet forest at
sea level. Greater Antilles; reported, probably in error, from
Panama.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 55

A large tree with thin bark; stipules narrow, long-attenuate, 1 cm. long or
less, glabrous; leaves on very short petioles, often almost sessile, coriaceous, lance-
oblong or elliptic-oblong, mostly 8-15 cm. long and 3-5 cm. wide, rather abruptly
obtuse-acuminate, acute or obtuse at the base, somewhat paler beneath ; staminate
heads globose, 4 mm. in diameter; fruit ovoid, 1-1.5 cm. long, turning bright red at
maturity.

Called "cherry" in British Honduras. The red fruits are reported
to have a delicious flavor and are much eaten in regions where the
tree occurs. The trunk is said to yield a latex that flows easily but
is hard to collect. Quite probably it is used to adulterate chicle.
The wood is light brown, hard, heavy, tough, coarse- textured,
splintery, not durable. So far as we know, it is not utilized.

SOROCEA St. Hilaire

Shrubs or small trees with milky sap; leaves short-petiolate, entire or dentate,
penninerved; stipules small, caducous; flowers dioecious, in ament-like spikes or
racemes, usually rather lax or distant upon the rachis; staminate perianth 4-parted,
the segments broad, imbricate; stamens 4, the filaments free, erect, finally exserted,
the anthers ovate; pistillate perianth ovoid or tubular, with a small aperture at the
apex; ovary inferior, the style fleshy, ovoid-conic, short-attenuate at the apex, the
short branches exserted, spreading; ovule pendulous, affixed at or near the apex
of the cell; fruit enclosed in the accrescent perianth; seed pendulous, the testa
membranaceous; endosperm none, the embryo curved, the cotyledons unequal.

About 15 species in tropical America.

Sorocea pubivena Hemsl. Biol. Centr. Amer. Bot. 3: 150. 1883.
Type cited as from Guatemala, collected by Friedrichsthal.

Branchlets slender, glabrous; petioles 12-16 mm. long; leaf blades oblong-
elliptic, as much as 25 cm. long, caudate-acuminate, cuneate at the base, entire,
puberulent beneath, especially on the veins, glabrous above; staminate flowers on
slender pedicels 4-6 mm. long; fruits puberulent, oblong, about 8 mm. long, not
muricate.

We have seen no representation of this species, whose status is
altogether doubtful. It is quite possible that it does not belong to
the genus Sorocea, and even more probable that it. was not collected
in Guatemala. While all the Friedrichsthal plants were supplied
with labels bearing the heading "Guatemala," a large percentage
of them really came from Nicaragua and Costa Rica, an error that
has caused much confusion in the systematic botany of Central
America. Only by examination of the original labels at Vienna can
the localities be confirmed, and in some instances, unfortunately, the
Vienna labels do not bear accurate locality data.

56 FIELDIANA: BOTANY, VOLUME 24

TROPHIS L.

Trees or shrubs; stipules lateral, small, caducous; leaves alternate, short-
petiolate, membranaceous to coriaceous, entire or dentate, on young branches
sometimes lobate, penninerved; flowers dioecious, spicate or racemose, the inflores-
cences solitary or geminate in the leaf axils; flowers sessile or short-pedicellate, the
bracts minute; staminate perianth 4-parted or 4-lobate, the lobes valvate; stamens
4, the filaments in bud inflexed, in anthesis porrect and exserted; pistillate perianth
tubular, adnate to the ovary, 4-dentate at the orifice; ovary inferior; apex of the
style exserted, the branches short or elongate, filiform, usually recurved; fruit
globose, fleshy, concrete with the enlarged perianth; seed globose, with thin testa;
endosperm none; embryo straight, the cotyledons fleshy, equal, semiglobose, the
radicle very short, superior.

Probably 10 or more species, in tropical America. One other
Central American one is known from Panama and Costa Rica.

Leaves densely and softly short-pilose beneath T. cuspidata.

Leaves glabrous beneath or scabrous.

Leaves very scabrous on the upper surface and rough to the touch; fruit smooth,

not tuberculate T. racemosa.

Leaves not scabrous, smooth to the touch; fruit tuberculate or strongly rugose.
Leaf blades linear-lanceolate or very narrowly oblong-lanceolate, mostly 4-6

times as long as wide, 2.5 cm. wide or less T. chiapensis.

Leaf blades oblanceolate-elliptic to elliptic, usually 2-3 times as long as wide,
mostly 3-4.5 cm. wide T. chorizantha.

Trophis chiapensis Brandeg. Univ. Calif. Publ. Bot. 6: 178.
1915. T. nubium Standl. Field Mus. Bot. 22: 17. 1940. Cerezo de
montana.

Wet mixed mountain forest, 900-2,200 meters; Solola; Suchite-
pe"quez; Quezaltenango (type of T. nubium from Volcan de Zunil,
in second-growth thicket, Skutch 925); San Marcos. Chiapas, the
type from Cerro del Boqueron, Purpus 7091.

A shrub or a small tree 15 meters tall, the branches very slender, puberulent
or glabrous; stipules triangular, 3 mm. long; leaves on petioles 6-9 mm. long,
mostly linear-lanceolate or very narrowly oblong-oblanceolate, 8-14 cm. long,
1.5-4.5 cm. wide, very narrowly long-attenuate-acuminate, at the base obtuse or
subacute, glabrous, the lateral nerves about 15 pairs, divergent at a wide angle,
the margin closely serrate; pistillate racemes mostly 2 cm. long or less, lax and
few-flowered, short-pedunculate, the rachis densely tomentulose or in age glabrate,
the pedicels mostly 2-3 mm. long, or in age as much as 1 cm. long; fruit 6-8 mm.
long, subglobose, glabrate, densely and coarsely tuberculate (tubercles not always
apparent in young fruit).

There is a slight possibility that when more ample material is
available, it will be found that T. nubium is a distinct species, since
the pistillate inflorescences are densely tomentulose and the pedicels
short, but this is probably a mere matter of development.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 57

Trophis chorizantha Standl. Field Mus. Bot. 4: 302. 1929
(type from Lancetilla Valley near Tela, Honduras). Skutchia
caudata Pax & Hoffm. in Morton, Journ. Wash. Acad. Sci. 27: 307.
1937 (type from Costa Rica). T. Matudai Lundell, Lloydia 2: 81.
1939 (type collected on Mount Ovando, near Escuintla, Chiapas,
E. Matuda 2091). Palo morillo (fide Aguilar).

Moist or wet, mixed, mountain forest, 2,500 meters or lower;
Pete"n; Alta Verapaz; El Progreso; Izabal; Escuintla; Sacatepe"quez;
Chimaltenango; Suchitepe"quez ; Quezaltenango ; San Marcos.
Oaxaca(?); Chiapas; British Honduras; Atlantic coast of Honduras;
Costa Rica.

A large shrub or small tree, sometimes as much as 15 meters tall, the trunk
25 cm. or somewhat more in diameter, the branchlets slender, sparsely puberulent
or glabrate; stipules subulate, about 1.5 mm. long; leaves on very short petioles,
membranaceous, bright green above, somewhat paler beneath, oblong or obovate-
oblong, mostly 9-15 cm. long and 3.5-5 cm. wide, abruptly caudate-acuminate,
acute or subobtuse at the base, glabrous and smooth, the lateral nerves about
8 pairs, the margins subentire (especially in leaves of fertile branches) or often
coarsely dentate or serrate on young branches; staminate spikes solitary or gemi-
nate, almost sessile, short and dense; pistillate spikes or racemes very variable,
short or elongate and sometimes as much as 12 cm. long, remotely few-flowered,
the flowers sessile or often on stout pedicels; stigmas slender and elongate; fruit
red at maturity, glabrate, globose, coarsely tuberculate, 6-7 mm. in diameter.

A somewhat variable tree, of which a large number of specimens
have been collected, all of which seem undoubtedly conspecific.
There is some question as to whether T. chorizantha is different from
T. mexicana (Liebm.) Bureau, of Veracruz, but the two seem reason-
ably distinct in foliage characters, as shown by type material of
T. mexicana available for comparison.

Trophis cuspidata Lundell, Amer. Midi. Nat. 19: 427. 1938.

Type from Mount Ovando, near Escuintla, Chiapas, Matuda
1051; collected also on Volcan de Tacana, 2,000-4,000 meters, and
doubtless extending into San Marcos.

A tree, the young branchlets densely short-pilose; stipules 3-4 mm. long, the
petioles 10-14 mm. long; leaf blades oblong or narrowly oblong, 9-18 cm. long,
3.5-7 cm. wide, long-acuminate, obtuse or rounded and somewhat unequal at
the base, thick and firm, glabrous above, densely short-pilose beneath, the lateral
nerves 8-12 pairs; staminate spikes solitary in the leaf axils, 2.5-4.5 cm. long;
pistillate racemes solitary, short or much elongate, densely tomentulose, lax and
remotely flowered, the pedicels 1 cm. long or less; immature fruits globose-obovoid,
tomentulose, apparently somewhat tuberculate, the persistent stigmas short and
broad.

58 FIELDIANA: BOTANY, VOLUME 24

Trophis racemosa (L.) Urban, Symb. Antill. 4: 195. 1903.
Bucephalon racemosum L. Sp. PI. 1190. 1753. Trophis americana
L. Syst. Nat. ed. 10. 1289. 1759. Sahagunia urophylla Donn. Smith,
Bot. Gaz. 40: 11. 1905 (type from the north coast of Honduras,
Tela). Ramon Colorado (Pete"n); Yaxox, Catalox (Pete"n, Maya).

Moist or wet, sometimes dry, usually mixed forest, or in thickets,
ascending from sea level to about 1,500 meters; Pete"n; Alta Verapaz;
Izabal; Zacapa; Chiquimula; Santa Rosa; Escuintla; Suchitepe"quez ;
Retalhuleu; Quezaltenango; Huehuetenango. Mexico and British
Honduras to Panama; West Indies; northern coast of South America.

A tall shrub or a tree, sometimes 18 meters tall with a trunk 50 cm. in diameter,
the crown dense, the branches sometimes drooping, the bark brown; leaves short-
petiolate, oblong to oval or obovate, mostly 8-15 cm. long, short-acuminate or
cuspidate-acuminate, obtuse and somewhat unequal at the base, entire or obscurely
serrate, scabrous and rough above or sometimes smooth, beneath scabrous or
glabrous; staminate spikes elongate, dense or interrupted; pistillate spikes few-
flowered, the flowers sessile or nearly so, densely pubescent; fruit subglobose,
fleshy, red at maturity, 1 cm. or less in diameter.

The Maya name is reported from Yucatan as "chacox"; called
"white ramon" in British Honduras; in Honduras "ramon," "San
Ramon"(?), and "hoja tinta"; in Salvador "raspa-lengua," "ojushte,"
"ujushte," "chilujushte," "chulujushte," and "pilijushte" ; "ramon-
cillo" (Tabasco). The fruit is edible but not particularly palatable,
and has but scant flesh. The young branches and leaves are much
used in Pete"n, Yucatan, and elsewhere, like those of Brosimum, as
fodder for cattle and other stock during the dry season. The yellow-
ish wood is used as firewood and sometimes for other purposes.

PROTEACEAE

Shrubs or trees; leaves alternate, rarely opposite or verticillate, entire or
dentate, or sometimes simple and pinnate upon the same plant, commonly coria-
ceous; stipules none; flowers perfect, often large and showy, by abortion sometimes
polygamous or dioecious, capitate-spicate, racemose or rarely solitary, scattered
and solitary along the rachis, or in pairs and subtended by a bract, the whole
inflorescence in fruit sometimes strobiliform; perianth inferior, the 4 segments
valvately coherent at first and forming a cylindric tube, often separating in
anthesis and recurving; stamens 4, opposite the perianth segments and affixed to
them, shorter than the perianth, the filaments short or almost none; anthers erect,
all perfect or one of them abortive, the connective continuous with the filament,
the 2 cells introrsely adnate, parallel; squamellae or hypogynous glands often
present, alternate with the stamens; ovary free, sessile or stipitate, 1-celled, usually
oblique; style terminal, short or elongate, usually thickened at the apex, the stigma
small, terminal or sublateral; ovules solitary or geminate, or numerous and biseri-
ate, ascending or descending; fruit sometimes nut-like or drupaceous and inde-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 59

hiscent, sometimes dehiscent and follicular or capsular, the valves usually thick
and coriaceous; seeds 1-2 or few, with membranaceous or coriaceous testa, some-
times winged; endosperm none.

About 54 genera and 1,000 species or more, mostly in Australia
and South Africa, only a few species in other continents. One
other genus, Panopsis, is represented in Costa Rica and Panama.

Ovules ascending; leaves sericeous beneath, pinnate, the leaflets often deeply

cleft into narrow segments Grevillea.

Ovules pendulous; leaves simple or pinnate, the leaflets merely dentate or entire.

Roupala.

GREVILLEA R. Br.

Shrubs or trees, the leaves various in form, pinnate in the species cultivated
in Central America; flowers perfect, regular or irregular, geminate, pedicellate,
racemose, the racemes terminal and sometimes also axillary; perianth tube slender,
straight, sometimes dilated at the base and recurved or revolute below the limb,
usually cleft on the lower side in anthesis, the limb oblique; anthers sessile in pits
in the blades of the perianth segments, ovate or oblong, the connective not pro-
duced beyond the cells; disk carnose, sometimes none; ovary stipitate or subsessile;
style usually elongate and protruding from the cleft in the perianth tube, persis-
tent; ovules 2, collateral, laterally affixed; fruit follicular, dehiscent by the strongly
curved outer side, sometimes lignescent; seeds 2 or by abortion 1, plane-com-
pressed, usually winged.

More than 160 species, nearly all Australian, a few in New
Caledonia.

Leaf segments entire; flowers bright deep red G. Banksii.

Leaf segments deeply pinnatifid; flowers yellow G. robusta.

Grevillea Banksii R. Br. Trans. Linn. Soc. 10: 176. 1810.

Cultivated occasionally for ornament; seen in Guatemala, near
Chimaltenango, and at Santa Cruz, Alta Verapaz; probably also
elsewhere, but scarce. Native of Australia.

A shrub or small tree, flowering when only a meter high, the branches rather
densely sericeous or tomentose; leaves pinnate or deeply pinnatifid, with 3-11
segments, these linear or nearly so, green and thinly sericeous above, whitish and
densely sericeous beneath; racemes terminal, mostly 5-10 cm. long, the flowers
deep bright red, tomentose outside; follicles densely tomentose, 1.5-2 cm. long.

This species seems to be of recent introduction into Central
America, and still is infrequent. We have seen specimens also from
Costa Rica. In beauty it is far superior to G. robusta, because of the
brilliancy and attractive coloring of its flowers.

Grevillea robusta A. Cunn. Suppl. Prodr. Nov. Holl. 24. 1830.

Gravilea; Peineta; Talnete (flowers).

60 FIELDIANA: BOTANY, VOLUME 24

Native of eastern Australia. Cultivated for ornament or as a
shade tree in almost all parts of Guatemala, especially in the cooler
regions; abundantly planted for coffee shade in the central highlands;
often escaping and naturalized along roadsides and in thickets.

A medium-sized or often large tree, frequently 15 meters high, the branchlets
ferruginous- or grayish-tomentose; leaves large, petiolate, pinnate, the numerous
leaflets cleft into narrow long-attenuate lobes, gray-green above, sericeous beneath
with brown or silvery hairs; racemes often panicled, terminal, 12-18 cm. long, the
golden-yellow flowers long-pedicellate, glabrous; follicles 1.5 cm. long, glabrous.

In Guatemala, as in other parts of Central America, this is one
of the common ornamental and shade trees, and it is often planted
along streets and roads. It is of easy growth and survives neglect
and mistreatment. The young plants are handsome, and often are
grown in the United States as pot plants, under the name "Australian
silk oak," but the large trees, although bearing in winter and spring
great quantities of bright-colored flowers, are less attractive,
especially if they happen to be covered with dust, as often happens.
Some people, however, admire them, and in recent years many
young trees have been planted by the government along the roads
of Guatemala. In this country Grevillea also has an important part
in the coffee industry. Practically all the many cafetales of the valley
of Antigua (1,500 meters) are densely shaded with the tree, likewise
the scattered coffee plantations of the highlands of Chimaltenango,
some of them at as great an elevation as 1,800 meters. A few
cafetales in the higher parts of Quezaltenango have the same tree as
shade, but it is only in the Sacatepe"quez-Chimaltenango region
that it is important, and when one views the Antigua region from
some eminence, it appears one great forest of Grevillea. The coffee
here needs protection from cold misty nights and from cold winds,
and for this purpose this tree has been found more satisfactory than
anything else. So far as known, Grevillea is not used elsewhere for
this purpose, at least in Central America. The flowers are said to
give large amounts of honey, but of dark color and not particularly
good flavor. The wood is said to be elastic and durable, and used
in Australia for furniture and barrel staves, but no use is made of
it in Central America, although it could be grown easily in large
amounts. It is stated that in Australia trees are 6-9 meters high
at an age of 20 years, but in Central America growth is evidently
more rapid.

ROUPALA Aublet

Trees, glabrous or tomentose; leaves alternate, coriaceous, dimorphous, those
of adult flowering branches usually simple and entire or dentate, those of sterile

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 61

branches or of young plants pinnate; flowers perfect, regular, racemose, geminate
and pedicellate, the racemes axillary or lateral; bracts none; perianth cylindric,
straight, the segments separating in anthesis and revolute; stamens affixed at the
base of the perianth segments, the filaments short; anthers oblong-linear, the cells
imperfectly separated, the connective short-produced at the apex; hypogynous
scales 4, distinct, plane, obtuse or acute; ovary sessile; ovules 2, collateral, pendu-
lous from the apex of the cell; follicles hard and ligneous, obliquely bivalvate,
short-stipitate; seeds compressed, winged.

About 30 species in tropical America, chiefly in mountain regions.
It is doubtful whether more than two species occur in Central
America, although three others of questionable validity have been
recorded or described from Costa Rica and Panama. R. loranthoides
Meisn. (in DC. Prodr. 14: 425. 1856) was published as Guatemalan.
A photograph and fragment of the type, collected by Friedrichsthal,
are in the Herbarium of Chicago Museum. The locality of the label is
Monte Rincon, which may well be Rincon de la Vieja in Guanacaste,
Costa Rica, and probably is not Guatemalan. The species is note-
worthy for its very obtuse, emarginate leaves whose veins and nerves
are impressed on the lower surface. The type is not matched by any
Central American specimens.

Roupala borealis Hemsl. Biol. Centr. Amer. Bot. 3: 78. pi. 76.

1882. R. repanda Lundell, Amer. Midi. Nat. 29: 472. 1943 (type
from Monkey River, Toledo District, in hammock on pine ridge,
British Honduras, P. H. Gentle 4196). Zorrillo; Zorro.

Moist or wet forest, sometimes in open mountain pastures,
800-2,400 meters; Pete'n (near British Honduras boundary); Alta
Verapaz; Zacapa; Chiquimula; Santa Rosa; Escuintla; Guatemala;
Sacatepe'quez ; Quezaltenango; Huehuetenango. Southern Mexico;
British Honduras; Salvador, and probably south to Panama.

A small to rather large tree, usually 7-12 meters high or larger, the trunk
often 30 cm. or more in diameter, the branchlets pilose or tomentose with fer-
ruginous or grayish hairs, often glabrate; leaves coriaceous, very variable, part
of them on very long petioles, ovate to lance-elliptic or elliptic, 5-13 cm. long,
acute to long-acuminate, acute or obtuse at the base, glabrous or nearly so, at
least in age, undulate-dentate or coarsely serrate, sometimes entire; many or most
of the leaves pinnate, with 3-17 leaflets, these asymmetric, more or less rhombic,
coarsely dentate or undulate-serrate or sometimes almost laciniate; flowers white
or whitish, the racemes slender, mostly shorter than the leaves, many-flowered,
the rachis minutely puberulent; pedicels 4-5 mm. long, puberulent or sericeous
with whitish or brownish hairs, spreading at right angles; perianth slender, 12 mm.
long, sparsely and minutely puberulent outside; ovary densely short-pilose.

Sometimes called "chancho" or "palo de chancho" in Salvador.
The wood and foliage have a strong mephitic odor, hence the usual

62 FIELDIANA: BOTANY, VOLUME 24

name of "zorrillo." The Central American material of Roupala,
although rather voluminous, is not sufficiently ample to make pos-
sible a satisfactory disposition of the forms, if there are more than
two. It is quite possible that R. borealis should be united with R.
complicate, HBK., described from Venezuela, to which species some
of the specimens have been referred. One collection from Alta
Verapaz is notable for the abundant ferruginous pubescence on the
younger leaves and branches, and quite possibly represents a dis-
tinct and perhaps undescribed species. The wood is brown or red-
dish, hard, and heavy. We have not seen material of R. repanda,
but from description there is no reason to suppose that it differs in
any respect from R. borealis, of which several British Honduras
specimens are at hand.

LORANTHACEAE. Mistletoe Family

References: A. G. Eichler, Loranthaceae in Mart. Fl. Bras. 5,
pt. 2: 1-135. pis. 1-44. 1868. Ignatius Urban, Addimenta ad cogni-
tionem florae Indiae occidentalis, Particula IV. Loranthaceae, Bot.
Jahrb. 24: 10-76. 1897.

Parasitic shrubs, usually containing chlorophyll, growing on woody plants
and absorbing food from their sap through specialized roots called haustoria,
rarely terrestrial shrubs or small trees; branches terete or ungulate, usually articu-
late at the nodes, mostly glabrous but sometimes pubescent; leaves opposite,
sometimes reduced to scales, rarely alternate; flowers mostly very small, some-
times large and showy, perfect, or unisexual and monoecious or dioecious, in
axillary or terminal racemes, spikes, or panicles, sometimes solitary; perianth
1-2-seriate, symmetric, green, yellow, or red; calyx tube adnate to the ovary, the
limb usually much reduced; stamens 2-6; anthers 2-celled, the cells parallel,
longitudinally dehiscent, rarely 1-celled with the cells confluent and dehiscent by
a transverse pore or slit; disk usually present, sometimes none; ovary 1 and 1-celled,
the style simple or none, the stigma terminal; fruit generally small, baccate, the
pulp viscid; seeds mostly very small; embryo terete or angulate.

About 20 genera and 500 species, widely distributed, mostly in
tropical regions. The only other genus known in Central America
(Costa Rica) is Gaiadendron, a large terrestrial shrub or small tree.
The whole family has received little systematic attention in recent
years, and is seriously in need of careful revision. Both generic and
specific limits are often vague, and recognizable with difficulty.

Leaves all reduced to scales; perianth simple, no corolla present.

Flowers solitary in the axils of opposite connate scales Arceuthobium.

Flowers inserted on the joints of the flower spike between the nodes, usually

1-seriate and superposed Dendrophthora.

Leaves with well-developed blades; corolla present or absent.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 63

Perianth simple, no corolla present.

Flowers immersed in pits in the rachis of the spike; filaments short.

Phoradendron.
Flowers not immersed in the axis; filaments longer than the anthers.

Antidaphne.
Perianth double, both calyx and corolla present.

Corolla large, commonly 1.5-8 cm. long, usually bright red. . . .Psittacanthus.
Corolla small, much less than 1 cm. long, not red.

Flowers immersed in pits in the axis of the inflorescence Oryctanthus.

Flowers not immersed in the axis of the inflorescence.

Filaments subulate; plants usually not at all furfuraceous . . . Struthanthus.

Filaments stout; young plants usually densely ferruginous-furfuraceous

on the angles of the stems and margins of the leaves, in age usually

glabrous Phthirusa.

ANTIDAPHNE Poeppig & Endlicher

Small glabrous epiphytic shrubs; leaves alternate, broad, thick; flowers
spicate, monoecious or dioecious, the spikes sessile in the leaf axils, small, the
staminate subglobose or ovoid, strobiliform, the bracts scale-like, broadly imbri-
cate, caducous in an thesis, subtending 1-3 pedicellate flowers; pistillate spikes
subtended at the base by a few imbricate bracts, the floriferous part elongating,
ebracteate in anthesis, the flowers sessile in groups of 3-5, the rachis in age elongat-
ing into a leafy branchlet, the fruits often persistent on the branchlet below the
leaves; perianth none in the staminate flower, in the pistillate flower adnate to the
ovary, the margin minutely and remotely 3-4-dentate; stamens 3-5, inserted
about a small fleshy disk, the filaments elongate, very unequal; anthers ovate or
oblong, erect, the cells parallel, longitudinally dehiscent; berry ovoid, the pericarp
fleshy and viscid.

One other species is known, in South America.

Antidaphne viscoidea Poepp. & Endl. Nov. Gen. & Sp. 2: 70.
pi. 199. 1838. Liga.

On trees in forest, 1,400-2,600 meters; Alta Verapaz; Guate-
mala; Chimaltenango; Solola; Quiche"; San Marcos. Chiapas; Costa
Rica; Panama; southward to Bolivia.

A small, usually densely branched shrub, glabrous, the branches generally
50 cm. long or shorter, terete or nearly so, stout; leaves almost sessile or on very
short, thick petioles, obovate to suborbicular, 3-7 cm. long, broadly rounded at
the apex, acute at the base, the nerves and veins very prominent in the dry state
and openly reticulate; flowers cream-colored, the staminate spikes very small, 8
mm. long or shorter, their bracts at first conspicuous but soon deciduous; berry
oval.

ARCEUTHOBIUM Bieberstein

Parasitic shrubs, usually growing on Coniferae, commonly small, glabrous,
branched, the branches stout, articulate; leaves reduced to small scales, these

64 FIELDIANA: BOTANY, VOLUME 24

opposite, connate into small open sheaths; flowers dioecious, solitary in the axils
of the bracts, sessile or subsessile, not bracteolate; perianth tube almost obsolete
in the staminate flower, in the pistillate flower adnate to the ovary, the limb 2-5-
parted in the staminate flower, in the pistillate flower minute, 2-parted; anthers
sessile, transverse, the cells confluent, dehiscent by a single slit, in age almost
orbicular; disk carnose; ovary ovoid, the style short and thick, subconic, the stigma
obtuse; berry ovoid, short-stipitate, capped by the minute perianth lobes, the
pericarp fleshy, viscid, at maturity dehiscent at the base and elastically dehiscent,
often ejecting the seed to a considerable distance; seed ovoid-oblong; endosperm
carnose, copious.

About 6 species, one in southern Europe and western Asia, the
others North American. Only one occurs in Central America.

Arceuthobium vaginatum (HBK.) Eichler in Mart. Fl. Bras.
5, pt. 2: 105. 1868. Viscum vaginatum HBK. Nov. Gen. & Sp. 3:
445. 1820.

Parasitic on Pinus and Cupressus, 1,350-3,700 meters; Alta
Verapaz; Baja Verapaz; Zacapa; Huehuetenango; San Marcos.
Southwestern United States; mountains of Mexico.

Plants 10-30 cm. high, glabrous, yellowish brown, much branched, the stems
compressed-quadrangular or the older ones terete, as much as 8 mm. in diameter
at the base, lustrous, fragile, the branches opposite; leaf sheaths small, 2-dentate
at the apex, the teeth or lobes spreading; berries 5 mm. long, borne on stout
pedicels, recurved in age.

This plant is probably common on pine trees in the Guatemalan
mountains, but usually it grows so high on the branches that it is
unseen. It is noteworthy that in the Cuchumatanes the plants may
be found on almost any part of the tree, often in dense colonies
along and toward the base of the trunk. We have not observed
such distribution of the plants in the southwestern United States,
where they usually are confined to the upper branches.

DENDROPHTHORA Eichler

Parasitic shrubs, usually small and rather slender, generally glabrous, the
branches articulate at the nodes, the stems terete or angulate; leaves reduced to
small scales in the Guatemalan species; flowers monoecious or dioecious, sessile,
usually sunken in the rachis of the spike, solitary or several on each side of a joint,
usually superposed in 2 rows, the spikes axillary or terminal, articulate, bracteate
at each node; staminate perianth 3-lobate; filaments wholly adnate to the sepals,
the anthers sessile; pistillate calyx 3-lobate; ovary inferior; fruit baccate; embryo
small, surrounded by copious endosperm.

About 40 species, in tropical America. Two other species, with
well-developed leaves, are found in southern Central America
(Costa Rica and Panama).

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 65

Dendrophthora guatemalensis Standl. Field Mus. Bot. 22:
17. 1940. Paxte de palo.

Parasitic on broad-leafed trees, 350-1,200 meters; Alta Verapaz
(type collected on slopes above Finca Seamay, C. L. Wilson 204);
Suchitepe'quez ; endemic.

Plants slender and much branched, dense, dull dark olive-green, fragile,
glabrous, the branches slender above, terete, the base of the plant as much as
7 mm. in diameter, the ultimate branches scarcely 1 mm. thick, the internodes
7-15 mm. long, very minutely tuberculate; leaf scales scarcely 1 mm. long, rounded
at the apex; spikes axillary, short-pedunculate, 1-3-jointed, the pistillate spikes
usually terminated by a 1-flowered joint; sepals 3, closed, broadly triangular,
obtuse.

This probably is the plant reported from Guatemala by Eichler
as D. biserrula Engler. That species, common in Costa Rica and
Panama, probably is distinct from D. guatemalensis.

ORYCTANTHUS Eichler

Small or rather large shrubs, parasitic on dicotyledonous trees; leaves well
developed, opposite, the blades broad, thick, mostly palmate-nerved; flowers
small, perfect or rarely dioecious, spicate, the spikes sometimes paniculate, the
flowers solitary, opposite-decussate, immersed in pits in the fleshy rachis; bracts
scale-like, bordering the pits, in age obsolete, the bractlets rudimentary or abor-
tive; flowers 6-parted, the perianth 2-seriate, the margin of the calyx subentire,
the inner segments free, spreading in anthesis; filaments adnate below to the inner
perianth segments, free above, filiform-cylindric, attenuate or subulate above;
anthers more or less rounded, 2-celled, dehiscent by 2 longitudinal slits; ovary
obovoid, surrounded by a carnose annular disk, the style cylindric, the stigma
capitate; berry oblong, umbilicate at the apex, the epicarp carnose or subcoria-
ceous, the flesh viscid; endosperm copious, carnose; cotyledons semicylindric.

About 7 species, in tropical America. Two others are known
from southern Central America.

Leaves sessile or nearly so, broadly ovate, rounded or cordate at the base.

O. cordifolius.

Leaves distinctly petiolate, oblong or obovate-oblong, acute at the base.

0. guatemalensis.

Oryctanthus cordifolius (Presl) Urban, Bot. Jahrb. 24: 30.
1897. Viscum cordifolium Presl, Epim. Bot.' 253. 1849.

On various trees or shrubs, 800 meters or less; Pete'n; Izabal;
Quezaltenango. Southern Mexico; British Honduras to Salvador
and Panama.

A small glabrous shrub, usually erect and rather sparsely branched, the young
branches compressed and 2-edged, the older ones terete, sparsely furfuraceous or

66 FIELDIANA: BOTANY, VOLUME 24

glabrate, usually dark brown; leaves sessile or nearly so, broadly ovate, 7-14 cm.
long, 4.5-7 cm. wide, acuminate to obtuse, broadly rounded or cordate at the base,
thick when dried and conspicuously palmate-nerved; spikes fasciculate in the leaf
axils or at the ends of the branches, pedunculate, 10 cm. long or shorter, the small,
dark red or brownish flowers 4-ranked, inserted at a right angle with the rachis;
berries small, red.

Called "suelda con suelda" in Honduras; "hierba del pajaro"
(Salvador). A common parasite in the North Coast region, and
frequent in many parts of the Central American lowlands.

Oryctanthus guatemalensis (Standl.) Standl. & Steyerm.
Field Mus. Bot. 23: 40. 1944. Struthanthus guatemalensis Standl.
Field Mus. Bot. 17: 237. 1937. Liga.

Parasitic on shrubs or trees, 1,200-1,400 meters; endemic;
Suchitepe'quez (type from Finca Moca, J. Bequaert 46); Quezalte-
nango (southern slopes of Volcan de Santa Maria, near Finca
Pirineos).

A shrub about 25 cm. high, densely branched, the branches rather slender,
the younger ones tetragonous, densely ferruginous-furfuraceous on the angles,
the older ones subterete, glabrous, the internodes short; leaves small, on con-
spicuous petioles 3 mm. long, thin-coriaceous, oblong or obovate-oblong, some-
times oblong-ovate, 2.5-7 cm. long, 1-2.5 cm. wide, narrowly rounded or often
somewhat emarginate at the apex, acute at the base, obscurely 3-nerved or more
properly penninerved, when young densely ferruginous-furfuraceous on the mar-
gins and on the salient costa beneath, in age glabrous, the veins sometimes promi-
nent and reticulate beneath; spikes on peduncles 4-5 mm. long, axillary and
aggregate at the ends of the branches, simple or sometimes with 1-2 short basal
branches, scarcely more than 2 cm. long and often shorter, slender, glabrous,
densely flowered, the flowers 10 or more, inserted at about a right angle; bractlets
well developed at the base of the pits of the rachis; berry subglobose, smooth,
glabrous, 3 mm. long, rounded at base and apex.

PHORADENDRON Nuttall. Mistletoe

Reference: William Trelease, The genus Phoradendron, Urbana,
Illinois, 1916.

Small shrubs, parasitic on broad-leafed trees or shrubs or sometimes on Coni-
ferae, the stems easily broken at the nodes; leaves opposite, coriaceous, usually
with well-developed blades (in all Guatemalan species), sometimes reduced to
scales, the branches terete or angulate; flowers small, dioecious or monoecious,
usually sunken in the rachis of the spike, superposed in 2-6 or rarely 8 rows on
each joint of the spike; staminate calyx generally 3-lobate, with an almost sessile,
2-celled anther at the base of each lobe; pistillate calyx adnate to the inferior
1-celled ovary, the ovules solitary; style short, the stigma capitate; berry fleshy,
with viscid pulp; embryo small, the endosperm copious.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 67

Species about 200, or perhaps fewer, all American and mostly
in tropical America. Others are known in southern Central America.
In spite of the elaborate monograph published by Trelease, valuable
for its many illustrations of type specimens, the taxonomy of the
genus is in an unsatisfactory state. Some species are highly variable,
and their characters often inconstant and difficult to evaluate. The
species of the United States are much used for Christmas decora-
tions, being called "mistletoe," a name more properly belonging to
European species of the genus Viscum. The association of the plants
with Christmas is a sentimental one, and derives from ancient use
of the European plant in religious celebrations of the Druids. Plants
of this and other genera of the family often are highly destructive
to trees upon which they grow, ultimately killing them. The seeds
doubtless are spread by birds, which eat the usually more or less
translucent berries, and the seeds and fruits doubtless are spread
also because they adhere to the feet or feathers of birds, or to the
bodies of other animals. The mistletoe of Spain (Viscum) is known
in that country by the names "mue'rdago," "liga," and "visco."

Branches without scales on any of the joints.

Branches not or scarcely compressed, densely pubescent; leaves lanceolate.

P. velutinum.
Branches strongly compressed at the nodes, glabrous or nearly so; leaves

almost linear P. uspantanum.

Branches with scales at the base of the joints, at least on the lowest joint of each

branch.
Scales present on all the joints of the branches.

Scales of the branches bearing flower spikes in their axils .... P. crassifolium.
Scales without flower spikes in their axils.

Leaves palmate-nerved, the nerves all arising from the base of the blade.

P. supravenulosum.

Leaves penninerved P. piperoides.

Scales present only on the lowest joint of each branch.
Leaves penninerved.

Leaves lanceolate, 1.5-2 cm. wide P. aurantiacum.

Leaves ovate, all or mostly 3-5 cm. wide.

Flower spikes 3-5 cm. long P. Heydeanum.

Flower spikes almost 10 cm. long P. Gentlei.

Leaves palmate-nerved, the nerves all arising from the base of the blade (often

concealed by the thick leaf tissue and difficult to distinguish).
Flowers all or mostly 2-ranked on each joint, conspicuously stipitate.

P. cheirocarpum.

Flowers chiefly or all in 4 or 6 ranks on each joint, usually sessile.
Fruit tuberculate, often very conspicuously so.

Leaves suborbicular or broadly obovate, mostly 1.5-3.5 cm. long,

broadly rounded or emarginate at the apex P. mucronatum.

Leaves lanceolate or narrowly lanceolate, mostly 8-16 cm. long, long-
attenuate to the apex P. annulatum.

68 FIELDIANA: BOTANY, VOLUME 24

Fruit smooth, not tuberculate.

Branches terete or nearly so, sometimes somewhat compressed and

2-edged, not 4-angulate.
Leaves thick-coriaceous and heavy when dried.

Stems densely puberulent P. Treleaseanum.

Stems glabrous.

Leaves oblong-oblanceolate, mostly 1-1.5 cm. wide.

P. Aguilarii.
Leaves oblong to lanceolate, mostly 2-4 cm. wide.

P. robustissimum.

Leaves only moderately coriaceous, not very thick and heavy.
Scales inserted above the base of the joint of the branch.

P. crispum.
Scales inserted at the base of the branch.

Leaves rounded or very obtuse at the apex.

Leaves small, about 4 cm. long and 2 cm. wide, usually acute

at the base P. Rondeletiae.

Leaves larger, about 5 cm. long and 3 cm. wide, mostly

rounded or obtuse at the base P. vulcanicum.

Leaves acute to long-acuminate, the tip often obtuse.

Leaves small, mostly 5.5-7.5 cm. long P. huehuetecum.

Leaves large, mostly 9-16 cm. long or even larger.

P. nervosum.

Branches all or mostly distinctly quadrangular, the old branches some-
times terete.
Leaves very small, about 3 cm. long and 5 mm. wide.

P. libertadanum.
Leaves usually much larger P. quadrangular e.

Phoradendron Aguilarii Standl. & Steyerm. Field Mus. Bot.
23:40. 1944. Liga. ,

On Quercus, and perhaps other hosts, 1,500-2,000 meters; ende-
mic; Zacapa; Jutiapa (type from Volcan de Suchitan, northwest of
Asuncion Mita, Steyermark 31889); Guatemala; Chimaltenango;
Quiche.

A densely branched, glabrous shrub, yellowish brown when dried, the branches
stout, terete, more or less dilated and compressed at the nodes, the cataphylls
basal only; leaves thick-coriaceous, on short thick petioles, oblong-oblanceolate,
4-8 cm. long, 1-1.5 cm. wide, broadest above the middle, narrowly rounded or
very obtuse at the apex, attenuate to the base, basinerved, the nerves inconspicu-
ous, not elevated, the costa obscure, percurrent; spikes fasciculate, subsessile, in
fruit scarcely 2 cm. long, the joints 3-4, thick, mostly 6-flowered, the flowers
4-seriate with 2 smaller ones above; scales of the spikes minutely ciliate; sepals
closely inflexed.

Phoradendron annulatum Oliver, Vid. Medd. Naturh.
For. Kjoebenhavn 1864: 176. 1865. P. multiflorum Trel. Gen.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 69

Phorad. 59. pis. 66, 67. 1916 (type from Volcan de Acatenango,
Sacatepe"quez, W. A. Kellerman 5154, 5155). Liga; Liga de pajaro.

At 1,200-2,400 meters; Alta Verapaz; Guatemala; Sacatepe*quez ;
Suchitepe"quez; Quezaltenango; San Marcos. Costa Rica.

Plants glabrous, often much branched and forming large masses, erect or
pendent, stout, the branches with only basal cataphylls, 2-edged or somewhat
angulate at first, becoming terete; leaves short-petiolate, rather thin, narrowly
lanceolate or linear-lanceolate, 10-18 cm. long, 1-4 cm. wide, long-attenuate to
the obtuse apex, acute or attenuate at the base, rather thin, palmate-nerved, the
nerves inconspicuous; flower spikes 3-4 cm. long, 3-4-jointed, mostly solitary, the
flowers chiefly 4-ranked, orange-yellow; fruit subglobose, reddish, 3-4 mm. in
diameter, the sepals closely inflexed.

This has been reported from Guatemala as P. rubrum Griseb.

Phoradendron aurantiacum Trel. in Standl. Field Mus. Bot.
17: 236. 1937. Matapalo; Kimiche (Maya).

Known only from the type, Sabana Zis, Lago de Pete*n, Pete"n,
C. L. Lundell 3191.

Branches pseudodichotomous, granulose, golden brown when dried, the cata-
phylls basal, the internodes rather short, terete, 3-4 cm. long; leaves lanceolate,
on petioles 1 cm. long or shorter, very obtuse, 4-5 cm. long, 1.5-2 cm. wide, opaque,
obscurely penninerved, acutely contracted at the base; spikes mostly solitary,
almost sessile, slender, at maturity 3 cm. long, the joints about 10, short, each
with 12 or fewer flowers, these mostly 4-seriate.

We have seen no material of this species.

Phoradendron cheirocarpum Trel. Gen. Phorad. 94. pi. 129.
1916.

At 350 meters or less; Alta Verapaz (type from Cubilgiiitz,
Tuerckheim 7661). Chiapas; British Honduras.

Plants slender, the branches with cataphylls only at the base, the internodes
elongate, the upper ones compressed at the nodes, the older ones terete; leaves
slender-petiolate, thin when dried, falcate-oblanceolate or sometimes lanceolate,
usually broadest toward the apex, 5-9 cm. long and 1.5 cm. wide, obtuse or nar-
rowly rounded at the apex, attenuate to the base; spikes fasciculate, mostly less
than 2 cm. long, the joints about 4, slender, 2-flowered, short-pedunculate, the
flowers conspicuously stipitate; fruit obovoid, 6 mm. long, smooth, the sepals
erect or spreading.

Phoradendron crassifolium (Pohl) Eichler in Mart. Fl. Bras.
5, pt. 2: 125. 1868. Viscum crassifolium Pohl ex DC. Prodr. 4: 280.
1830. P. crassifolium var. Pittieri Trel. Gen. Phorad. 145. pi. 215.
1916. Icvolay quen (Alta Verapaz); Matapalo.

70 FIELDIANA: BOTANY, VOLUME 24

At 450 meters or less; Alta Verapaz; Izabal. Costa Rica; south-
ward to Brazil.

A rather large, glabrous shrub, the branches stout, terete; leaves almost sessile,
very thick and hard, lance-ovate to broadly ovate, 8-16 cm. long, 3-10 cm. wide,
acute or acuminate with a usually obtuse tip, rounded to acute at the base, basi-
nerved, the nerves visible but not elevated; spikes solitary or fasciculate, 2-3 cm.
long, about 5-jointed, the joints 4-6-flowered, the principal flowers 4-ranked, with
2 smaller ones above; fruit yellowish, smooth, subglobose, 4 mm. in diameter, the
sepals closely inflexed.

Phoradendron crispum Trel. Gen. Phorad. 77. pi. 99. 1916.

At 1,700-2,400 meters; Zacapa; Chiquimula; San Marcos. Costa
Rica; Panama.

Usually a small shrub, glabrous, yellowish green when dried, the branches
stout, terete, the cataphylls a single pair, inserted above the base of the branch;
leaves on short or rather long petioles, rounded-obovate, mostly 3-5 cm. long and
1.5-2.5 cm. wide, rounded at the apex, abruptly or cuneately contracted at the
base, basinerved; spikes mostly solitary and 1.5 cm. long or shorter, the joints
usually 2-3, slender, the flowers 4-seriate; fruit small, smooth, white.

Phoradendron Gentlei Trel. in Standl. Field Mus. Bot. 12:
410. 1936.

Known only from the type, Corozal District, British Honduras,
P. H. Gentle 505.

Cataphylls basal, the internodes short and rather thick, obscurely somewhat
papillate, quadrangular; leaves short-petiolate, lustrous, elliptic or subobovate,
3-4 cm. long, 1-2 cm. wide, obtuse, cuneate at the base, minutely rugulose;
spikes solitary (?), short, the nodes about 3, few-flowered, the peduncle very short;
berries ellipsoid, apparently red, the sepals open.

We have seen no material of this species.

Phoradendron Heydeanum Trel. Gen. Phorad. 135. pi. 199.
1916.

Known in Guatemala only from the type, San Miguel Uspantan,
Quiche", 2,000 meters, Heyde & Lux 3140.

Plants with elongate branches, glabrous, the cataphylls basal only, compressed
and 2-edged, dilated at the nodes; leaves short-petiolate, lance-ovate, 10 cm. long
and 5 cm. wide or smaller, sometimes obovate and smaller, subobtuse, thick,
penninerved, the nerves very slender and inconspicuous; spikes often fasciculate,
3-5 cm. long, the joints 4-5, thick, somewhat turbinate, short-pedunculate, the
flowers 4-seriate, with 2 smaller flowers above the principal 4.

This has been reported from Guatemala as P. nervosum Oliver.

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 71

Phoradendron huehuetecum Stand!. & Steyerm. Field Mus.
Bot. 23:41. 1944.

Known only from the type, on Quercus, near Tachique, east of
Huehuetenango, Dept. Huehuetenango, 1,900 meters, Standley
82597.

A glabrous shrub, 30 cm. high or more, yellowish brown when dry, the branches
terete or subterete, rather slender, not thickened at the nodes, the cataphylls basal
only; leaves yellowish when fresh, only moderately coriaceous, on stout petioles
6 mm. long, lanceolate, sometimes somewhat falcate, mostly 4-7 cm. long and 1-1.5
cm. wide, gradually rather long-attenuate to the narrowly obtuse apex, attenuate
to the base, palmately 5-nerved, somewhat lustrous, the nerves very slender, evi-
dent and prominulous on both surfaces; young flower spikes solitary, stout, sessile,
1.5 cm. long, 2-3-jointed, few-flowered, the flowers mostly 4-seriate.

Phoradendron libertadanum Trel. in Standl. Field Mus. Bot.
17: 236. 1937. Matapalo.

Known only from the type, Pete"n, on Cochlospermum vitifolium,
La Libertad, C. L. Lundell 2401.

Plants glabrous, much branched, sometimes obscurely granulate, the cata-
phylls basal only; internodes of the branches 3-6 cm. long, 2-6 mm. thick, acutely
quadrangular, the upper ones ancipital; leaves on petioles 5 mm. long, oblong,
about 3 'cm. long and 5 mm. wide, mucronate-acute, cuneately narrowed at the
base, crispate, very obscurely basinerved.

Perhaps a form of P. quadr angular e, but apparently distinct in
its very reduced leaves. We know this species only from the original
description.

Phoradendron mucronatum (DC.) Krug & Urban, Bot.
Jahrb. 24: 34. 1897. Viscum mucronatum DC. Prodr. 4: 282. 1830.
P. yucatanum Trel. Gen. Phorad. 118. pi. 173. 1916 (type from
Yucatan).

At 500-600 meters; Jutiapa (between Asuncion Mita and Lago
de Giiija, Steyermark 31834). Yucatan Peninsula of Mexico; West
Indies; South America.

A stout, glabrous, often densely branched shrub, yellowish green when dried,
the branches usually sharply quadrangular, with basal cataphylls only, the inter-
nodes rather short; leaves on short thick petioles, moderately coriaceous, orbicular
to broadly obovate, mostly 1.5-3.5 cm. long and 1.5-2.5 cm. wide, broadly rounded
at the apex or often deeply emarginate, acute to rounded at the base, often abruptly
contracted, basinerved, the nerves slender, prominulous or often obscure; spikes
usually fasciculate, almost sessile, generally 1 cm. long or less, 3-4-jointed, the
joints 4-6-flowered, the flowers 4-ranked; scales of the spike ciliate; sepals erect;
fruit subglobose, orange, 3-4 mm. long, very densely and conspicuously verrucose.

72 FIELDIANA: BOTANY, VOLUME 24

Probably several species of the Aequitoriales-Emarginatae recog-
nized by Trelease are referable to the synonymy of this species. The
single Guatemalan collection is most like the species he recognized
as P. emarginatum Eichler, which is scarcely distinct from the com-
mon West Indian plant to which the name mucronatum was applied
originally.

Phoradendron nervosum Oliver, Vid. Medd. Naturh. For.
Kjoebenhavn 1864: 175. 1865; Trel. Gen. Phorad. pi. 74. 1916.
Liga; Sarapa (Quezaltenango).

On Quercus and probably other hosts, 1,200-3,000 meters; Alta
Verapaz; Zacapa; Guatemala; Sacatepe"quez; Chimaltenango; Solola;
Quiche" ; Huehuetenango; Quezaltenango; San Marcos. Southern
Mexico.

Plants glabrous, usually much branched, the branches erect or often pendent
and forming large dense masses, sometimes a meter long or more, with basal cata-
phylls only, at first compressed but in age terete, the plants often blackening
when dried; leaves short-petiolate, moderately coriaceous or often thin, obliquely
lanceolate, mostly 9-16 cm. long or even larger, usually long-attenuate to a narrowly
obtuse apex, acute or attenuate at the base, basinerved, the stout petioles 1 cm.
long or shorter, the nerves very slender, usually evident and often prominulous;
spikes mostly fasciculate, 2-6 cm. long, short-pedunculate, mostly 4-6-jointed, the
joints turbinate, the flowers 4-seriate, with often 2 smaller flowers above the princi-
pal 4, the scales ciliate; flowers greenish yellow; fruit brick-red, subglobose, 3 mm.
in diameter, minutely granular, the sepals inflexed.

From Mexico this plant is reported as occurring on Annona,
Liquidambar, and Quercus. It is one of the commonest species of
Guatemala.

Phoradendron piperoides (HBK.) Trel. Gen. Phorad. 145.
pis. 217-222. 1916. Viscum latifolium Swartz, Fl. Ind. Occ. 1: 268.
1797, not Lam. 1789. Loranthus piperoides HBK. Nov. Gen. & Sp.
3: 443. 1818. P. latifolium Griseb. Fl. Brit. W. Ind. 314. 1860. Liga.

At 1,400 meters or less; Alta Verapaz; Izabal; Escuintla; Guate-
mala; Solola; Quezaltenango; San Marcos. Mexico; British Hon-
duras to Salvador and Panama; West Indies; tropical South America.

A small or rather large shrub, erect or pendent, glabrous, often much branched
and forming dense masses, the branches stout or slender, terete or when young
slightly compressed, with cataphylls at the base of all the joints; leaves on short
thick petioles, moderately coriaceous, lanceolate to ovate, mostly 5-12 cm. long
and 1.5-7 cm. wide, acute or acuminate with an obtuse tip, acute at the base,
penninerved; spikes mostly fasciculate, 2.5-6 cm. long, about 6-jointed, the joints
rather slender, 10-15-flowered, the flowers mostly 4-ranked, yellowish green; fruit

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 73

yellow to orange or brown, ovoid or ellipsoid, smooth or somewhat granulate,
5 mm. long; sepals ascending, usually somewhat separated.

Called "matapalo" and "anteojos" in Salvador; "suelda con
suelda" (Honduras); "God Almighty" (British Honduras). The
Guatemalan hosts are not indicated but in Honduras the species
sometimes grows upon Ficus. Many Phoradendron species are not
confined to any one specific or generic host, while others are limited
in their occurrence.

Phoradendron quadrangulare (HBK.) Krug & Urban, Bot.
Jahrb. 24: 35. 1898. Loranthus quadrangularis HBK. Nov. Gen. &
Sp. 3: 444. 1818. P. Rensoni Trel. Gen. Phorad. 105. pi. 149. 1916
(type from San Salvador, Salvador). P. Gaumeri Trel. Gen. Phorad.
114. pi. 167. 1916 (type from Izabal, Yucatan). P. zacapanum Trel.
op. cit. 115. pi. 168. 1916 (type from Gualan, Zacapa, W. A. Keller-
man 5612). P. Millspaughii Trel. Bull. Torrey Club 54: 475. 1927
(type from Yucatan). P. belizense Trel. in Standl. Field Mus. Bot.
12: 409. 1936 (type from Belize, British Honduras, C. L. Lundell
1820). P. cayanum Trel. loc. cit. (type from El Cayo, British
Honduras, H. H. Bartlett 11997). P. cocquericotanum Trel. op. cit.
410 (type from Cocquericot, British Honduras, H. H. Bartlett
12073). P. manatense Trel. loc. cit. (type from Cornhouse Creek,
Manatee River, British Honduras, Bartlett 11304). P. franciscanum
Trel. in Standl. Field Mus. Bot. 17: 236. 1937 (type from Sabana
San Francisco, near La Libertad, Pete"n, C. L. Lundell 2398). P.
petenense Trel. op. cit. 237 (type from La Libertad, Pete"n, on Cura-
tella americana, Lundell 2400). Matapalo; Liga; Nigilita.

On various broad-leafed trees, 1,500 meters or less, chiefly below
1,000 meters; Pete"n; Alta Verapaz; Zacapa; Chiquimula; Jutiapa;
Santa Rosa; Escuintla; Suchitepe"quez; Retalhuleu; Quezaltenango;
San Marcos; Huehuetenango; doubtless in all the lower departments.
Southern Mexico; British Honduras to Salvador and Panama; West
Indies; South America.

A small glabrous shrub, usually much branched, erect or often pendent, mostly
50 cm. long or less, the branches usually slender and with elongate internodes,
quadrangular or the oldest ones subterete, with basal cataphylls only, these at or
near the base of the branch; leaves rather thin when dried or only moderately
coriaceous, dull or yellowish green, obovate to oblong-oblanceolate, broadest
toward the apex, mostly 4-7 cm. long and 1-2 cm. wide but variable in size and
shape, often slightly falcate, obtuse or rounded at the apex, attenuate to the base,
basinerved, the nerves very slender but usually evident and often prominulous;
spikes generally fasciculate, 3-4 cm. long, the joints generally 3-5, rather slender,
turbinate, the flowers pale yellow or yellow-green, mostly 4-ranked, the spikes

74 FIELDIANA: BOTANY, VOLUME 24

short-pedunculate; fruit brown to white or orange, subglobose, smooth or obscurely
papillate, 3 mm. in diameter; sepals usually closely inflexed.

Known in Salvador by the names "suelda con suelda," "cunegiie,"
and "sobrepalo." As this group of the genus (Aequitoriales-Quad-
rangulares) is treated by Trelease in The genus Phoradendron, it
contains 15 species, separated mainly by their geographic occurrence,
six of them being described from Mexico and Central America.
Since that publication the same author has described a rather large
number of additional species from the same area. It is obvious that
the species of this relationship have been fantastically multiplied and
that most of those recently published will have to be reduced to
synonymy. It is possible that the material now referred to P.
quadrangulare does represent more than a single species, but it is
not apparent where specific lines, if there are any, may be drawn.
It seems likely that all the names cited above, with a good many
more based on material from other regions, represent a single not
exceptionally variable species.

Phoradendron robust issimum Eichler in Mart. Fl. Bras. 5,
pt. 2: 122. 1868. P. robustissimum var. simulans Trel. Gen. Phorad.
78. pi. 102. 1916. P. falcifolium Trel. op. cit. 79. pi. 100. 1916 (type
from Santa Rosa, Baja Verapaz, Tuerckheim 11.2168).

On Quercus, Sapium, Dipholis, and probably other trees, 1,900
meters or less; Baja Verapaz; Jutiapa; Escuintla; Guatemala; Retal-
huleu; Huehuetenango. Campeche; British Honduras to Salvador
and Costa Rica.

Usually a rather large shrub, glabrous, the branches very stout, cellular-
papillate, compressed at first, in age terete, with only basal cataphylls, the inter-
nodes short or elongate; leaves on short thick petioles, very thick, coriaceous, and
stiff, oblong to elliptic-oblong or lance-ovate, mostly 5-12 cm. long and 2.5-5 cm.
wide, obtuse or rounded at the apex, acute to rounded at the base, basinerved
but the nerves usually obsolete; spikes mostly fasciculate and 3-5 cm. long, about
5-jointed, the joints about 16-flowered, on peduncles 3-5 mm. long, the flowers
yellowish green, 4-seriate; scales scarcely ciliate; fruit smooth, 5 mm. long; sepals
closely inflexed.

Called "matapalo" in Salvador; "suelda con suelda" (Honduras).
A large and showy plant with extremely thick and heavy leaves, the
foliage often tinged with brownish red.

Phoradendron Rondeletiae Trel. Gen. Phorad. 76. pi. 98.
1916.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 75

On Rondeletia, 1,300-1,450 meters; endemic; Alta Verapaz (type
from Coban, Tuerckheim 11.2045; collected also at Samac near
Coban).

Branches rather short, with only basal cataphylls, the internodes short, 1-3
cm. long, glabrous, somewhat compressed at first, dilated at the nodes, terete in
age; cataphylls inserted near the base of the branch; leaves on stout petioles 5 mm.
long or less, obovate or cuneate-obovate, 3-4 cm. long, 1-2 cm. wide, rounded at
the apex, cuneate at the base, thick, basinerved; spikes solitary, sessile or nearly
so, 1-1.5 cm. long, 2-3-jointed, the joints 4-10-flowered, the flowers 4-seriate;
sepals erect, spreading.

Phoradendron supravenulosum Trel. Gen. Phorad. 154.

W 9^9 1Q1fi

pi/, X/OiC. -LJ7-LU.

Known in Guatemala only from the vicinity of the type locality,
Cubilguitz, Alta Verapaz, 350 meters, Tuerckheim 8574. British
Honduras; Nicaragua; Costa Rica; Panama.

A rather large shrub, bright yellowish green when dry, glabrous, the branches
rather slender, with cataphylls on all the joints, the internodes granular, somewhat
hexagonal or subterete; cataphylls inserted 5-10 mm. above the nodes, deltoid and
pointed; leaves short-petiolate, rather thin or moderately coriaceous, broadly
lanceolate to ovate, mostly 9-14 cm. long and 3-6 cm. wide, acute or acuminate,
obtuse or acute at the base, palmately 5-nerved, the nerves slender but prominent
and very conspicuous on both surfaces, the veins also often elevated and conspicu-
ously reticulate; spikes mostly fasciculate, sessile, 3-7 cm. long, about 10-jointed,
the joints short, the flowers usually 6-seriate; fruit somewhat granular, the sepals
closely inflexed.

This has been reported from Guatemala as P. nervosum Oliver.

Phoradendron Treleaseanum Standl. & Steyerm. Field Mus.
Bot. 23:41. 1944.

Known only from the type, Dept. Baja Verapaz, Sierra de las
Minas opposite El Rancho (El Progreso), 700 meters, W. A. Keller-
man 7630.

A branched shrub, the branches stout, terete, densely and minutely puberu-
lent, the cataphylls basal only, subtruncate, puberulent; leaves sessile, narrowly
oblong, 3.5-5 cm. long, 1-1.5 cm. wide, very obtuse, shortly somewhat narrowed
at the base, the point of attachment of the base very broad, minutely puberulent
and granular, thick-coriaceous and rigid, slightly paler beneath, basinerved but
the nerves scarcely visible; spikes little more than 1.5 cm. long, very thick, sub-
sessile, fasciculate, densely puberulent, the joints 1-2 and 8-10-flowered, the
flowers 4-seriate; fruit globose-ovoid, 4 mm. long, very densely puberulent; sepals
open in fruit.

Phoradendron uspantanum Trel. Gen. Phorad. 53. pi. 61.
1916.

76 FIELDIANA: BOTANY, VOLUME 24

Known certainly only from the type, from San Miguel Uspantan,
Quiche1, 2,100 meters, Heyde & Lux 3141; probably conspecific is
Steyermark 47432 from Volcan de Atitlan, Suchitepe"quez, at 2,500
meters.

Branches rather long and stout, without cataphylls, the internodes rather
long, like the leaves sparsely hispidulous at first, glabrate in age, more or less com-
pressed, dilated at the nodes and as much as 13 mm. broad; leaves petiolate,
narrowly oblong-lanceolate or almost linear, about 15 cm. long and 1.5 cm. wide,
obtuse, the narrow basal portion 10-15 mm. long, the blades conspicuously and
palmately 5-nerved; spikes fasciculate, 2.5-4 cm. long, almost glabrous, the joints
3-5, the peduncle 2-4 mm. long; scales ciliate, glabrate.

This was reported once from Guatemala as P. angustifolium
Eichler.

Phoradendron velutinum Nutt. Journ. Acad. Phila. n. ser. 1:
185. 1847.

At 2,400 meters, on Prunus; Sacatepe"quez (Volcan de Agua,
W. A. Kellerman 4541). Mexico.

Plants rather large and stout, yellowish green when dried, the branches with-
out cataphylls, densely yellowish-pubescent like the leaves; leaves on petioles 1 cm.
long or shorter, rather thin or only moderately coriaceous, lanceolate or narrowly
lanceolate, often falcate, 7-17 cm. long, 1-4 cm. wide, long-attenuate to the acute
or obtuse apex, acute at the base, basinerved, the nerves slender but prominent
and conspicuous on both surfaces; spikes mostly fasciculate, 1.5-2 cm. long, villous,
the joints about 3, subglobose, the peduncle 3 mm. long; fruit subglobose, glabrous,
4 mm. in diameter; sepals separated.

This is reported from Mexico on Crataegus and Cornus.

Phoradendron vulcanicum Trel. Gen. Phorad. 77. pi. 99. 1916.

On Leguminosae (genera not recorded) and perhaps other hosts,
2,700-3,000 meters; Sacatepe"quez (type from Volcan de Acatenango,
W. A. Kellerman 4829; collected also on Volcan de Fuego); San
Marcos(?); endemic.

Plants glabrous, the cataphylls basal only, the branches somewhat compressed
or subterete, somewhat dilated below the nodes; leaves short-petiolate, elliptic or
oval, 4-6.5 cm. long, 2.5-3 cm. wide, rounded at the apex, rounded or obtuse at
the base, basinerved; spikes usually fasciculate, 1 cm. long or often longer, sub-
sessile, 2-3-jointed, the joints about 10-flowered; flowers 4-seriate, with 2 smaller
flowers above the principal ones.

PHTHIRUSA Martius

Parasitic shrubs, growing on dicotyledonous trees or shrubs, the stems often
elongate and pendent; leaves well developed, broad, opposite, coriaceous or car-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 77

nose; flowers small, usually perfect, solitary or in groups of 3, in terminal or axillary
spikes, racemes, or panicles, the bractlets connate into a small cupule; calyx limb
truncate or dentate; petals usually 6, free, spreading in anthesis; stamens alter-
nately unequal, the filaments fleshy, inserted on the petals below their middle;
ovary surrounded by an annular disk, the style stout, columnar; fruit a small
fleshy berry with viscid pulp; embryo straight.

Species about 45, in tropical America. One other Central Ameri-
can species is known from Panama.

Spikes few-flowered, scarcely longer than the petioles; leaves small, 2-4 cm. long,
very obtuse or rounded at the apex, conspicuously brown-marginate.

P. phaneroloma.

Spikes many-flowered, mostly equaling or longer than the leaves; leaves acute or
subacute, much larger, not brown-marginate P. pyrifolia.

Phthirusa phaneroloma Standl. Carnegie Inst. Wash. Publ.
461: 55. 1935. Struthanthus phaneroloma Lundell, Lloydia 2: 83.
1939.

Known only from the type, Sibun River, British Honduras,
P. H. Gentle 1426.

Plants branched, 35 cm. long or more, the older branches terete, glabrate,
the young ones densely ferruginous-furfuraceous, the internodes shorter than the
leaves; leaves on petioles 2-4 mm. long, bright green when dry and rigid, elliptic
or oblong-elliptic, 2-4 cm. long, 1.5-2.2 cm. wide, obtuse or narrowly rounded at
the apex, obtuse or subacute at the apex, glabrous above, densely furfuraceous
beneath on the costa when young, glabrate in age, obscurely 5-plinerved, the
margins densely ferruginous-furfuraceous; inflorescences axillary, 3-5-flowered,
short-pedunculate, scarcely longer than the petioles, glabrous, the flowers sessile;
berries oblong-cylindric, 5 mm. long, glabrous, subtruncate at the apex.

This species is referable to the genus Dendropemon, as the family
was divided by Urban, a group unknown otherwise in Central
America.

Phthirusa pyrifolia (HBK.) Eichler in Mart. Fl. Bras. 5, pt.
2: 63. 1868. Loranthus pyrifolius HBK. Nov. Gen. & Sp. 3: 441.
1818.

On broad-leafed trees, 350 meters or less; Alta Verapaz; Izabal.
British Honduras to Salvador and Panama; tropical South America.

Plants erect or often pendent, frequently much branched and forming dense
masses, the branches somewhat compressed or in age terete, when young fer-
ruginous-furfuraceous; leaves short-petiolate, ovate to elliptic, mostly 7-14 cm.
long and 3-6 cm. wide, acute or obtuse and usually short-cuspidate, obtuse or
rounded at the base, often decurrent, glabrous, penninerved; flower spikes rather
slender and remotely flowered, simple, ferruginous-furfuraceous, often longer
than the leaves, pedunculate; flowers brown or dark red, the perianth 1-1.5 mm.

78 FIELDIANA: BOTANY, VOLUME 24

long; berries oblong, spreading or reflexed, glaucous or glaucescent, 5-6 mm. long,
rounded at the apex.

Called "suelda con suelda" in Salvador and Honduras, and
probably also in Guatemala; "matapalo" (Salvador), a name given
to parasites or epiphytes of various families.

PSITTACANTHUS Martius

Parasitic shrubs, growing on broad-leafed trees; leaves all or mostly opposite,
well developed, with flat blades, usually thick-coriaceous when dried, very fleshy
when fresh, palmate-nerved or penninerved; flowers perfect, mostly 6-parted with
a 2-seriate perianth, very large and showy, usually red, racemose, corymbose, or
umbellate, in groups of 2-3, pedicellate, subtended by a cupular bractlet; calyx
usually urceolate, entire, crenate, or dentate; petals free or connate at the base
into a tube, spreading in anthesis; filaments filiform, partially united with the
petals, free and subulate above; anthers mostly versatile, elliptic to linear, 2-celled,
introrse, dehiscent by 2 longitudinal slits; ovary obovoid or subglobose, surrounded
by a usually annuliform disk, the style cylindric-filiform, generally 6-striate,
equaling the petals, often flexuous or geniculate, the stigma capitate or rarely
somewhat 2-lobate; berry fleshy, viscid; seed without endosperm; cotyledons
plano-convex.

About 50 species, in tropical America.

Corolla 6.5-8 cm. long, the segments almost filiform in anthesis. . . .P. Schiedeanus.
Corolla 3-5 cm. long, the segments linear in anthesis.

Corolla in bud conspicuously dilated near the apex, acute, conspicuously curved.

P. calyculatus.

Corolla in bud obtuse, not dilated near the apex, of equal breadth throughout,
straight or nearly so P. mayanus.

Psittacanthus calyculatus (DC.) G. Don, Gen. Syst. 3: 415.
1834. Loranthus calyculatus DC. Coll. M&m. pi. 10. 1830. Liga;
Liga dejocote; Anteojo;Gallito; Matapalo; Andilla (Huehuetenango) .

On broad-leafed trees, usually on Spondias purpurea, 1,500
meters or less; Pete*n; Zacapa; Chiquimula; Jutiapa; Santa Rosa;
Escuintla; Guatemala; Suchitepe"quez; Retalhuleu; Solola; Hue-
huetenango. Mexico; British Honduras to Salvador and Panama.

A small or rather large, parasitic shrub, usually erect, sparsely or much
branched, the branches very stout, quadrangular or compressed, the oldest ones j
subterete; leaves short-petiolate, coriaceous, short-petiolate, lanceolate and some-
what falcate to oblong or elliptic, 6-15 cm. long, attenuate to an acute apex or
often rounded or very obtuse; flowers very showy, red or orange-red, very numer-
ous, corymbose, long-pedicellate, the buds conspicuously outcurved, thickened
near the apex, acute; fruit black or purple-black, very juicy, oval, 1-1.5 cm. long.

Called "suelda con suelda" and "gallinago" in Honduras;
"chacxiu" (Yucatan, Maya). The plants of this genus are well

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 79

known in Central America because they produce the so-called flores
de palo, or are probably the principal source of them. These are
curious scars, somewhat resembling conventional rosettes of archi-
tectural decorations, left upon the woody host plant when the base
of the mistletoe plant is pulled away from it. These "wood flowers"
are often kept in houses for decorations, sometimes embellished with
gold and silver paint(!), and they occasionally are sold in tourist
shops. It is believed that some of them are produced by plants of
other genera of Loranthaceae, and they are said to be found often
on trees of Quercus, orange, Spondias, and other groups. In Guate-
mala P. calyculatus is often said to be confined to trees of Spondias
purpurea, and this is the most common host but certainly not the
only one. In British Honduras it is reported as occurring on Ficus.
The plants are very showy when in flower, but they often grow high
on the branches of tall trees, where they can be studied only from a
distance. The name "liga" is given in central Guatemala to all
plants of this family. The viscid fruits are employed as bird lime
or liga for catching sensontles and other birds that are kept in cages.
About Antigua it was stated that bird lime was prepared also from
Grevillea and avocado branches, the young twigs being chewed
thoroughly, buried in the ground for a few days, dug up and chewed
again, then applied to the branches of bushes or trees on which small
birds might alight. P. calyculatus has been confused in recent years
with P. americanus (L.) Mart., a species probably confined to the
Lesser Antilles and northern South America. The species of this
group are closely related, and the differences between them none too
well marked, or perhaps only misunderstood.

Psittacanthus mayanus Standl. & Steyerm. Field Mus. Bot.
23:41. 1944.

Type from Santa Rita, British Honduras, growing on Bursera
Simaruba, Percy Gentle 116. Southern Mexico; Honduras.

A glabrous branched shrub 30 cm. high and larger, the branches stout, more
or less compressed and rather acutely quadrangular, the older ones ochraceous,
subterete; leaves opposite or the uppermost subopposite, on short thick petioles,
coriaceous when dry, falcate-lanceolate to oblong or oblong-elliptic, 4.5-7 cm.
long, 1-3 cm. wide, attenuate to an acute apex or more often obtuse or narrowly
rounded, acute or attenuate at the base, 3-5-plinerved, the nerves prominent on
both surfaces; flowers red, corymbose, the corymbs mostly dense and many-
flowered, rarely lax and few-flowered, the pedicels ternate, umbellate; bractlets
cupular, 1.5 mm. long; calyx campanulate subtruncate 3 mm. long and broad;
corolla 3-5 cm. long, in bud linear, almost straight, of uniform length throughout,
obtuse, glabrous, the petals in anthesis almost filiform, revolute; anthers 2-2.5
mm. long; berries oval, 6 mm. long, capped with the persistent calyx.

80 FIELDIANA: BOTANY, VOLUME 24

There is referred here with much doubt a British Honduran
collection said to have been taken from a pine tree. One would
expect this to represent a distinct species, but there are no obvious
characters for separating it, especially since the material is in poor
condition for study.

Psittacanthus Schiedeanus (Schlecht. & Cham.) Blume ex
Schult. Syst. Veg. 7: 1730. 1830. Loranthus Schiedeanus Schlecht. &
Cham. Linnaea 5: 172. 1830.

On Pinus and perhaps other genera of trees, 1,700 meters or less;
Chiquimula; reported from Suchitepe"quez and Sacatepe'quez.
Southern Mexico; Salvador; Costa Rica; Panama.

A stout stiff shrub, usually erect, the branches thick, acutely quadrangular or
the older ones terete, greenish or ochraceous, glabrous throughout; leaves short-
petiolate, narrowly falcate-lanceolate to ovate, 6-16 cm. long, usually very asym-
metric, attenuate to an obtuse apex or merely obtuse, attenuate to obtuse at the
base, very thick; flowers numerous, in dense corymbs, orange-red, 6.5-8.5 cm.
long; corolla in bud linear, little dilated at the apex, almost straight, obtuse; fruit
black at maturity, oval, 2 cm. long or shorter.

Called "matapalo" in Salvador.

STRUTHANTHUS Martius

Shrubs, usually glabrous, growing upon dicotyledonous trees or shrubs, erect
or often scandent or pendent, sometimes with twining stems, the stems terete or
quadrangular; leaves opposite or mostly so, with well developed blades, usually
rather thinly coriaceous, penninerved; flowers small, green or yellow, commonly
dioecious and 6-parted, ternate, the groups of flowers racemose, corymbose, or
pseudocymose, sometimes paniculate, sometimes in axillary glomerules, accom-
panied by bracts and bractlets; calyx small, entire or obsoletely dentate; petals
free; stamens unequal, alternately long and short, the filaments filiform-subulate;
anthers versatile, elliptic or cordate, 2-celled, dehiscent by longitudinal slits;
ovary obovoid or depressed-globose, surrounded by a fleshy disk; style cylindric,
usually equaling the petals, the stigma discoid-capitate, papillose; fruit a small
berry with viscid pulp.

Species perhaps 50, in tropical America. A few others are known
from southern Central America.

Leaves all or chiefly rounded, retuse, or very obtuse at the apex, mostly orbicular,

obovate, oblanceolate, or obovate-elliptic.
Leaves orbicular or nearly so, abruptly contracted into the petiole and mostly

rounded or very obtuse at the base of the blade S. orbicularis.

Leaves mostly obovate or oblanceolate-oblong, broadest above or rarely at the

middle, long-attenuate to the base.

Inflorescences all or mostly 3-flowered; branches relatively short, stout, not
at all flexuous S. oliganthus.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 81

Inflorescences mostly several-many-flowered; branches long and slender,

flexuous, generally more or less twining or scandent S. cassythoides.

Leaves all or mostly acute or acuminate, sometimes attenuate to an obtuse apex,

all or nearly all of them broadest at or below the middle.
Inflorescences head-like, sessile in the leaf axils, scarcely or not at all longer

than the petioles S. Johnstonii.

Inflorescences not head-like, often pedunculate, much longer than the petioles.
Leaves thick-coriaceous when mature, not or scarcely blackening when dried;

branches usually not emitting aerial roots S. tacanensis.

Leaves thin, blackening when dried; branches usually emitting numerous

thick aerial roots.
Leaves abruptly acute or acuminate, with an acute tip.

Branches of the inflorescence and mature calyx densely and minutely

whitish-tuberculate S. papillosus.

Branches of the inflorescence and mature calyx smooth, not tuberculate.

S. marginatus.

Leaves merely acute or subacute, rarely obtuse, never abruptly acute or
acuminate, often attenuate to an obtuse tip, the tip rarely if ever acute.

Leaf blades mostly elliptic or broadly ovate and 2.5-5 cm. long.

S. Matudai.

Leaf blades various in shape, mostly lanceolate or lance-oblong or ovate-
oblong, mostly 5-10 cm. long.

Leaves sessile or nearly so, the petiole marginate to the base; inflores-
cence short and few-flowered, usually about as long as broad.

S. brachybotrys.

Leaves conspicuously petiolate, the petiole often 1 cm. long; inflores-
cence generally elongate and many-flowered.

Flowers slender-pedicellate S. tenuifolius.

Flowers sessile.

Leaves lanceolate or linear-lanceolate, very acute or attenuate at

the base S. Haenkei.

Leaves lance-oblong or ovate-oblong, abruptly contracted at the
base and commonly rounded or very obtuse . . . S. tacanensis.

Struthanthus brachybotrys Standl. & Steyerm. Field Mus.
Bot. 23: 42. 1944.

Parasitic on Quercus and "Acacia," 1,200-1,800 meters; endemic;
Guatemala (Lago de Amatitlan); Huehuetenango (type from Rio
Pucal, about 14 km. south of Huehuetenango, Standley 82420).

An erect or pendent shrub, the branches straight, not emitting aerial roots,
terete, striate, ochraceous or grayish, the internodes short; leaves sessile or sub-
sessile, thin-coriaceous, when dry pale brownish or sometimes fuscous, lance-oblong,
ovate-oblong, or oblong-elliptic, broadest usually at the middle, 4-7.5 cm. long,
1.2-3 cm. wide, acute or subobtuse, cuneately narrowed at the base, the lateral
nerves slender, prominent on both surfaces or sometimes obsolete beneath, ascend-
ing at a narrow angle; inflorescences solitary, 1.5-2 cm. long (including a peduncle
7-8 mm. long), almost head-like, densely few-flowered, the groups of flowers
(ternations) almost sessile, their peduncles very short and thick, the flowers green,

82 FIELDIANA: BOTANY, VOLUME 24

sessile, crowded; calyx little more than 1 mm. broad, subtruncate, smooth; corolla
clavate-obovate in bud, gradually dilated upward, 4 mm. long, the tube very thick;
fruit ellipsoid, orange-colored, 6-8 mm. long, rounded at the base and apex.

It is possible that this is S. Oerstedii (Oliver) Standl., described
from Granada, Nicaragua, and supposed to occur in Costa Rica.
Of that we have seen no authentic material and the too brief original
description does not agree satisfactorily with the Guatemalan plant.

Struthanthus cassythoides Millsp. ex Standl. Field Mus.
Bot. 8: 7. 1930 (type from Progreso, Yucatan). S. Gentlei Lundell,
Contr. Univ. Mich. Herb. 6: 7. 1941 (type from Stann Creek,
British Honduras, P. H. Gentle 2660). Matapalo.

On Byrsonima, Conocarpus, and doubtless other genera of shrubs
and trees, 300 meters or less; Pete"n; Alta Verapaz; Izabal; Santa
Rosa. Yucatan Peninsula of Mexico; British Honduras.

A glabrous parasite, usually pendent or scandent, the stems rarely as much as
6 meters long, slender, glaucous-green, terete, the internodes short or elongate;
leaves mostly on short thick petioles 3-4 mm. long or less, obovate to narrowly
obovate-oblong, mostly 2.5-5.5 cm. long and 1-2.5 cm. wide, very variable in
shape and size, rounded at the apex, cuneate or cuneate-attenuate at the base,
grayish or fuscous when dried, moderately coriaceous, the lateral nerves few,
ascending at a very narrow angle, conspicuous or often obsolete; inflorescences
solitary or fasciculate, on stout peduncles 2-5 mm. long, mostly 3-10-flowered
and short but not very dense, sometimes more elongate, the ternations pedunculate,
the flowers sessile, yellowish green; calyx truncate; corolla in bud almost linear,
slightly dilated near the apex, smooth, the linear petals 3-4 mm. long; filaments
stout, equaling the petals; style thick, straight, equaling the petals; fruit ellipsoid,
reddish green, about 7 mm. long.

The type of S. Gentlei is a form with unusually large and broad
leaves. At first glance it appears distinct from the typical form with
relatively small and narrow leaves, but there are so many apparently
intergrading forms that it is not practical to recognize here two
species, unless further collections should reveal distinctive characters
not now apparent.

Struthanthus Haenkei (Presl) Engler, Nat. Pflanzenfam.
Nachtr. 1: 134. 1897. Spirostylis Haenkei Presl ex Schult. Syst.
Veg. 7: 163. 1829. Matapalo; Suelda con suelda.

OnQuercus, Pinus, and probably other hosts, 1,000-1,800 meters;
Baja Verapaz; El Progreso; Zacapa; Chiquimula; Jalapa. Southern
and western Mexico.

A glabrous shrub, the branches straight or nearly so, usually not emitting aerial
roots, grayish or ferruginous, slender or rather stout, often elongate and pendent;

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 83

leaves usually thick-coriaceous, on rather slender petioles as much as 1 cm. long,
lanceolate or linear-lanceolate, mostly 7-12 cm. long and 1.2-3 cm. wide, long-
attenuate to a narrow obtuse apex, acute at the base, grayish green when dried,
paler beneath, rather conspicuously 3-nerved and also penninerved above the base,
the nerves slender but often prominulous; inflorescences solitary or fasciculate,
elongate and few-many-flowered, 7 cm. long or less, much interrupted, the terna-
tions short-pedunculate, the flowers closely sessile, the bracts rather large and con-
spicuous in the young inflorescence but soon deciduous; fruit oblong-ovoid or
ellipsoid, 5-6 mm. long, probably black at maturity.

This is perhaps the plant reported by Loesener as Struthanthus
spirostylis, growing on Juniper us, from Huehuetenango (Seler 3064).

Struthanthus Johnstonii Standl. & Steyerm. Field Mus. Bot.
23:43. 1944. Matapalo.

OnQuercus and perhaps other hosts, 1,350-2,300 meters; endemic;
Huehuetenango (type collected along the road between Aguacatan
and Huehuetenango, at km. 12, John R. Johnston 1887).

Plants glabrous, erect or pendent, the branches stout, not emitting aerial
roots, subterete, ferruginous, the internodes shorter than the leaves; leaves on short
stout petioles 7 mm. long or less, ovate, oblong-ovate, or elliptic-ovate, mostly
5-9 cm. long and 2.5-4 cm. wide, acute or short-acuminate with an acute tip,
abruptly contracted at the base and obtuse or almost rounded, blackish green
when dry, more or less rugulose above, paler beneath and densely and minutely
granular; pistillate inflorescences sessile, few-flowered, head-like, the flowers
closely sessile; calyx glaucescent, truncate, 2 mm. broad, smooth; fruit oblong or
ellipsoid, 7-12 mm. long, 4-6 mm. thick, closely sessile, broadly rounded or sub-
truncate at the apex.

Struthanthus marginatus (Desr.) Blume ex Schult. Syst.
Veg. 7: 1731. 1830. Loranthus marginatus Desr. in Lam. Encycl. 3:
596. 1791. Liga; Anteojos.

Parasitic on various trees or large shrubs, often on Coffea,
400-2,400 meters, mostly at 1,200-1,500 meters; Alta Verapaz; El
Progreso; Chiquimula; Jalapa; Jutiapa; Santa Rosa; Guatemala;
Sacatepe"quez; Chimaltenango; Solola; Quiche"; Huehuetenango;
Suchitepe"quez; Quezaltenango; San Marcos. Chiapas; Salvador to
Panama; South America.

A glabrous shrub, often glaucous green, usually darkening when dried, the
branches generally long and pendent, often twining and scandent, frequently
emitting conspicuous aerial roots; leaves on short slender petioles, generally thin
when dried, broadly ovate to lanceolate, mostly 6-11 cm. long, rather abruptly
acute or acuminate with an acute tip, abruptly contracted and broadly rounded
to obtuse at the base, penninerved, the nerves usually conspicuous and promi-
nulous, very slender, the veins often evident and closely reticulate; inflorescences
solitary or fasciculate, racemose, usually much shorter than the leaves, slender,

84 FIELDIANA: BOTANY, VOLUME 24

interrupted or rarely dense, the ternations pedunculate, the flowers sessile or very
shortly pedicellate, greenish yellow or green; corolla about 3 mm. long; fruits oval
or ellipsoid, red or brown.

Called "matapalo" in Salvador and doubtless also in Guatemala.
This is one of the species that often infests coffee bushes.

Struthanthus Matudai Lundell, Lloydia 4: 45. 1941.

At 2,500-3,000 meters; San Marcos (northeastern slopes of
Volcan de Tacana, Steyermark 36216). Chiapas, the type from
Cerro Ovando.

A small glabrous shrub, the branches stout, terete, ferruginous, the young ones
often angulate, not emitting aerial roots, the internodes short; leaves short-
petiolate, thin-coriaceous, lance-oblong to ovate or ovate-elliptic, mostly 2-4 cm.
long and 1-2.7 cm. wide, usually acute, with an acute or sometimes obtuse tip,
acute at the base or sometimes rounded and abruptly contracted, the nerves
obsolete or nearly so; inflorescences mostly fasciculate, 2 cm. long or less, densely
few-flowered, short-pedunculate, sometimes head-like, the ternations sessile or
short-pedunculate, the flowers sessile, 6 mm. long or less; calyx truncate; petals
linear, 5 mm. long or shorter; style contorted, 4 mm. long.

Struthanthus oliganthus Standl. & Steyerm. Field Mus.
Bot. 23: 154. 1944. Liga.

At 1,350-2,300 meters; endemic; Huehuetenango (type collected
above San Ildefonso Ixtahuacan, Steyermark 50672; also on Cerro
Chiquihui, northwest of Cuilco).

A small glabrous branched shrub, the branches stout, terete, not at all flexuous
or twining, when young ochraceous or pale brown;. leaves small, coriaceous, yel-
lowish when dried, borne on short stout petioles, obovate-oblong or broadly
cuneate-oblong, about 2.5 cm. long, 9-14 mm. wide, rounded at the apex, cuneately
narrowed below and decurrent almost to the base of the petiole, penninerved, but
the lateral nerves obscure; inflorescences very small, axillary, on stout peduncles
scarcely more than 3 mm. long, 3-flowered, the flowers greenish, sessile; calyx
short, 1.2 mm. broad; corolla in bud clavate-cylindric, 3.5 mm. long.

Struthanthus orbicularis (HBK.) Blume ex Schult. Syst. Veg.
7: 1731. 1830. Loranthus orbicularis HBK. Nov. Gen. & Sp. 3: 434.
1818. S. belizensis Lundell, Lloydia 2: 81. pi. 2. 1939 (type from
Valentin, El Cayo District, British Honduras, C. L. Lundell 6973).
S. escuintlensis Lundell, Phytologia 2: 1. 1941 (type from Escuintla,
Chiapas). Liga; Matapalo; Bejuco secapalo (Pete"n); Liga cazadora;
Liga de cortina.

On many groups of trees and shrubs, chiefly or wholly at 1,100
meters or less; Pete'n; Alta Verapaz; Izabal; Jutiapa; Santa Rosa;
Escuintla; Guatemala; Suchitepe'quez; Retalhuleu; Suchitepe'quez;

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 85

Solola; San Marcos. Chiapas; British Honduras to Salvador and
Panama; South America.

A glabrous parasite, glaucous-green, the branches terete, or when young
angulate or compressed, long and slender, often greatly elongate and twining or
scandent, usually emitting numerous coarse aerial roots, the internodes generally
much elongate; leaves slender-petiolate, only moderately coriaceous when dry,
fleshy when fresh, mostly orbicular or nearly so, varying to rounded-obovate,
chiefly 4-7 cm. long, broadly rounded at the apex, often conspicuously mucronate,
abruptly contracted at the base and rounded or obtuse, the costa prominent
beneath, the nerves inconspicuous or obsolete; inflorescences sessile or short-
pedunculate, much interrupted, slender, mostly many-flowered, generally longer
than the leaves but sometimes shorter, the ternations short-pedunculate or sessile,
the flowers sessile, green or whitish; corolla 6 mm. long or shorter; fruit oval, red
at maturity, 1 cm. long or shorter.

Sometimes called "hierba de rosario" in Salvador. This is the
most luxuriant in growth of all the Loranthaceae of Guatemala.
It often is a large vine, completely covering with its festoons of
branches large shrubs or even small trees, so that little of the proper
foliage of the host may be seen. In Alta Verapaz it is particularly
abundant in abandoned or neglected coffee plantations, covering
the bushes and extending from one to another. It is needless to say
that when it occurs in such abundance it soon kills the hosts, and in
well-tended cafetales this and other members of the family are
removed regularly from the bushes.

Struthanthus papillosus Standl. & Steyerm. Field Mus. Bot.
23: 43. 1944. Matapalo.

Parasitic on Erythrina (the type) and other hosts, 1,200-1,600
meters; endemic; Alta Verapaz (type from Coban, Tuerckheim
11.1240); Baja Verapaz; Guatemala(?).

A pendent glabrous shrub, the branches slender, often much elongate, terete,
ferruginous or grayish, generally emitting coarse aerial roots, the internodes
elongate; leaves thin and subcoriaceous, generally blackening when dried, on
slender petioles 5-10 mm. long, lanceolate to rather broadly ovate or oblong-ovate,
mostly 6-8 cm. long and 2-3.5 cm. wide, abruptly acute or rather long-acuminate,
with an acute tip, abruptly contracted at the base and rounded or obtuse, penni-
nerved, somewhat paler beneath, the very slender nerves often conspicuous, very
slender, the veins often conspicuous beneath and closely reticulate; inflorescences
axillary, solitary or more often fasciculate, mostly 6.5 cm. long or shorter, equal-
ing or shorter than the leaves, short-pedunculate, slender, much interrupted, the
branches minutely whitish-papillose, the ternations slender-pedunculate, the
peduncles about 3 mm. long, the flowers sessile, green; calyx little more than 1 mm.
broad, densely and minutely but conspicuously whitish-papillose; corolla slender-
cylindric in bud, not or scarcely dilated at the apex, 4 mm. long, the petals linear;
stamens about equaling the petals.

86 FIELDIANA: BOTANY, VOLUME 24

It is questionable whether this is a distinct species, but the
papillosity of the inflorescence is sometimes conspicuous, particularly
in the type collection. It remains to be determined whether this is
a good specific character.

Struthanthus tacanensis Lundell, Lloydia 4: 46. 1941.

Parasitic onQuercus and perhaps other hosts, 2,500-2,900 meters;
Quezaltenango. Chiapas, the type from Chiquihuite, Volcan de
Tacana, E. Matuda 2840.

A large coarse shrub, the branches stout, terete, ferruginous and furfuraceous,
not emitting aerial roots, the nodes often conspicuously enlarged, the internodes
short; leaves on thick petioles 8 mm. long or less, thick-coriaceous, oblong-lanceo-
late or ovate-oblong, 5-12 cm. long, 2-5 cm. wide, attenuate to a narrow obtuse
apex, abruptly contracted and obtuse or rounded at the base, drying dark yellowish
green or fuscescent, the lateral nerves slender, evident or almost obsolete; inflores-
cences 5 cm. long or shorter, sessile or short-pedunculate, often densely fasciculate,
interrupted or rather dense, the ternations on very short, thick peduncles or sub-
sessile, the flowers closely sessile, 10 mm. long or shorter; calyx truncate, obscurely
denticulate; petals linear, as much as 9 mm. long; style contorted, 7 mm. long;
fruit ovoid or ellipsoid, 7-8 mm. long.

Struthanthus tenuifolius Standl. & Steyerm. Field Mus.
Bot. 23: 155. 1944.

Known only from the type, Huehuetenango, Cie*naga de Lagar-
tero, 300 meters, parasitic on Taxodium mucronatum, Steyermark
51538.

A slender glabrous pendent shrub, laxly branched, the branches terete, appar-
ently not emitting aerial roots; leaves coriaceous, on slender petioles about 4 mm.
long, linear-lanceolate, 3.5-6.5 cm. long, 3-5 mm. wide, gradually attenuate to an
acute or subacuminate apex, long-attenuate to the base, 1-nerved; inflorescences
axillary and terminal, rather lax and open, about 2 cm. long and broad, few-
flowered, subcymose; flowers ternate, on stout pedicels 2-3 mm. long; calyx short,
truncate, slightly more than 1 mm. broad.

OPILIAGEAE

Shrubs or trees; leaves alternate, entire; stipules usually none; flowers small,
white or greenish, regular, perfect, spicate, racemose, or umbellate; calyx entire
or obscurely 4-5-dentate; petals 4-5, free; stamens as many as the petals and
opposite them, free or adnate at the base; disk present; ovary superior or nearly
so, 1-celled, with a thick central placenta; ovule 1, pendulous from the apex of the
placenta; style simple; fruit fleshy; seed without a testa, the endosperm copious;
embryo large, the radicle superior.

About 5 genera, in the tropics of both hemispheres, with only a
few species. Only the following genus reaches North America.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 87

AGONANDRA Miers

Reference: Paul C. Standley, The North American species of
Agonandra, Journ. Wash. Acad. Sci. 10: 505-508. 1920.

Shrubs or trees, glabrous or sparsely pubescent, the branches often slender
and pendulous; leaves thin, stipulate, short-petiolate, the lateral nerves usually
obscure; flowers very small, whitish or greenish, in bracteate axillary racemes,
usually dioecious; calyx minute, cupular, 4-lobate; petals 4 in the staminate flower,
villosulous outside; stamens 4, exserted, alternating with the same number of
glands, the filaments filiform; anthers ovate, suberect; petals and stamens none
in the pistillate flowers; ovary sessile, glabrous, the stigma sessile, discoid; fruit
fleshy, drupaceous.

About 6 species, in tropical America. There may be one or two
additional Central American species in southern Central America.

Agonandra racemosa (DC.) Standl. Journ. Wash. Acad. Sci.
10: 506. 1920. Schaefferia racemosa DC. Prodr. 2: 41. 1825.

Moist or wet forest, 1,300 meters or less; Izabal; Jutiapa; Retal-
huleu; Quiche". Mexico; Salvador; perhaps extending southward
into South America.

Usually a tree of 4-9 meters, glabrous throughout, the branches very slender,
green when young; leaves thin, on petioles 4-9 mm. long, lanceolate to ovate or
broadly elliptic-ovate, sometimes rounded, mostly 4-8 cm. long and 1-4.5 cm.
wide, usually acute to long-acuminate, often abruptly so, cuneate to broadly
rounded at the base, papillate beneath when dry, the lateral nerves scarcely
perceptible; racemes longer or shorter than the leaves, the flowers pedicellate;
bracts acute or acuminate, covering the buds but caducous in anthesis; petals
2.5 mm. long; fruit subglobose, about 8 mm. long.

Probably some of the South American species will have to be
reduced to the synonymy of A. racemosa, giving the species a wide
range. There are at least 2 and probably 3 good species of the genus
in Mexico. The wood is of good quality in this genus, but seldom
procurable in sizes large enough to be of importance. The heart-
wood is orange-yellow, the sap wood pale yellow; very hard, heavy,
compact, and strong, fine-textured, usually straight-grained, finishes
very smoothly. The senior author once saw some trees probably
of this genus and species in the lowland forest of the Atlantic coast
of Honduras. The trunk was about 9 meters high, simple, about
25 cm. in diameter at the base and tapering very gradually to a long
and slender tip like a buggy whip. The crown consisted of only a
few weak, more or less pendent branches. The habit was some-
what suggestive of the curious genus Idria (Fouquieriaceae) found
in Baja California, although of course the two are not related and
grow under very different conditions.

88 FIELDIANA: BOTANY, VOLUME 24

OLACACEAE

Trees or shrubs; leaves usually alternate and entire, penninerved, without
stipules; inflorescence usually axillary and few-flowered, the flowers solitary,
fasciculate, cymose, or racemose, small, greenish or white, regular, perfect or
unisexual; calyx small, with 4-6 teeth or lobes, sometimes greatly enlarged in
fruit; petals 4-6, free or more or less united, valvate or subimbricate; stamens
4-12, inserted with the petals and more or less adnate to them, all fertile or part
of them sterile, the filaments free or rarely monadelphous; anthers 2-celled; disk
various; ovary free, 1-celled or imperfectly 3-5-celled; ovules usually 2-3; fruit
drupaceous, commonly 1-celled and 1-seeded.

About 25 genera, widely dispersed in tropical regions. Two
other genera, Chaunochiton and Minquartia, are known from Costa
Rica and Panama.

Corolla lobes densely barbate within; plants armed with spines Ximenia.

Corolla lobes not barbate; plants unarmed.

Stamens twice as many as the corolla lobes; calyx accrescent in age and saucer-
shaped, bright red; flowers fasciculate in the leaf axils Heisteria.

Stamens as many as the corolla lobes; calyx not accrescent, small, green;
flowers in small racemes Schoepfia.

HEISTERIA Jacquin

Glabrous trees or shrubs; leaves membranaceous or coriaceous, short-petiolate,
entire; flowers very small, short-pedicellate or sessile, fasciculate in the leaf axils;
calyx minute, 5-6-dentate or 5-6-lobate, persistent and greatly enlarged in fruit,
erect and enclosing the fruit or often reflexed and exposing it, usually bright red
or purple, subentire to deeply lobate, often rotate and orbicular; petals small,
more or less villous within; stamens usually 10-12, rarely 5-6, hypogynous or
adnate at the base to the petals; ovary depressed-globose, 3-celled; fruit drupa-
ceous, globose to oblong, often black, the flesh thin, the endocarp crustaceous.

Species about 50, mostly in tropical America, a few in west
Africa. Three or four other species are found in southern Central
America. The Guatemalan species are easily recognized when in
fruit by the deep red, circular calyx in whose middle is seated the
small black drupe. The wood in this genus is moderately to decid-
edly heavy, hard, and strong, usually fine- textured. The trees are
too small to be of commercial importance, and no use is known to be
made of the wood locally.

Fruiting calyx deeply lobate, more or less enclosing the fruit; drupe about 14 mm.

thick; petioles mostly 1 cm. long or longer H. media.

Fruiting calyx subentire or shallowly lobate, rotate, not enclosing the fruit; drupes

about 8 mm. thick; petioles mostly 6-7 mm. long H. macrophylla.

Heisteria macrophylla Oerst. Vid. Medd. Kjoebenhavn 1856:
40. 1857 (type from San Juan del Norte, Nicaragua). Arito de
montana (Quezaltenango) ; Palo de baston (Quezaltenango).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 89

Mostly in dense, moist or wet, mixed forest, 240-2,700 meters;
Escuintla; Suchitepe"quez ; reported from Sacatepe"quez ; Chimalte-
nango; Solola; Quezaltenango ; San Marcos. Salvador and Honduras
to Panama.

A shrub or small tree 1.5-6 meters high, the branches slender, green, angulate at
first; leaves short-petiolate, oblong-lanceolate or lanceolate, 9-20 cm. long, acumi-
nate or long-acuminate, obtuse or rounded at the base, membranaceous or char-
taceous, bright green; pedicels solitary or fasciculate in the leaf axils, mostly 4-6
mm. long; fruiting calyx bright deep -red, about 2 cm. broad, very shallowly
5-lobate with rounded lobes or subentire, widely spreading or even reflexed in
fruit; stamens 10; fruit subglobose, black, 8-10 mm. long.

A frequent and rather conspicuous (when in fruit) shrub of the
understory in the mountain forests of the Pacific bocacosta. This
species has been reported from Guatemala as H. acuminata (Humb.
& Bonpl.) Benth. & Hook., a Colombian species. The shrub is
sometimes known in Salvador by the name "sombrerito."

Heisteria media Blake, Contr. U. S. Nat. Herb. 24:*3. 1922.
H. Chippiana Standl. Field Mus. Bot. 11: 130. 1932 (type from 19
Mile, Stann Creek Valley, British Honduras, W. A. Schipp 970).

Moist or wet forest, 800 meters or less; Pete*n; Alta Verapaz.
Chiapas; British Honduras; Honduras (type from Los Ranches,
Dept. Copan).

A shrub or tree as much as 15 meters high with a trunk diameter of 45 cm.,
the branches slender, terete or somewhat angulate; leaves subcoriaceous, lustrous,
the petioles mostly 10-15 mm. long; leaf blades lance-oblong to oblong-elliptic,
10-15 cm. long, gradually or abruptly and shortly obtuse-acuminate, acute at the
base; flowers usually densely fasciculate in the leaf axils, subsessile or on short
thick pedicels; fruiting calyx 3-4 cm. broad, ascending and involving the fruit,
at first green, turning purple-red, lobate to about the middle, the lobes broadly
rounded; fruit ochroleucous, subglobose, about 1.5 cm. long, rounded at each end.

Known in British Honduras by the names "copalche" macho,"
"nance cimarron," and "wild cinnamon"; "pate macho" (Honduras).

SCHOEPFIA Schreber

Glabrous shrubs or small trees; leaves usually coriaceous; flowers small and
inconspicuous, in few-flowered racemes, these axillary, solitary or fasciculate;
calyx small, cyathiform, obscurely denticulate, unchanged in fruit; disk entire,
adnate to the ovary; petals 4-6, inserted on the margin of the disk, coalescent to
form a tubular-campanulate corolla, the segments valvate in bud; stamens as
many as the corolla segments, adnate to the corolla, the anthers small, dorsifixed;
ovary semi-immersed in the disk, imperfectly 3-celled, the style short or elongate,
the stigma 3-lobate; ovules 3, pendulous from the apex of the placenta; fruit
drupaceous, annulate near the apex, the stone crustaceous or chartaceous; seed
falsely erect, the embryo minute, the endosperm carnose.

90 FIELDIANA: BOTANY, VOLUME 24

About 10 species, in the tropics of America and Asia. Only the
following species are known from Central America but four others
occur in Mexico.

Flowers 4-5 mm. long; corolla lobes fully half as long as the tube; leaves with
usually 3-4 pairs of lateral nerves S. Schreberi.

Flowers about 8 mm. long; corolla lobes scarcely one-third as long as the tube;
leaves with about 6 pairs of lateral nerves S. vacciniiflora.

Schoepfia Schreberi Gmel. Syst. Veg. 2: 376. 1791. Limoncillo
(Pete"n); Shivecurs-tziquin (Guatemala).

Moist or dry forest or rocky thickets, often in thickets near
streams, 1,200 meters or less; Pete"n: El Progreso; Chiquimula;
Jalapa; Jutiapa; Santa Rosa; Escuintla; Guatemala. Florida;
Mexico; Salvador; Honduras; Panama; West Indies; South America.

A shrub or small tree, usually 9 meters high or less, the branches whitish, sub-
angulate< leaves short-petiolate, mostly ovate, 3-8 cm. long, acute or obtuse,
acute at the base, the venation irregular, the veins usually prominulous and laxly
reticulate; flowers subsessile or short-pedicellate, in few-flowered short-peduncu-
late racemes scarcely longer than the petioles; corolla usually red and 4-lobate;
fruit ovoid or oval, 1 cm. long or smaller, red.

Called "sombra de armado" in Honduras. The Maya name
"sac-bace" is reported from Yucatan.

Schoepfia vacciniiflora Planch, ex Hemsl. Diag. PI. Mex. 5.
1878. Cafe silvestre (Guatemala); Nance de montana (Zacapa).

Moist or dry forest or thickets, sometimes in pine or oak forest,
1,300-2,500 meters; type from Volcan de Fuego, Salvin; Baja
Verapaz; Zacapa; Escuintla; Guatemala; Sacatepe"quez; Chimalte-
nango; Solola; Quiche"; Huehuetenango; Quezaltenango; San Marcos.
Costa Rica; Panama; reported from Venezuela.

A large shrub or often a tree of 6-12 meters, the older branches pale; leaves
coriaceous, often lustrous, on short thick petioles, often blackening when dried,
mostly lance-oblong, rarely lance-ovate, 4-7 cm. long, mostly acute or acuminate
with obtuse tip, acute at the base; racemes mostly cyme-like, few-flowered, short-
pedunculate; corolla greenish or dull red or red tinged with yellow outside, greenish
yellow within; fruit oval, 10-12 mm. long, red or red and yellow.

A common and characteristic shrub or tree in mountain forests,
especially in the central region.

XIMENIA L.

Shrubs or trees, glabrous or pubescent, often armed with spines, these formed
from abortive branchlets; leaves often fasciculate on short spurs, deciduous;

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 91

flowers larger than in most genera of the family, whitish, mostly in short axillary
cymes; calyx small, with 4-5 lobes or teeth, unchanged in fruit; petals 4-5, valvate,
narrow, densely white-barbate within; stamens twice as many as the petals, the
filaments filiform; anthers linear, erect, dehiscent by slits; ovary partially 3-celled,
the style entire, the stigma subcapitate; ovules 3, linear, pendulous; fruit drupa-
ceous, ovoid or globose, with abundant pulp, the stone crustaceous or subligneous;
seed falsely erect, the embryo minute, the endosperm carnose.

At least 8 species, 5 of them in Mexico, one in South Africa,
another in the Pacific islands. Only one is found in Central America.
The generic name commemorates Francisco Xime'nez, native of
Luna in Aragon, who went as a soldier in 1605 to New Spain, where
he later became a lay brother in the Convento de Santo Domingo
de Mexico. In 1615 there was published in the City of Mexico
under his authorship a volume entitled Quatro libros de la naturaleza
y virtudes de las plantas y animates, which is important for the large
amount of original information it contains regarding Mexican plants.

Ximenia americana L. Sp. PI. 1193. 1753. Limoncillo; Man-
zanilla; Putzil (Huehuetenango) ; Tocote de monte (Pete'n) ; Tepenance
(fide Aguilar) ; Abalche, Saaxnic (Pete'n, Maya) ; Membrillo de monte.

Chiefly in dry thickets, rarely in wet or moist places, sometimes
in coastal thickets or mangrove swamps, ascending from sea level
to 2,000 meters; Pete'n; Izabal; Baja Verapaz; El Progreso; Zacapa;
Chiquimula; Jutiapa; Escuintla; Sacatepe"quez; Huehuetenango;
San Marcos. Mexico; British Honduras to Salvador and Panama;
West Indies; South America; Old World tropics.

A densely branched shrub or small tree, rarely more than 6 meters high, the
bark smooth, reddish, the branches abundantly armed with stout sharp spines;
leaves short-petiolate, oblong to elliptic, 3-7 cm. long, rounded or obtuse at each
end, glabrous; flowers yellowish white, fragrant, in few-flowered short-pedunculate
cymes; corolla 4-lobate, subcoriaceous, the linear lobes reflexed; fruit yellow or
reddish, globose or ovoid, 14-17 mm. long.

Sometimes called "cagalero" and "chocomico" in Honduras and
"pepenance" in Salvador; the Maya name "xcuche" is reported
from Yucatan. The fruit is edible, either raw or cooked, having an
acid flavor. Oil is reported to have been extracted from the seeds
in Brazil. The wood has been employed in India as a substitute for
sandalwood. It is fragrant, reddish yellow, fine-textured, very hard
and heavy, the specific gravity about 0.92. The astringent bark has
been employed in some parts of the tropics for tanning. The fruits
are said to contain hydrocyanic acid. Although the shrub is usually
deciduous, it was noted as one of the few shrubs with green leaves in
the Zacapa-Chiquimula region during late April. It is remarkable

92 FIELDIANA: BOTANY, VOLUME 24

for its wide altitudinal distribution, from sea level into the high
mountains.

BALANOPHORACEAE

Fleshy herbs, parasitic on the roots of other plants, usually yellowish, without
chlorophyll; rhizomes tuberous, often very large, simple or lobate, sometimes
emitting cylindric branches, these glabrous or tomentose, naked or squamose,
epigaean or hypogaean; peduncles short or elongate, cylindric, naked or surrounded
by an annulus; flowers small and numerous, unisexual, densely crowded in simple
or very rarely branched, unisexual or androgynous inflorescences (spadices), these
ovoid, clavate, cylindric, globose, or fusiform; staminate flowers naked or with a
3-8-lobate valvate perianth; stamens solitary or binate in the naked flowers, in
those with a perianth usually as many as the perianth lobes and opposite them;
filaments free or connate into a tube or column; anthers in the naked flowers
attached by the base or dorsal surface, 2-celled, dehiscent by lateral or anterior
slits, in the perigoniate flowers basifixed, free or connate, 2-celled or 4-many-celled;
perianth none in the pistillate flowers or adnate to the ovary, the limb small,
truncate, 2-labiate, or tubular; ovary globose or ellipsoid and compressed, or
prismatic-obovoid, 1-3-celled; styles terminal, either 1 and filiform or subclavate,
or 2 and short or elongate, the stigmas simple or capitellate, or the stigma rarely
sessile and discoid; ovules solitary in the cells, usually pendulous; fruit small,
nut-like, crustaceous or somewhat fleshy or coriaceous, 1-celled, 1-seeded; seed
globose or compressed, the testa very thin or none; endosperm usually oily.

About 15 genera and twice as many species, in both hemispheres,
mostly in tropical regions. Two other genera, Corynaea and Langs-
dorffia, are represented in Costa Rica.

HELOSIS L. Richard

Glabrous fleshy herbs, reddish or yellowish; rhizomes tuberous, emitting
elongate naked subterranean branches; peduncles erect, naked, short or elongate,
annulate at the base or higher; spadices broadly ovoid or globose, covered with
peltate, hexagonal, valvately connected bracts, these deciduous; flowers of either
sex crowded in mammillae corresponding to the bracts, mixed with very numerous,
linear-clavate hairs; bractlets none; tube of the staminate perianth cylindric, the 3
lobes ovate, concave, valvate; stamens 2, the filaments connate into a tube, their
apices free; anthers basifixed, ovate-cordate, connate; pistillate perianth superior,
2-labiate, the lobes triangular, obtuse; ovary ellipsoid, 1-celled; styles 2, elongate,
filiform, deciduous, the stigmas capitellate; ovule 1, pendulous from the apex of
the cell; fruit nut-like; seed oblong or subglobose, the endosperm oily.

Three species have been described, all of them perhaps to be
reduced to H. cayennensis (Swartz) Spreng. of northern South
America. Only the following is known from Central America.

Helosis mexicana Liebm. Forh. Vid. Skand. Nat. 4: 181. 1844.
Mazorca de culebra (Huehuetenango).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 93

Moist or wet, dense, mixed forest, usually in dark places among
rotting leaves, 1,400 meters or less; Pete'n; Alta Verapaz; Izabal;
Huehuetenango. Southern Mexico; British Honduras; Honduras;
Costa Rica.

Plants white to brown or dull orange, glabrous, arising from a much-branched
mass of coralline rootstocks; peduncles solitary or often several together, stout,
erect, 6-10 cm. long; spadix oval or oblong, 1.5-4.5 cm. long, 1.5-2.5 cm. broad,
rounded at the apex.

In general appearance this plant resembles some of the mush-
rooms, with which it is likely to be confused at first glance. In
habit and habitat it is suggestive also of such Orobanchaceae as
Conopholis. It is rather frequent in the lowlands of the Atlantic
coast of Central America.

ARISTOLOCHIACEAE

Herbs or rarely shrubs, often scandent, frequently strong-scented; leaves
alternate, petiolate, often cordate, entire or lobate; stipules none, but pseudo-
stipules sometimes present; flowers medium-sized or large, mostly green, yellowish,
or brown-purple, terminal, axillary, or lateral at the base of the stem, solitary or
in cymes or racemes, perfect; perianth simple, adnate below to the ovary, variously
produced above the ovary, equally 3-lobate or asymmetric and entire, dentate, or
3-lobate, the lobes valvate; stamens 6 or numerous, affixed about the apex of the
ovary or the style column in 1-2 series, free or adnate to the column, erect, the
anther cells parallel, distinct, dehiscent by longitudinal slits; disk none; ovary
inferior or rarely semi-superior, perfectly or imperfectly 4-6-celled, the placentae
intruded from the cell walls and connivent or coalescent in the center of the ovary;
styles united to form a short thick column, this divided at the apex into 3-8 stigma-
tose lobes; ovules numerous in each cell, anatropous, horizontal or pendulous;
capsule irregularly opening or often septicidally or placenticidally dehiscent; seeds
numerous, horizontal or pendulous; endosperm copious, carnose.

Six genera are recognized, widely distributed. The only other
American one, Asarum, has a small number of species in the United
States and Canada.

ARISTOLOCHIA L.

Herbs, often scandent, or sometimes scandent shrubs, rarely erect shrubs or
small trees; leaves usually petiolate, entire or lobate, often cordate at the base;
peduncles axillary or lateral, solitary, fasciculate, or racemose; bracts none or
present at the bases of the peduncles and simulating stipules, sometimes present
on the peduncle below the ovary; perianth adnate to the base of the ovary, more
or less distinctly articulate above the ovary, around the stamens and gynoecium
utricular, globose or oblong, above the androecium constricted or contracted and
often annulate within, above this tubular, then expanded into a limb, this highly
variable in form, entire, 1-2-labiate, or 3-lobate; stamens usually 6 and 1-seriate;

94 FIELDIANA: BOTANY, VOLUME 24

anther cells extrorsely dehiscent; ovary inferior, usually perfectly 6-celled; stigma
lobes usually 3 or 6; capsule septicidally or placenticidally dehiscent, usually
from the base upward; seeds horizontal, compressed.

More than 200 species, widely distributed but chiefly in the
tropics. A few additional species grow in southern Central America.

Leaves 3-lobate A. trilobata.

Leaves not lobate, entire.

Stems densely hirsute with long brown spreading hairs A. pilosa.

Stems not hirsute.
Pseudostipules present, large and conspicuous.

Leaves broadly rounded at the apex; calyx limb bilabiate A. ringens.

Leaves acute or obtuse; calyx limb entire A. anguitida.

Pseudostipules none, or minute and inconspicuous.

Flowers very large, the calyx limb 8-10 cm. broad or larger, long-caudate
at the apex. Plants scandent, herbaceous; leaves long-petiolate, large,

broadly ovate-cordate, acute A. grandiflora.

Flowers much smaller or, if large, the calyx limb not caudate.

Leaves broadly ovate-cordate or deltoid-cordate, much the broadest
near the base, glabrous or essentially so.

Limb of the calyx 5-7 cm. wide; leaves conspicuously deltoid.

A. odoratissima.

Limb of the calyx much less than 5 cm. wide; leaves not noticeably
deltoid.

Stems woody; leaves mostly 12-18 cm. wide A. Schippii.

Stems herbaceous; leaves mostly 4-6 cm. wide A. inflata,

Leaves chiefly oblong, elliptic-oblong, or obovate, usually broadest at
or above the middle, sometimes broadest near the base but then
conspicuously pubescent, at least beneath.
Leaves cordate at the base, usually deeply so.

Leaves, at least the younger ones, densely lanate beneath .A. sericea.
Leaves puberulent or merely hirtellous beneath.
Leaves acute or acuminate A. sp.

Leaves rounded to obtuse at the apex or subacute.

A. Chapmaniana.

Leaves obtuse to truncate at the base, some of the leaves rarely sub-
cordate.

Leaves densely pilose or hirsute on the upper surface A. mollis.

Leaves glabrous on the upper surface or practically so.

Leaves obtuse or rounded at the apex A. maxima.

Leaves acuminate.

Leaf blades glabrous beneath A. Steyermarkii.

Leaf blades puberulent, hirtellous, or pilose beneath .A. arborea.

Aristolochia anguicida Jacq. Enum. PI. Carib. 30. 1760.
A. loriflora Masters, Bot. Jahrb. 8: 220. 1887 (type from Chiquimula,
F. C. Lehmann 1702). Guaco.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 95

Moist or wet thickets of the Oriente, 180-900 meters; Zacapa;
Chiquimula; Jutiapa; Santa Rosa; Escuintla. Salvador; Nicaragua;
Costa Rica; West Indies.

A small or large, herbaceous vine, climbing over shrubs or small trees, the
stems puberulent or almost glabrous; pseudostipules conspicuous, large, orbicular
or reniform, broadly rounded at the apex, mostly 1.5-2 cm. broad; leaves on long
slender petioles, oval-ovate or deltoid-ovate, 4.5-10 cm. long, 3.5-8 cm. wide, very
obtuse to acute, glabrous above, puberulent beneath, the ultimate veins thickened,
prominent, and forming a close reticulation; bracts oval; pedicels 2.5-3.5 cm. long;
perianth yellow-green, the lower utricle-like inflated portion 1 cm. long or less,
the tube 1.5-2 cm. long, slender below, dilated above, the limb linear from a
broader base, 1.5-3 cm. long; capsule oval, about 2.5 cm. long and almost 2 cm.
broad, rounded at base and apex, glabrous, conspicuously costate and transverse-
striate.

Known in Salvador also by the names "chompipito" and "chom-
pipe," in reference to the form of the flowers. The stems are used
there by laundresses to rub dirt from clothes, and the plant is a
domestic remedy for pains in the stomach. The name "guaco" often
applied in Central America to Aristolochia species would indicate
that they were employed as remedies for snake bites. The perianth
in this species often has dark brown-purple stripes, especially within.

Aristolochia arborea Linden, Cat. 13. 1858; Hooker, Bot. Mag.
pi. 5295. 1862.

Wet forest, about 350-1,250 meters; Alta Verapaz; Solold. Type
from Chiapas.

Described as either a small tree or a large vine, the young branches densely
pilose with appressed brownish hairs, the old branches covered with thick corky
ridged bark; leaves large, on stout petioles 1 cm. long, oblong or lance-oblong, 20-35
cm. long, 6-9 cm. wide, long-acuminate, rather obliquely rounded at the base,
penninerved, glabrous above, densely pilose beneath with weak hairs; flowers
clustered on the lower part of the trunk or stem below the leaves; perianth purple-
brown, 8-9 cm. long, densely and finely pubescent, the tube inflated, striate, the
limb broadly cordate, abruptly inflexed-acuminate at the apex, the throat of the
tube closed by a large orbicular puberulent-glandular disk; capsule clavate, 10 cm.
long or larger.

The plant is in cultivation in the Jardin Botanico of Guatemala.

Related to this species but doubtless distinct are three collections
from the Pacific lowlands, all unfortunately sterile, and all probably
representing undescribed species. One is a low erect shrub with
smaller leaves glaucescent beneath, plentiful in mixed forest between
Retalhuleu and the coast. Another collection from Dept. Guate-
mala is noteworthy for its long and very narrow leaves whose pubes-

96 FIELDIANA: BOTANY, VOLUME 24

cence is unlike that of A. arborea; and the last is a tree of San Marcos,
said to be 8 meters high, whose leaves somewhat resemble those of
A. arborea in size and form, but have different pubescence.

Aristolochia Chapmaniana Standl. Contr. Arnold Arb. 5: 60.
pi. 9. 1933. A. maxima L. var. cordata Standl. Field Mus. Bot. 8:
136. 1930.

British Honduras, 60 meters, and probably extending into Peten;
sterile collections from the lowlands of Alta Verapaz, Izabal, and
Retalhuleu perhaps represent the same species. Panama.

A large or small, woody vine, the stems striate, sparsely hispidulous; leaves
short-petiolate, subcoriaceous, oblong or narrowly oblong, 9-20 cm. long, 2.5-7.5
cm. wide, acute or subobtuse, sometimes rounded and abruptly pointed, deeply
and narrowly cordate at the base, more or less lustrous above and glabrous or
nearly so, 5-7-nerved at the base and penninerved above, minutely hispidulous or
puberulent, sometimes glabrate; flowers axillary, the peduncles elongate, 1-flow-
ered; bracts linear-lanceolate, 10-15 mm. long; perianth dark brownish white and
yellow, sparsely pilose, the utricular basal portion 4-5 cm. long and 2 cm. wide,
the tube subrefracted, 2.5-3.5 cm. long, the limb lance-oblong, about 7 cm. long
and 2 cm. wide, acute and filiform-caudiculate; capsule long-stipitate, obovoid,
about 6 cm. long, blackish-ferruginous, conspicuously costate.

Called "guaco" in British Honduras.

Aristolochia grandiflora Swartz, Fl. Ind. Occ. 1566. 1806.
A. gigas Lindl. Bot. Reg. pi. 60. 1842 (based on plants of Guate-
malan origin cultivated in England). A. gigas var. Sturtevantii W.
Watson, Card. & For. 4: 546. 1891. Chompipe; Guegiiecho; Hedion-
dilla; Chumpa; Alcatraz (north coast); Bonete de fraile, Bonete del
diablo (Pete"n) ; Guegiiecho de zope; Chompipona (fide Aguilar); Flor
de pato.

Usually in wet thickets, often on stream banks, common in
second growth in the tierra caliente, chiefly at 600 meters or less,
rarely ascending to 1,300 meters; sometimes planted in gardens;
Pete"n; Alta Verapaz; El Progreso; Izabal; Jalapa; Santa Rosa;
Escuintla; Suchitepe"quez; Retalhuleu; San Marcos. Southern
Mexico; British Honduras to Panama; West Indies.

A large herbaceous vine, often covering medium-sized trees, the stems puberu-
lent or glabrous; leaves long-petiolate, broadly ovate-cordate, 8-25 cm. long, acute
to long-acuminate, with a deep basal sinus, puberulent or glabrate, thin, slightly
paler beneath; flowers axillary, pendent, solitary, huge; tubular portion of the
perianth 12-20 cm. long, sparsely pilose outside, the limb oval, commonly 15-45
cm. long and very broad, hairy and dark purple within, whitish outside, bearing
at the apex a slender linear tail-like pendent appendage as much as a meter long;
capsule oblong, about 10 cm. long and 4.5 cm. thick.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 97

This remarkable vine is often cultivated in United States green-
houses under such names as duck flower and pelican flower, the form
of the perianth just before opening suggesting a duck and being of
about the same size. The flower is one of the largest produced by
any plant, and doubtless is the largest flower of America. It has a
strong and disgusting odor that in Guatemala sometimes is the basis
of rather fantastic tales. It is stated that it "draws" insects — which
it probably does — and "eats" them. The plant is well known in
those parts of the country where it grows naturally, since such a
strange blossom would attract attention anywhere. The plant is
sometimes called "guaco" in Salvador, and the roots are one of the
reputed remedies for bites of snakes and other poisonous animals.
The roots have been reported as poisonous to hogs, the name "poison
hog-meat" being formerly applied to the plant in Jamaica. Des-
courtilz goes further and states that in the West Indies the roots were
sometimes used for criminal poisoning of human beings.

Aristolochia inflata HBK. Nov. Gen. & Sp. 2: 145. pi. 111.
1817. A. gibbosa Duchartre, Ann. Sci. Nat. IV. 2: 53. 1854 (type
from San Antonio, Retalhuleu, Hartweg 566). (?)A. podocarpa
Bertol. Fl. Guat. 437. 1840 (type from Escuintla, Velasquez).

Moist or rather dry thickets and forest, 600 meters or less;
Jutiapa; Santa Rosa; Escuintla (?); Suchitepe"quez; Retalhuleu; San
Marcos. Panama; Colombia.

A small or rather large, herbaceous vine, glabrous throughout or nearly so;
leaves long-petiolate, broadly ovate-cordate, 6-10 cm. long, acute or obtuse, with
a rather deep basal sinus and rounded basal lobes; flowers axillary, solitary, long-
pedicellate; perianth 3-3.5 cm. long, almost glabrous, pale greenish white outside,
pale yellow within, the inflated basal portion almost 1 cm. long, semiglobose, the
tube short, the limb about 3.5 cm. long and 2.5 cm. wide, acute or very obtuse;
capsule about 3.5 cm. long and 8 mm. thick.

Aristolochia maxima Jacq. Enum. PI. Carib. 30. 1760. A.
geminiflora HBK. Nov. Gen. & Sp. 2: 118. pi. 117. 1817. Guaco;
Canastilla; Tecolotillo.

Dry to wet thickets, sometimes in forest, 1,200 meters or less;
El Progreso; Baja Verapaz; Zacapa; Chiquimula; Jutiapa; Santa
Rosa; Escuintla; Guatemala. Southern Mexico; British Honduras
to Panama; northern South America.

Often a large woody vine; older stems covered with large corky ridges, when
young puberulent; leaves short-petiolate, subcoriaceous, mostly oblong or obovate-
oblong, 7-18 cm. long, usually rounded or very obtuse at the apex, sometimes

98 FIELDIANA: BOTANY, VOLUME 24

apiculate, obtuse or rounded at the base or occasionally shallowly and broadly
cordate, glabrous above, sparsely or densely puberulent or pilose beneath, the
venation prominent and reticulate; racemes few-many-flowered, axillary or more
often crowded in dense masses at the base of the stem or on its lower part; perianth
densely puberulent outside, commonly 8-10 cm. long, purple-brown, the basal
portion utricular, the tube short, abruptly reflexed, the limb broadly ovate, acute
or obtuse and mucronate; capsule about 10 cm. long and 4 cm. thick, dark fer-
ruginous or blackish, coarsely costate.

A common and characteristic vine in many of the drier areas of
Central America, conspicuous during the dry season because of its
large seed pods, which hang for a long time upon the branches. The
flowers are likely to be overlooked, massed as they usually are at
the base of the main stem, although often they are produced along
the terminal branches. The tender young pods are reported to be
cooked and eaten in Costa Rica.

Aristolochia mollis Standl. & Steyerm. Field Mus. Bot. 23:
155. 1944. Hoja del aire.

Known only from the type, Huehuetenango, canyon tributary
to Rio Trapichillo, between Democracia and canyon of Chamushu,
about 1,000 meters, Steyermark 51269.

A scandent shrub, the branches stout, very densely pilose-tomentose with
brownish hairs, the internodes rather short; leaves large, short-petiolate, thick-
membranaceous or chartaceous, the stout petioles 6-8 mm. long, densely pilose-
tomentose; leaf blades oval or oval-elliptic, 13-18 cm. long, 6-10 cm. wide, rounded
to very obtuse at the apex, sometimes apiculate, narrowly rounded at the base, the
very base sometimes emarginate, entire, densely pilose-hirsute on the upper sur-
face, the nerves and veins prominulous, laxly reticulate, slightly paler beneath,
very densely velutinous-pilose, the hairs spreading, pale brownish, the costa rather
stout, the lateral nerves about 7 on each side, the veins prominent, laxly reticulate;
peduncles axillary, apparently 1-flowered, about 7 mm. long; perianth brown-
purple, glabrous within, densely short-pilose outside with spreading hairs, the
basal portion utricular, 2.5 cm. long, the tube short, abruptly reflexed, the limb
rounded-ovate, 3.5 cm. long; young capsule borne on a pedicel 1 cm. long, clavate-
oblong, 2.5 cm. long, 5-7 mm. broad near the apex, rounded at the base, densely
pilose with soft ascending brownish hairs.

The species is probably closely related to A. asclepiadifolia
Brandeg., of Veracruz, which it much resembles in general appear-
ance. A decoction of the leaves is used in domestic medicine in
Huehuetenango as a remedy for stomach ailments.

Aristolochia odoratissima L. Sp. PI. ed. 2. 1362. 1763. Guaco;
Patito.

Pete*n. Southern Mexico; Honduras; Costa Rica; Panama;
West Indies; South America.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 99

A large, often woody vine, the stems glabrous; leaves long-petiolate, deltoid-
cordate, 7-15 cm. long, obtuse to short-acuminate, usually with a very shallow
basal sinus, glabrous or nearly so, paler beneath; flowers axillary, solitary, long-
pedicellate; perianth puberulent outside, brown-purple and cream, the basal
portion utricular, the narrow tube short, the limb broadly cordate-ovate, 6-11
cm. long, rounded and mucronate at the apex; capsule 7-10 cm. long, 1-2 cm.
thick, angulate.

Called "cocoba" in Tabasco.

Aristolochia pilosa HBK. Nov. Gen. & Sp. 2: 116. pi. 113.
1817. A. pilosa var. ligulifera Masters in Bonn. Smith, Bot. Gaz.
33: 256. 1902 (type from Cubilgiiitz, Alta Verapaz, Tuerckheim
7768). Sombrerito; Hediondilla; Hicac (Cacchiquel).

Moist thickets, 1,500 meters or less; Alta Verapaz; Izabal;
Quezaltenango; San Marcos. Southern Mexico; British Honduras
to Panama; South America.

A scandent herb, the stems hirsute with brown hairs; leaves long-petiolate,
cordate-ovate, 7-18 cm. long, obtuse or rounded at the apex, deeply cordate at the
base, glabrous above, densely brown-pilose or hirsute beneath; flowers axillary,
solitary, long-pedunculate; perianth 5-7 cm. long, hirsute, the basal portion in-
flated, the tube slender, the limb ovate or oblong-ovate, obtuse, pale green with
purple-brown dots, the throat dark purple-brown, the limb smooth or muricate-
ligulate within; capsule narrow, 6 cm. long or more.

In the typical form of the plant the perianth limb is smooth
within; in var. ligulifera it is ligulate-appendaged.

Aristolochia ringens Vahl, Symb. Bot. 3: 99. 1794.

Cultivated for ornament, Sacatepe"quez; a sterile collection from
Rio Guacalate, Escuintla, 700 meters, probably represents the same
species. Jamaica and Cuba; Colombia and Venezuela.

A large glabrous vine; pseudostipules very large and conspicuous, reniform,
pale green; leaves long-petiolate, reniform-cordate, 6-15 cm. wide, broadly rounded
at the apex, pale beneath; flowers axillary, long-pedunculate, the basal portion
large and inflated, 5 cm. long, the broad tube refracted, the limb bilabiate, the
upper lip lanceolate, obtuse or subobtuse, the lower lip with a long narrow base
abruptly expanded into an ovate obtuse blade, the whole perianth pale greenish,
with dark purple veins, or the lips dark purple.

Aristolochia Schippii Standl. Field Mus. Bot. 8: 8. 1930.

Type from Big Creek, British Honduras, 15 meters, Schipp 75;
probably extending into Pete*n. Veracruz.

A large woody vine, sometimes 10 meters long, glabrous throughout or nearly
so, the older branches covered with thick corky ridged bark; leaves long-petiolate,

100 FIELDIANA: BOTANY, VOLUME 24

subcoriaceous, lustrous, rounded-cordate, about 24 cm. long and 18 cm. wide or
smaller, acute or short-acuminate, deeply cordate at the base, 5-nerved; flowers
apparently arising from naked stems, fasciculate, the peduncles 2-2.5 cm. long;
perianth glabrous, yellowish with reddish brown veins, 5 cm. long, slightly curved,
the inflated basal portion 1 cm. long and 7 mm. broad, the tube 1.5 cm. long, the
blade almost 3.5 cm. long, 14 mm. wide, long-acuminate; capsule 11 cm. long, 1 cm.
thick, abruptly contracted and stipitate, subterete, 6-costate, glabrous, contracted
and acuminate at the apex.

Called "con tray erba" in Veracruz.

Aristolochia sericea Benth. PI. Hartw. 81. 1841.

Moist forest or thickets, 1,500-1,800 meters; Guatemala; Saca-
tepe'quez. Type from Comitan, Chiapas.

A woody vine, the stems lanate or tomentose with whitish or brownish hairs;
leaves short-petiolate, oblong or oval-oblong, 5-12 cm. long, subcoriaceous,
rounded to short-acuminate at the apex, cordate at the base, tomentulose or
glabrate above, usually densely whitish-tomentose beneath; peduncles axillary,
bracteate, shorter than the leaves; perianth villous, the tube gibbous and recurved,
the limb oblong, about 2.5 cm. long, trilobate at the apex, the lobes lanceolate,
linear-acuminate, 6 mm. long; capsule 3.5 cm. long, tomentose.

Although Bentham describes the leaves as subacute, a photograph
of a specimen of the original collection in the Berlin herbarium
shows them as rounded at the apex. In the recent specimens referred
here the leaves vary from obtuse to short-acuminate. The latter
material, all without flowers, agrees fairly well in foliage with the
original collection of A. sericea although it is quite possible that
flowers will show the Guatemalan plant to be specifically distinct.

Aristolochia Steyermarkii Standl. Field Mus. Bot. 22: 329.
1940. Guaco de montana.

Type from Quezaltenango, Quebrada Geronimo, Finca Pirineos,
southern slopes of Volcan de Santa Maria, 1,300-2,000 meters,
Steyermark 33455.

A tree of 6 meters, the slender branches glabrous; leaves on petioles 1-1.5 cm.
long, narrowly oblong or lance-oblong, 15-23 cm. long, 4-6 cm. wide, short- or
long-acuminate, obtuse and somewhat unequal at the base, glabrous, pale and
glaucescent beneath, penninerved; flowers axillary, solitary, the peduncles in fruit
2-3 cm. long; capsule narrow, glabrous, lustrous, the valves recurved after dehis-
cence, 4-4.5 cm. long, 6 mm. wide.

Aristolochia trilobata L. Sp. PL 960. 1753.

British Honduras, and to be expected in Pete"n or Izabal. Hon-
duras; Costa Rica; Panama; West Indies; South America.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 101

A small or large vine, the slender stems puberulent or glabrous; pseudostipules
large and conspicuous, green; leaves long-petiolate, broader than long, subcordate
at the base, 3-lobate to the middle or more deeply, the lobes oblong or obovate,
obtuse or rounded at the apex, green and glabrous above, glaucous and puberulent
beneath; flowers axillary, long-pedunculate; perianth glabrous, yellowish green
outside, dark red or purple within, the inflated basal portion 4-5 cm. long, the
broad tube 5-6 cm. long, the limb ovate, contracted at the apex into a slender cord-
like appendage 12-15 cm. long; capsule cylindric, 5-7 cm. long, acute at the base,
costate.

Called "media-luna" in Honduras; known in British Honduras
as "contrayerba," "country ebo," and "contrebo." The foliage is
much like that of some species of Passiflora. Used in British Hon-
duras as a domestic remedy for fevers.

Aristolochia sp.

Plants scandent, apparently low but perhaps in age much elongate, the stems
slender, pilose with short, mostly reflexed hairs; leaves short-petiolate, oblong or
narrowly triangular-oblong, 4.5-9 cm. long, acute or acuminate, cordate at the
base, with short rounded lobes, pilose above or glabrate, short-pilose beneath,
3-nerved at the base and penninerved above the base, the venation prominent
and reticulate beneath; flowers and fruit unknown.

Represented by three collections from Volcan de Quezaltepeque,
Chiquimula, Sierra de las Minas, El Progreso, and Volcan de Agua,
Sacatepe"quez, at 1,800-2,100 meters. Probably a new species, but
possibly only juvenile plants of A. sericea or some related species.
Sterile material of one or two other species, probably undescribed,
also has been collected in Guatemala.

RAFFLESIACEAE

Fleshy or almost dry parasites, on roots, stems, and branches of trees and
shrubs, the leaves reduced to scales; chlorophyll none; flowers often large, but in
American genera often almost minute, solitary, or sometimes spicate, by abortion
unisexual, sometimes polygamous or perfect; calyx more or less epigynous, of
4-10 imbricate segments; anthers sessile in 1-3 series about a fleshy column,
2-celled, opening by longitudinal slits or terminal pores; pollen often viscous;
ovary inferior or subinferior, 1-celled but the placentae sometimes extending
almost to the middle; stigma undivided, discoid or lobate, or the stigmas numerous
at the apex of the ovary; ovules very numerous, on parietal placentae or from the
apex of the cell; fruit fleshy, indehiscent or irregularly ruptured; seeds very numer-
ous, minute; endosperm cellular; embryo minute.

Genera about 6, in the tropics of both hemispheres. Only the
following are known from America. Rafflesia Arnoldii R. Br. of
Sumatra is believed to bear the largest flower of any plant, about a
meter broad. In contrast, flowers of the genera Apodanthes and

102 FIELDIANA: BOTANY, VOLUME 24

Pilostyles have exceedingly small flowers, and the entire plants are
among the smallest known.

Plants terrestrial, on the roots of trees, mostly 5-8 cm. high Mitrastemon.

Plants parasitic on the branches of shrubs or trees, mostly less than 5 mm. high.
Perianth segments epigynous, unguiculate, deciduous, the inner ones slightly

connate; plants parasitic on Flacourtiaceae Apodanthes.

Perianth segments perigynous, broad at the base, the inner ones free; plants
parasitic on Leguminosae Pilostyles.

APODANTHES Poiteau

Minute plants, parasitic on branches of Flacourtiaceae, usually growing in
colonies, 1-flowered, arising from a ligneous cupule, whitish or brownish in age;
pistillate flowers almost sessile, the stem bearing 3 whorls of scales, the lowest
verticel of 2 free scale-like leaves, the second whorl of 4 scales connate at the base;
segments of the third verticel petaloid, rounded, unguiculate at the base, decidu-
ous, leaving punctiform scars; ovarial column cylindric, narrowed above and
surrounded by an annular stigma; ovary semisuperior, 1-celled, many-ovulate;
ovules very numerous, anatropous, on long funicles; fruit like the flower except for
the deciduous perianth segments, baccate; testa of the seed osseous.

Two species, the other in Venezuela.

Apodanthes Caseariae Poit. Ann. Sci. Nat. 3: 422. 1824.

British Honduras, Temash River, little above sea level, on
Casearia, W. A. Schipp S916. Known in Central America also from
Barro Colorado Island, Canal Zone; Guianas; Brazil.

Buds subglobose, 3-4 mm. long; perianth segments suborbicular, entire, sub-
coriaceous, whitish at first, sometimes reddish in age.

The senior author has spent much time in Guatemala and else-
where in Central America searching for this parasite, but without
success. It is probably rare, although there are many Flacourtia-
ceous plants on which it might well be found.

MITRASTEMON Makino

Low stout plants, parasitic on the roots of trees, often forming dense colonies,
the thick stems covered with large coriaceous obtuse scales; flowers perfect, solitary
and terminal, erect; perianth hypogynous, gamophyllous, cupuliform, truncate or
somewhat 4-lobate, persistent; stamens hypogynous, united to form a caducous
tube; anthers numerous, in several series; ovary superior, sessile, 1-celled, with 9-13
or more parietal placentae, these fleshy; ovules numerous, more or less stipitate,
anatropous; style terminal, short, very thick, the stigma conic; fruit baccate,
indehiscent; seeds numerous, small, with a hard testa.

Three other species are known, in Japan and Formosa.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 103

Mitrastemon Matudai Yamamoto, Bot. Mag. Tokyo 50: 539.
ill. 1936.

Dense wet forest, near a stream bank, 1,500 meters; Alta Verapaz
(along road between Tactic and the divide on the road from Tactic
to Tamahu, Standley 91455). Chiapas, the type from Mount
Ovando, Escuintla, at 1,500-1,900 meters.

Plants glabrous, somewhat fleshy, pale yellowish or whitish at first but soon
darkening, forming dense colonies, 4-8 cm. high, with the scales 3-3.5 cm. thick;
stem cylindric, 1 cm. in diameter or more; scales few, imbricate, opposite, lustrous,
unequal, the lower ones 'smallest, the upper ones gradually larger, ovate or broadly
ovate, 1.5-3 cm. long, obtuse; perianth short-cylindric, about 6 mm. high and 17
mm. broad; androecium calyptriform, the stamen tube 2 cm. long, striate outside,
the anther tube 6 mm. long; fruit cylindric, terete, about 1 cm. long and 1.5 cm. in
diameter; placentae about 15; style 2 mm. long, the stigma conic, 7 mm. long and
of equal diameter; seeds very numerous, minute, reticulate.

The original collection was parasitic on Quercus. The Guate-
malan plants were believed to be attached to the roots of Carpinus.
The plant is a curious one, and remarkable for its isolated occurrence,
far distant from the range of the Asiatic species. The senior author
was able to find the plant at only one locality, where under a single
tree there were numerous individuals, barely exserted above the sur-
face of the wet soil. He first noted them while he was in the tree
gathering epiphytes. Looking down he saw what appeared to be
Geaster or some other fungus, and was very much surprised — and
puzzled — when he discovered that the strange objects were flowering
plants, whose affinities were not at once apparent.

PILOSTYLES Guillemin

Minute plants parasitic on branches of Leguminosae, in Guatemala on
Calliandra, generally in dense colonies and appearing like warts on the branches,
usually reddish or purplish, arising from depressed cupules in the branches; leaves
scale-like, in 2-3 verticels; flowers solitary, terminal, dioecious, the perianth seg-
ments attached by a broad base; anthers transversely dehiscent; ovary inferior,
the ovules scattered irregularly over its inner surface, the stigma annular; ovules
anatropous; fruit very small, baccate, surrounded by the dry perianth segments.

About a dozen species, reported from other regions also on Inga,
Bauhinia, Dalea, Galactia, and perhaps other hosts. Only one species
has been found in Central America but several are known from
Mexico. The species have not been studied recently and it is not
known how many of those described are valid. The plants are hard
to find but once one has found them it is easier to locate them a
second time, and despite their small size they may be seen from some

104 FIELDIANA: BOTANY, VOLUME 24

distance because of the peculiar warty appearance they give to the
branches.

Pilostyles mexicana (Brandeg.) Rose, Contr. U. S. Nat. Herb.
12: 264. 1909. Apodanthes mexicana Brandeg. Zoe 5: 245. 1908.

On Calliandra of two or more species, 1,000-1,900 meters;
Zacapa; Guatemala (Fiscal); Chimaltenango (between Chimalte-
nango and San Martin Jilotepeque) ; Huehuetenango. Southern
Mexico; Honduras.

Plants ovoid, brownish or reddish, about 3 mm. long; bracts and perianth
segments all much like, about 12, unequal, orbicular or ovate, minutely erose.

POLYGONACEAE. Knotweed Family

Herbs, shrubs, or trees, sometimes scandent; leaves alternate or sometimes
opposite, variable in form, rarely lobate or divided, the petiole often dilated and
clasping, its base often membranaceous-marginate, the margin continuous with
an intrapetiolar ocrea that sheathes the stem; flowers usually small, solitary or
commonly fasciculate within a cuplike bract (ocreola), the flower fascicles axillary
or disposed in spikes or racemes; pedicels usually articulate; flowers perfect or
sometimes unisexual, regular; perianth inferior, calyx-like or colored, the lobes or
segments 4-6, imbricate in 1-2 series, equal, or the outer ones smaller or larger,
unchanged in fruit or some of them accrescent and embracing the fruit; stamens
usually 6-9, crowded on a central disk, the filaments filiform or dilated at the base,
free or connate at the base; anthers 2-celled, usually versatile, the cells parallel or
subparallel, dehiscent by longitudinal slits, an annular disk often present at the
base of the perianth, entire, crenate, or dentate; ovary superior, usually sessile,
trigonous or compressed, 1-celled; styles mostly 3 or 2, apical, distinct or somewhat
connate, the stigmas capitate, peltate, or fimbriate; ovule 1, orthotropous, sessile
or erect at the apex of an elongate funicle; fruit an achene, trigonous or compressed,
usually surrounded by the persistent perianth, the pericarp crustaceous or rarely
coriaceous or indurate; seed erect, sessile or short-stipitate, often sulcate or lobate,
the testa membranaceous; endosperm abundant, farinose, uniform or ruminate;
embryo usually somewhat ex centric or lateral, curved or straight; cotyledons
plane, narrow or broad, rarely very broad and convolute, the radicle long or
short, superior or ascending.

About 30 genera, widely distributed in tropical and temperate
regions of both hemispheres. All the Central American genera are
represented in Guatemala. The family contains but few plants of
great economic importance. One of the most important is buck-
wheat (trigo negro, trigo sarraceno), cultivated in Europe, Asia, and
North America for its seeds, from which a kind of flour is made.
This flour is much used in the United States for making a special
kind of griddle cakes, and it probably may be found in the delica-
tessen shops of Guatemala City for sale to foreigners. So far as we

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 105

know, buckwheat (Fagopyr'um esculentum Moench) is never grown
in Central America, although it might be expected to thrive in
mountain regions.

Plants scandent by tendrils terminating the inflorescences Antigonon.

Plants erect or, if scandent, without tendrils.
Plants herbaceous, never scandent.

Leaves palmately nerved Rheum.

Leaves penninerved.

Stigmas capitate; inner sepals not accrescent, not bearing tubercles.

Polygonum.
Stigmas fimbriate; inner sepals usually accrescent and bearing a tubercle

on the outer surface Rumex.

Plants woody or, if herbaceous, scandent.

Plants leafless, the stems compressed and ribbon-like Muehlenbeckia.

Plants with normal leaves, the stems not compressed and ribbon-like.
Perianth normally 5-parted.
Perianth lobes winged.

Pedicels not winged; filaments pubescent; leaves orbicular.

Neomillspa ughia .
Pedicels winged; filaments glabrous; leaves ovate or narrower.

Podopterus.
Perianth lobes not winged.

Flowers perfect; plants trees or large shrubs, not scandent . . . Coccoloba.

Flowers usually polygamo-dioecious; plants vines or small prostrate

shrubs Muehlenbeckia.

Perianth 6-parted or rarely 3-parted.

Flowers perfect; outer perianth segments broadly ovate. . .Gymnopodium.

Flowers dioecious; outer perianth segments of pistillate flowers narrowly
spatulate.

Stamens numerous; fruit acutely triquetrous Triplaris.

Stamens 9; fruit obtusely angulate Ruprechtia.

ANTIGONON Endlicher

Scandent herbs, sometimes suffrutescent below; leaves alternate, cordate or
deltoid, the petioles somewhat amplexicaul; ocreae small or reduced to a trans-
verse line; flowers perfect, fasciculate within a small bract, the fascicles racemose,
the racemes terminal or arising from the upper axils, the rachis often prolonged
into a tendril; pedicels short, often elongate in fruit, the flowers at first small,
usually pink, the perianth accrescent in fruit, 5-parted, the segments erect, mem-
branaceous-scarious, the outer 3 larger, broadly cordate, the 2 inner ones narrower,
oblong; stamens 7-8, the filaments filiform, connate at the base, the anthers ovate;
ovary 3-angulate, narrowed to the 3 short styles, the stigmas capitate or peltate;
ovule at first pendulous from a long funicle, finally erect; achene trigonous, hidden
by the accrescent perianth; seed subglobose, 3-6-1 obate, the endosperm strongly
ruminate; cotyledons narrowly oblong.

106 FIELDIANA: BOTANY, VOLUME 24

Five or fewer species in Mexico and Central America. The fol-
lowing are the only species of the genus, except for A. macrocarpum
Britt. & Small, which is known only in cultivation, in Costa Rica
and Puerto Rico. It is distinguished by having very large, orbicular
fruiting bracts, and is otherwise like A. leptopus.

Outer sepals broadly ovate, not cordate at the base, at least in anthesis; leaf

blades abruptly decurrent at the base upon the petiole A. guatemalense.

Outer sepals rounded-ovate or suborbicular, conspicuously cordate at the base.
Leaf blades abruptly contracted at the base, not decurrent upon the petiole.

A. leptopus.

Leaf blades abruptly contracted at the base and decurrent upon the petiole.
Sepals yellowish or greenish, in fruit usually longer than broad, acute or

acutish; plants usually almost glabrous. A. flavescens.

Sepals bright rose, in fruit nearly or quite as broad as long, obtuse or rounded
at the apex and apiculate; plants usually copiously pubescent.

A. cinerascens.

Antigonon cinerascens Mart. & Gal. Bull. Acad. Brux. 10,
pt. 1:354. 1843.

Moist or dry thickets, 250-1,300 meters; Zacapa; Jutiapa.
Southern Mexico; Honduras; Salvador.

A large vine, densely pubescent or sometimes glabrate, the stems angulate;
leaves slender-petiolate, the blades ovate-cordate or broadly deltoid-cordate,
mostly 6-9 cm. long, obtuse to acuminate, with a broad shallow basal sinus,
abruptly and narrowly decurrent upon the petiole; racemes paniculate, the panicles
small or large; flowers dull dirty pink or purplish pink, the outer sepals in anthesis
about 8 mm. long, in fruit rounded-cordate and 1.5 cm. long; achene brown,
lustrous, almost 1 cm. long.

Called "bejuco de colacion" in Salvador. This is quite as hand-
some as the better known A. leptopus.

Antigonon flavescens Wats. Proc. Amer. Acad. 22: 446. 1887.
Moist or dry thickets of the Oriente, about 400 meters; Chiqui-
mula. Jalisco to Oaxaca.

A small or large vine, the stems wholly or chiefly herbaceous (as in other
species), angulate, puberulent or glabrate, green; leaves on rather short, slender
petioles, the blades ovate-cordate, mostly 6-11 cm. long and 3-6.5 cm. wide, long-
acuminate, deeply and openly cordate at the base, green and glabrate, the margins
somewhat undulate; racemes lax, rather few-flowered, the lower on long slender
pedicels; sepals yellowish or greenish white, green and accrescent in age and then
about 1.5 cm. long, acute or subacute, reticulate- veined; achene 8 mm. long, ovoid,
brownish, glabrous, acuminate.

This seems to be a valid species rather than a mere color form.
The plant is, of course, much less ornamental than other species.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 107

Antigonon guatemalense ("guatimalense") Meisn. in DC.
Prodr. 14: 184. 1856. Polygonum grandiflorum Bertol. Fl. Guat. 412.
1840, not P. grandiflorum Willd. 1799. A. grandiflorum Robinson,
Proc. Amer. Acad. 44: 513. 1909.

Dry or moist thickets, 1,300 meters or less, chiefly in the foot-
hills and on the Pacific plains; El Progreso; Zacapa; Chiquimula;
Jutiapa; Santa Rosa; Escuintla (type from Escuintla, Velasquez);
Sacatepe"quez; Suchitepe'quez ; Huehuetenango; doubtless in all the
Pacific coast departments. Southern Mexico; Salvador; Nicaragua.

Usually a large vine, the stems angulate, puberulent; leaves on rather short
petioles, broadly ovate-cordate or rounded-cordate, mostly 5-9 cm. long, rounded
to short-acuminate at the apex, with a deep sinus at the base, usually more or less
decurrent upon the petiole, densely pubescent beneath or sometimes glabrate;
racemes lax or dense, paniculate, the flowers very showy, mostly bright rose-pink;
sepals puberulent, the outer ones in anthesis 8-10 mm. long, in fruit suborbicular
and much larger, rounded or obtuse at the apex, reticulate- veined; achene about
1 cm. long, brown, lustrous.

Known in Salvador by the names "eolation," "confite," and
"San Andre's." The plant is plentiful in many parts of the Pacific
plains, where it often forms dense tangles over thickets or hedges
and affords fine displays of handsome color.

Antigonon leptopus Hook. & Arn. Bot. Beechey Voy. 308.
pi. 69. 1839-40. A. cordatum Mart. & Gal. Bull. Acad. Brux. 10,
pt. 1: 14. 1843. Confite; Flor de San Miguel; San Diego.

Damp thickets and hedges, probably naturalized but perhaps
native in the Pacific coast; Izabal; Alta Verapaz; Retalhuleu; cul-
tivated for ornament through most of the warmer parts of Guate-
mala and in the central and western mountains up to 1,500 meters
or more. Mexico.

A large or small vine; leaves ovate-cordate or most often broadly deltoid-
cordate, mostly 5-10 cm. long, acute or acuminate, shallowly and openly cordate
at the base, pubescent beneath or often almost glabrous; outer sepals rose-pink,
at first 8-10 mm. long, in fruit much larger and usually rounded-ovate, about 1.5
cm. long, reticulate-veined.

This is the only species noted in cultivation in Guatemala, and
in most regions of the country where it is planted it is apparently
introduced, although it may well be native along the Pacific coast.
It is common in western Mexico, but there could have been no reason
for importing it into Guatemala, since it is in no way superior to
local species. The Maya name reported from Yucatan is "chaclo-
macal." In Honduras the vine is called "bellisima"; in Salvador

108 FIELDIANA: BOTANY, VOLUME 24

"eolation" and "confite rojo." In Florida, where it is much planted,
it is known as "Confederate vine." Its habit and general appearance
are somewhat suggestive of Bougainvillea. It is a good ornamental
vine because the flowers retain their color for a long time. The most
remarkable character of the vines of this genus is found in the ten-
drils, which are borne in the inflorescences, a most unusual place
for tendrils. The roots bear tubers that usually are small, but some-
times weigh as much as fifteen pounds. They are said to be edible
and to have an agreeable nutlike flavor.

COCCOLOBA L.

Trees or shrubs, usually glabrous or nearly so; ocreae coriaceous-membrana-
ceous, cylindric, not ciliate, truncate, deciduous; leaves persistent or deciduous,
mostly coriaceous; flowers perfect, in spike-like, axillary or subterminal, simple or
rarely branched racemes, the bracts ocreiform, subtending several flowers, the
pedicels short or elongate, articulate at the apex; calyx green or whitish, small,
the 5 segments subequal, united at the base, either the tube or the lobes accrescent
with age and enclosing the fruit, usually becoming much thickened and succulent;
stamens 8, equal; achene subtrigonous-globose, small or large.

Probably more than 150 species, all in tropical America. A few
additional ones are known from southern Central America.

Leaves peltate C. acapulcensis.

Leaves not peltate.

Leaves densely hirsute on both surfaces with long spreading hairs. . .C. hirsuta.
Leaves glabrous or merely puberulent, at least not hirsute, sometimes pilose

along the nerves and veins.
Flower spikes paniculate.

Leaves 3-6 cm. wide C. Gentlei.

Leaves mostly 10-15 cm. wide or larger.

Leaf blades cordate or subcordate at the base C. belizensis.

Leaf blades cuneately narrowed to the base, the base acute or obtuse.

C. Tuerckheimii.
Flower spikes simple, not branched.

Leaves all or mostly broadly rounded or very obtuse at the apex.

Rachis of the inflorescence glabrous. Leaf blades longer than broad,

obtuse or acute at the base C. corozalensis.

Rachis of the inflorescence puberulent or hirtellous.

Leaves hirtellous or pilose beneath, at least along the nerves, mostly
large and suborbicular, obtuse to broadly rounded or sometimes
subcordate at the base, with elevated and reticulate venation

beneath C. caracasana.

Leaves glabrous or merely puberulent beneath.

Flower spikes very dense and crowded, the pedicels none or very
short.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 109

Leaves broadest above the middle, not cordate at the base, mostly

3-6 cm. wide C. floribunda.

Leaves broadest at or usually below the middle, often cordate or

subcordate at the base, often much wider.
Leaves mostly cordate at the base and 7-9 cm. wide . . C. spicata.
Leaves mostly rounded at the base and 4-6 cm. wide.

C. mayana.

Flower spikes lax and open, the flowers not crowded, the pedicels
conspicuous, often elongate.

Leaf blades mostly obovate, broadest above the middle, mostly

3.5-5.5 cm. wide, narrowed to the base C. reflexiflora.

Leaf blades mostly orbicular or nearly so.

Leaf blades all or mostly broader than long, conspicuously

cordate at the base C. Uvifera.

Leaf blades fully as long as broad, rounded at the base.

C. Lundellii.

Leaves acute or acuminate at the apex, or at least subacute.
Leaf blades conspicuously cordate at the base.

Leaves ferruginous-pilose beneath along the costa C. montana.

Leaves glabrous beneath.

Racemes mostly 4 cm. long or less, the slender pedicels much longer

than the flowers; leaves relatively thin C. Browniana.

Racemes 8-12 cm. long or larger, the pedicels shorter than the

flowers; leaves coriaceous.
Leaves mostly 7-9 cm. wide, mostly rather deeply cordate at the

base • C. spicata.

Leaves mostly 4-6 cm. wide, subcordate at the base . . C. mayana.
Leaves acuminate to very obtuse or rounded at the base.
Rachis of the inflorescence glabrous.

Flowers sessile or subsessile, the pedicels 1 mm. long or less.

C. cozumelensis.

Flowers on slender elongate pedicels C. laurifolia.

Rachis of the inflorescence puberulent or pilose.

Leaves broadest above the middle, mostly oblanceolate-oblong or

obovate-oblong C. floribunda.

Leaves broadest at or below the middle.

Flowers conspicuously pedicellate, twice as long as the ocreolae or

longer, at least in full anthesis C. escuintlensis.

Flowers sessile or subsessile.

Leaf blades mostly acute at the base, barbate beneath in the

axils of the nerves C. acuminata.

Leaf blades rounded or very obtuse at the base, not barbate

beneath.
Leaves long-acuminate, usually gradually so.

Lateral nerves of the leaves about 14 pairs, obscure, not or
scarcely elevated; racemes mostly 7-13 cm. long.

C. Steyermarkii.

Lateral nerves of the leaves 6-7 pairs, conspicuous and
elevated beneath; racemes mostly 2-4 cm. long.

C. Schippii.

Leaves rounded or obtuse at the apex and abruptly short-
acuminate or apiculate C. Schiedeana.

110 FIELDIANA: BOTANY, VOLUME 24

Coccoloba acapulcensis Standl. Proc. Biol. Soc. Wash. 33:
66. 1920.

Moist or dry, often rocky, brushy hillsides, 600-1,400 meters;
El Progreso; Jutiapa; Huehuetenango. Guerrero, Mexico, the type
from Acapulco.

A shrub 2 meters tall, or sometimes a tree of 5 meters, glabrous throughout,
the branches rather slender, dark ferruginous; leaves on slender petioles 2.5-3 cm.
long, the blades peltate far above the base, orbicular to rounded-ovate, 6-11 cm.
wide, rounded to subacuminate at the apex, broadly rounded or emarginate at
the base; pedicels fasciculate, the racemes stiff, 8 cm. long or less, rather dense;
fruit obovoid, about 2.5 cm. long and 1.5 cm. in diameter.

A remarkable plant because of the peltate leaves, unique among
at least the North American species of the genus. The fruits are
abnormally large.

Coccoloba acuminata HBK. Nov. Gen. & Sp. 2: 141. 1817.

Moist or wet thickets at or little above sea level; Izabal (near
Bananera, Steyermark 38986). Honduras to Panama; Colombia.

Usually a shrub of 2-3 meters, the branchlets very slender, puberulent or
glabrate; ocreae 1 cm. long; leaves short-petiolate, oblong, lance-oblong, or elliptic-
oblong, 10-20 cm. long, 4-6 cm. wide, long-acuminate, acute or obtuse at the base,
glabrous and lustrous above, densely barbate beneath in the nerve axils; racemes
very long and slender, often recurved and pendulous, dense or lax, the rachis
hirtellous or puberulent; pedicels mostly shorter than the ocreolae; fruit sub-
globose, obtusely trigonous, 6 mm. long, bright red; perianth tube accrescent and
enclosing the achene.

Apparently scarce in Guatemala but very common in many parts
of the Atlantic coast of Central America. The shrub is a showy and
ornamental one when in mature fruit. Known in Honduras by the
names "rabo de leon" and "tapatamal."

Coccoloba belizensis Standl. Trop. Woods 16: 38. 1928. Uva
de monte (Pete"n).

Wet forest or thickets, 900 meters or less; Pete"n; Alta Verapaz;
Izabal. British Honduras; Atlantic coast of Honduras.

A small or large tree, sometimes 25 meters high with a trunk 45 cm. in diame-
ter, the thick branchlets densely puberulent; ocreae large and conspicuous, ferru-
ginous-puberulent or tomentulose; leaves large, thick-coriaceous, short-petiolate,
the blades broadly oval to broadly oblong or obovate, often 30 cm. long and 24
•cm. wide, but many of the leaves smaller, usually very obtuse or rounded at the
apex and abruptly pointed, sometimes acute, shallowly cordate at the base or
merely obtuse, puberulent or glabrate beneath, the lateral nerves coarse and
prominent, glabrous on the upper surface; flower spikes few or numerous, panicu-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 111

late, 20 cm. long or less, usually very dense, the stout rachis densely hirtellous or
puberulent, the flowers sessile or nearly so, whitish, slightly odorous; fruits sub-
globose, 5 mm. in diameter when dry.

Called "uva" and "bul" (an Indian name) in Honduras, and
"wild grape" in British Honduras.

Coccoloba Browniana Standl. Trop. Woods 10: 4. 1927.
C. cardiophylla Standl. Field Mus. Bot. 8: 8. 1930 (type from
Yucatan).

Northern British Honduras, and to be expected in Pete"n; Hon-
duras, the type from Olanchito.

A tree about 7 meters tall, the branches dark ferruginous or sometimes in age
whitish, glabrous; ocreae sheathing, 8-10 mm. long, deciduous; leaves slender-
petiolate, usually rounded-ovate, 5.5-11 cm. long, 4.5-8 cm. wide, commonly
rounded or obtuse at the apex and abruptly short-acuminate, shallowly cordate
at the base, rather thin, glabrous, paler beneath; racemes short, in an thesis mostly
2-4 cm. long, the rachis glabrous; pedicels short in flower but in fruit 5 mm. long
or more, stiff, divaricate; calyx tube accrescent and enclosing the fruit; fruit sub-
globose, 1.5 cm. long and almost as broad.

Called "tolondron" in Honduras.

Coccoloba caracasana Meisn. in DC. Prodr. 14: 157. 1856.
Papaturro bianco; Papaturro.

Moist thickets or forest of plains and hillsides, often in dry
regions, sometimes along roadsides, 600 meters or less; Jutiapa;
Santa Rosa; Escuintla; Suchitepe"quez; Retalhuleu; doubtless also
in the other Pacific coast departments. Salvador to Panama;
Colombia and Venezuela.

A small or medium-sized tree, usually 6-15 meters high, the crown dense and
rounded, the trunk short, the branchlets hirtellous or puberulent or glabrate;
ocreae 1.5-2.5 cm. long; leaves short-petiolate, generally suborbicular, not very
thick, 10-30 cm. long and almost as wide, broadly rounded or emarginate at the
apex, rounded or shallowly cordate at the base, glabrous above or nearly so,
beneath short-pilose or puberulent, in age sometimes glabrate, the veins prominent
and closely reticulate; racemes simple, usually longer than the leaves, the rachis
puberulent or hirtellous; flowers green or greenish white, sweet-scented, the pedi-
cels shorter than the ocreolae, often almost none, the racemes usually dense;
lobes of the perianth accrescent and enclosing the fruit, this 5 mm. long when dry,
in fruit much larger, white, juicy.

Sometimes called "paparron" and "papalon" in Salvador. The
usual name for this and related species along the Pacific coast of
Central America is "papaturro," a term utilized in the name of an
aldea, Papaturro, of Jutiapa. The tree is plentiful at many places

112 FIELDIANA: BOTANY, VOLUME 24

on the Pacific plains, where it is rather attractive because of the
very dense, rounded crown of large and handsome leaves. The
juicy white fruits have an agreeable acidulous flavor and often are
eaten by people. Their weight at maturity causes the spikes to
become pendent.

Coccoloba corozalensis Lundell, Bull. Torrey Club 66: 587.
/. 2. 1939.

Known only from British Honduras, the type from Xiabe,
Corozal District, Lundell 4908; to be expected in Pete"n.

A tree with a trunk 10-15 cm. in diameter, the branchlets glabrous; ocreae
5-9 mm. long; leaves glabrous, slender-petiolate, oblong-elliptic to broadly
obovate, 5-12 cm. long, 2.5-7.5 cm. wide, rounded or very obtuse at the apex,
rounded to subacute at the base; racemes simple, 7-14 cm. long, the flowers rather
remote, the rachis glabrous, the pedicels shorter than the ocreolae; fruit black-
purple, 7-9 mm. long.

Known in British Honduras by the names "uva cimarron,"
"pigeon plum," and "wild grape."

Coccoloba cozumelensis Hemsl. Biol. Centr. Amer. Bot. 4: 108.
1887 (type from Cozumel Island, Yucatan). C. yucatana Lindau,
Bot. Jahrb. 13: 190. 1890 (type from Yucatan).

Moist or wet thickets, Pete"n. British Honduras; Yucatan;
Campeche.

A shrub or small tree, 9 meters high or less, glabrous throughout or nearly so,
the branchlets slender, ferruginous or blackish; leaves short-petiolate, mostly
ovate-oblong or lance-oblong, 3-10 cm. long, rather thin, acute or acuminate, often
with an obtuse tip, obtuse at the base, usually barbate beneath in the axils of the
nerves; racemes slender, simple, mostly 13 cm. long or less, often recurved above
the middle, rather densely flowered, the flowers pale green, the pedicels very short
or almost none; perianth tube accrescent and enclosing the fruit, this 4-5 mm.
long when dry, globose-ovoid.

Called "wild grape" and "manzanilla" in British Honduras.

Coccoloba escuintlensis Lundell, Phytologia 1: 213. 1937.
Cacho de ternero (San Marcos).

Moist or dry forest or thickets, often in second growth, sometimes
in pastures or cafetales, Pacific slope, ascending from sea level to
1,400 meters, mostly at 900 meters or less; Escuintla; Retalhuleu;
Quezaltenango; San Marcos; doubtless also in Suchitepe"quez.
Chiapas, the type from Escuintla, Matuda 413.

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 113

A small or large tree, sometimes 25 meters high, with a trunk 65 cm. in diame-
ter, usually smaller, the bark rather rough, light brown, the branchlets glabrous or
essentially so; leaves on rather short petioles, mostly lance-oblong to ovate-lanceo-
late, 10-25 cm. long and 4-9 cm. wide, or in young plants often larger, long-
acuminate or acute, rarely subobtuse, rounded or obtuse at the base, rather thick,
glabrous; racemes simple, mostly 6-14 cm. long, the rachis minutely puberulent;
pedicels about twice as long as the ocreolae; perianth lobes accrescent and enclosing
the fruit; dry fruit 7-8 mm. long, subglobose, dull dark red when fresh and larger.

The leaves probably are deciduous at the end of the dry season.
The young leaves are often coppery red.

Coccoloba floribunda (Benth.) Lindau, Bot. Jahrb. 13: 217.
1890. Campderia floribunda Benth. Bot. Voy. Sulph. 159. pi. 52.
1844. Papaturro.

Moist or dry thickets or forest, often in coastal thickets, 850
meters or less, chiefly on the Pacific plains; Zacapa; Chiquimula;
Jutiapa; Santa Rosa; Guatemala; Retalhuleu. Southern Mexico;
Salvador to Costa Rica, the type from Tigre Island, Honduras;
Colombia to Brazil.

A densely branched shrub or tree, sometimes 9 meters high, with a broad
spreading crown, the low trunk often gnarled and twisted, sometimes a meter in
diameter, the bark light or medium brown, the inner bark darker brown, glabrous
throughout or nearly so; ocreae 8 mm. long or less; leaves on very short petioles,
obovate or obovate-oblong, 5-12 cm. long, rounded to subacute at the apex, some-
what narrowed to the subacute to rounded base, coriaceous, the veins prominulous
and reticulate; racemes mostly 4-10 cm. long, very dense, the rachis minutely
puberulent or glabrate, the pedicels shorter than the ocreolae; perianth lobes
accrescent and enclosing the fruit, this ovoid-globose, 5-6 mm. long in the dry
state, bluish black or purplish red at maturity, sometimes dull dark red before
maturity.

In Salvador known by the names "papaturro," "iron," "irire,"
and "juril." The fruit is juicy and edible. The sapwood is pale
yellow, the heartwood brownish. This, like some other members
of the genus, makes a good shade tree and often is seen about
dwellings on the Pacific plains.

Coccoloba Gentlei Lundell, Bull. Torrey Club 66: 591. /. 3.
1939.

Izabal (sterile material collected near Escobas is probably
referable here). British Honduras, the type collected along the
Belize-Sibun River road, Belize District, Percy H. Gentle 56.

A small tree, glabrous or nearly so; ocreae 12 mm. long; leaves short-petiolate,
coriaceous, oblong-ovate or ovate, 7-16 cm. long, 3.5-8 cm. wide, subacute at the
apex, usually rounded and often unequal at the base, lustrous above, somewhat

114 FIELDIANA: BOTANY, VOLUME 24

paler beneath, the lateral nerves conspicuous beneath, the veins prominulous and
reticulate; racemes few, paniculate, 16 cm. long or less, the rachis glabrous, the
flowers rather distant, the flowers subsessile.

Coccoloba hirsuta Standl. Field Mus. Bot. 4: 303. 1929.

Wet forest, at or near sea level; Izabal. Atlantic coast of Hon-
duras, the type collected in Lancetilla Valley, near Tela.

A shrub or small tree, the branchlets very thick, densely hirsute with fulvous
hairs; ocreae 1-1.5 cm. long, hirsute; petioles stout, 4-8 cm. long, fulvous-hirsute;
leaf blades oblong-oval or elliptic-obovate, about 40-50 cm. long and 17-28 cm.
wide, short-acuminate, rounded or shallowly cordate at the base, thin, green
above and hirsute, somewhat paler beneath and fulvous-hirsute, the lateral nerves
about 11 pairs, conspicuous.

The species is known only from sterile material but is easily
recognized by the abundant pubescence of long spreading hairs.
It seems rather probable that it will be found to have panicled
racemes, and to be closely related to C. Tuerckheimii. In Honduras
the tree is called "uva" and "uva de monte."

Coccoloba laurifolia Jacq. PI. Hort. Schoenbr. 3: 9. pi. 267.
1798. C. lancifolia Lundell, Bull. Torrey Club 66: 593. 1939 (type
from Jacinto Hills, Toledo District, British Honduras, W. A. Schipp
1200).

Wet forest or thickets, 500 meters or less; Alta Verapaz; Izabal.
British Honduras; southern Florida; West Indies; Venezuela.

A large shrub or a small tree, sometimes 10 meters tall with a trunk 12 cm. in
diameter, glabrous throughout; ocreae 3-6 mm. long; leaves short-petiolate, sub-
coriaceous, lance-oblong to oblong-ovate, 5-13 cm. long, 2-6.5 cm. wide, acuminate
to subobtuse, subacute to almost rounded at the base and often unequal; racemes
stout and stiff, 7 cm. long or less, rather lax, the stout fruiting pedicels divaricate,
5 mm. long or less; fruit ovoid, 1 cm. long, yellow or at maturity blue-black.

Coccoloba Lundellii Standl. Field Mus. Bot. 8: 138. 1930.
C. suborbicularis Lundell, Lloydia 2: 84. 1939 (type from Stann
Creek District, Stann Creek Railway, British Honduras, Percy H.
Gentle 2687).

Type from Honey Camp, British Honduras, Lundell 649; to be
expected in Pete"n.

A shrub or small tree, the branchlets pale, glabrous; ocreae 5-7 mm. long;
leaves short-petiolate, coriaceous, usually orbicular or nearly so, 8-19 cm. long
and almost or fully as wide, broadly rounded at the apex, rounded at the base and
often emarginate or subcordate, somewhat unequal, glabrous above, very minutely
puberulent beneath or almost glabrous; racemes simple, rather lax, 18-27 cm.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 115

long, the rachis minutely puberulent; pedicels in fruit divaricate, stout, 2-2.5
mm. long; perianth tube accrescent and enclosing the fruit, this globose-ovoid, as
much as 1 cm. long.

Called "wild grape" in British Honduras.

Coccoloba mayana Lundell, Bull. Torrey Club 64: 547. 1937.

Moist or rather dry, often rocky thickets, usually along streams
or about waterholes, 700 meters or less; Pete"n; Izabal; Escuintla;
Suchitepe"quez. Veracruz to Chiapas; British Honduras.

A small tree, usually 9 meters high or less, sometimes 15 meters tall, the young
branchlets puberulent or glabrate; ocreae 4-8 mm. long, puberulent; leaves short-
petiolate, deciduous, rounded-ovate to ovate-oval or ovate-oblong, 6-12 cm. long,
3-7 cm. wide, subcoriaceous, acute to rounded and apiculate at the apex, rounded
or shallowly cordate at the base, glabrous or essentially so; racemes simple, 8-25
cm. long, open, the rachis minutely puberulent, slender, often curved, the nodes
1-2-flowered; pedicels short, about equaling the ocreolae; fruit ovoid, 8 mm. long.

Coccoloba montana Standl. Journ. Wash. Acad. Sci. 13: 368.
1923. Papaturro.

Moist lowland forest of the Pacific slope, 900-1,400 meters;
Quezaltenango; San Marcos. Salvador, the type from Finca Colima,
Ahuachapan.

A shrub or tree, sometimes 7 meters tall, the young branches pale, glabrous;
ocreae brown, glabrous, 6-7 mm. long; petioles stout, glabrous, 1-4 cm. long; leaf
blades ovate or oblong-ovate, 10-35 cm. long, 6-16 cm. wide, acuminate or long-
acuminate, at the base rounded on one side and semicordate on the other, glabrous
above, beneath brownish-pilose along the costa, especially in the nerve axils,
elsewhere glabrous or nearly so, the lateral nerves prominent and conspicuous
beneath, the veins prominulous and closely reticulate.

This "species" is known only from sterile material, and the
Guatemalan specimens have been determined by comparison with
the original description. It is suspected that the material referred
here represents juvenile foliage or leaves from vigorous sterile
branches of possibly C. escuintlensis, or perhaps of one of the other
species listed here.

Coccoloba reflexiflora Standl. Field Mus. Bot. 4: 203. 1929.

Pete'n (region of Uaxactun). Campeche; Yucatan; British
Honduras.

A large shrub or a small tree, glabrous or nearly so, the trunk 5-8 cm. in
diameter; ocreae 4-12 mm. long; leaves on very short (2.5-5 mm.) petioles, coria-
ceous, rounded-obovate to oblong-obovate, mostly 6-8 cm. long and 2.5-5 cm. wide,

116 FIELDIANA: BOTANY, VOLUME 24

broadly rounded to obtuse at the apex, somewhat narrowed to the obtuse or nar-
rowly rounded base, barbate beneath along the costa; racemes simple, mostly
8-12 cm. long, often reflexed or recurved, rather lax and many-flowered, the rachis
minutely puberulent; pedicels 1-1.5 mm. long, divaricate or, especially in anthesis,
reflexed; fruits black at maturity, when dry ellipsoid and 6 mm. long.

Coccoloba Schiedeana Lindau, Bot. Jahrb. 13: 187. 1890.
C. hondurensis Lundell, Bull. Torrey Club 66: 591. 1939 (type from
Little Cocquericot, Belize River, British Honduras, Lundell 3996).
Papaturro; Garner o.

Dry or moist thickets or forest, often along rocky stream banks,
900 meters or less, mostly near sea level, sometimes in coastal
thickets; Pete"n; Alta Verapaz; Izabal; Santa Rosa; Escuintla;
Retalhuleu; Quezaltenango; San Marcos; Huehuetenango. Southern
Mexico; British Honduras; Honduras; Salvador.

A small or medium-sized tree, sometimes 15 meters high but commonly lower,
the bark dark, rough, the crown spreading, the trunk often crooked, sometimes
60 cm. or more in diameter, the branchlets puberulent or glabrate; ocreae 5-9
mm. long; leaves short-petiolate, mostly broadly oval to oblong-elliptic, 10-20
cm. long and 6-13 cm. wide or sometimes larger, often coriaceous, generally
rounded or very obtuse at the apex and abruptly short-acuminate or at least
protracted, obtuse to cordate at the base, glabrous; racemes simple, usually equal-
ing or longer than the leaves, the rachis minutely puberulent; pedicels short,
usually shorter than the ocreolae and often almost obsolete, the racemes often
very densely flowered, often recurved and pendulous, the flowers whitish; fruit
ovoid or subglobose, almost 1 cm. long or shorter; perianth tube accrescent and
enclosing the fruit.

Called "wild grape" and "iril" in British Honduras. The
material referred here is slightly variable in foliage, but not unreason-
ably so and we find no basis for dividing it into two or more species.
Called "uvero" in Oaxaca. The sapwood is cream-colored, the heart-
wood light or medium brown. The branches in this and related
species are sometimes hollowed and inhabited by ants that bite
severely.

Coccoloba Schippii Lundell, Bull. Torrey Club 66: 594. 1939.

Known only from the type, Schipp S687, collected at Camp 31
on the boundary between Pete"n and British Honduras, 630 meters.

A tree of 9 meters, the trunk 15 cm. in diameter, the slender branchlets
glabrous; ocreae barbate at the apex; leaves glabrous, rather thin, on slender
petioles 9-14 mm. long, lance-oblong, 8-12.5 cm. long, 3-5 cm. wide, acuminate,
obtuse or narrowly rounded at the base and slightly oblique; spikes simple, the
young ones 2.5-4.5 cm. long, the rachis hirtellous, the flowers cream-colored,
rather crowded, the nodes mostly 1-flowered, the pedicels very short.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 117

Coccoloba spicata Lundell, Bull. Torrey Club 66: 594. /. 4. 1939.

Dry upland forest, or about lake borders, 300 meters or less;
Pete"n; Alta Verapaz. British Honduras; Yucatan; Quintana Roo;
Campeche.

A tree of 5-15 meters, the trunk 10-20 cm. or more in diameter, the branch-
lets glabrous; ocreae 5-9 mm. long, at first rufous-hirsute but soon glabrate;
leaves on stout petioles 1-2.5 cm. long, coriaceous, ovate-oblong to rounded-oval,
7-15 cm. long, 4-10 cm. wide, obtuse to broadly rounded at the apex and often
apiculate, rounded to cordate at the base, barbate beneath in the axils of the nerves,
elsewhere glabrous or nearly so, the leaves of sterile branches sometimes as much
as 29 cm. long and 21 cm. wide, the lateral nerves elevated and conspicuous
beneath; spikes simple, 9-25 cm. long, densely flowered, often recurved or pendu-
lous, the rachis minutely puberulent, the flowers sessile or nearly so; perianth tube
accrescent and enclosing the fruit, this globose-ovoid or subglobose, when dry
about 7 mm. long.

The Maya names of Yucatan are "boob," "bob," and "bobche";
called "wild grape" in British Honduras, and "bochiche" (a Maya
name) in Campeche. The larger leaves are reported to be used in
Yucatan for wrapping certain dulces.

Coccoloba Steyermarkii Standl. Field Mus. Bot. 22: 138. 1940.

Known only from the type, Steyermark 39533, Dept. Izabal, Rio
Dulce, 2-4 miles west of Livingston, at sea level.

A tree, the branchlets glabrous, ochraceous; ocreae 9 mm. long, minutely
puberulent; leaves on stout petioles 1.5 cm. long, coriaceous, very narrowly lance-
oblong, 14-19 cm. long, 4-5 cm. wide, narrowly attenuate-acuminate, obtuse or
narrowly rounded at the base, glabrous, somewhat paler beneath, the ultimate
venation prominulous and closely reticulate on both surfaces; spikes simple, 4.5-8
cm. long, dense and many-flowered, the rachis stout, densely puberulent, the
stout pedicels twice as long as the ocreolae.

The flowers are pale green.

Coccoloba Tuerckheimii Donn. Smith, Bot. Gaz. 37: 213. 1904.
Irayol de montana; Pojchic (Alta Verapaz).

Wet forest or thickets, sometimes on limestone, 1,100 meters or
less; Alta Verapaz (type from Cubilgiiitz, Tuerckheim 8493); Izabal.
British Honduras; Honduras; Costa Rica; Panama.

A tree, sometimes 15-20 meters high, the trunk rarely 120 cm. in diameter,
the bark very dark brown, corky, checkered and flaking, the branchlets stout,
puberulent or glabrate; ocreae about 3 cm. long, lax; leaves short-petiolate, sub-
coriaceous or thin, oblong-obovate to broadly obovate-elliptic, mostly 15-35 cm.
long and 8-18 cm. wide, rounded or obtuse and abruptly short-acuminate at the
apex, narrowed to the acute or obtuse base, glabrous above, the veins often

118 FIELDIANA: BOTANY, VOLUME 24

depressed and the blades somewhat bullate, puberulent beneath, the nerves
slender, elevated; racemes forming a large sessile panicle equaling or shorter than
the leaves; pedicels solitary, twice as long as the ocreolae or longer; flowers greenish
white; fruit ovoid, 1 cm. long.

Called "wild grape" in British Honduras and "uva" and "al-
mendro de monte" in Honduras.

Coccoloba Uvifera (L.) Jacq. Enum. PI. Carib. 19. 1760.
Polygonum Uvifera L. Sp. PL 365. 1753. Uva.

Thickets along the edges of sea beaches, Izabal. Southern
Florida; Mexico; British Honduras to Panama, along the Atlantic
coast; West Indies; northern South America.

A densely branched shrub or tree, usually less than 10 meters high, the trunk
rarely a meter in diameter, the bark thin, smooth, brown; ocreae 1 cm. long;
leaves short-petiolate, thick-coriaceous and rigid, orbicular or transverse-oval,
mostly 8-20 cm. wide, rounded or truncate at the apex, often emarginate, cordate
at the base, minutely puberulent or glabrate beneath, often red or tinged with red
or purple, the nerves and veins not very conspicuous; racemes simple, equaling
or longer than the leaves, the rachis minutely puberulent; flowers whitish, fra-
grant, the pedicels twice as long as the ocreolae; fruit ovoid, 2 cm. long or less,
purplish.

Called "grape" in British Honduras; "niiche" (Yucatan, Maya);
"uva," "uva de la playa," "papaturro" (Honduras). The wood is
red or dark brown tinged with red, sometimes violet or streaked,
the sapwood pinkish; odorless, with slightly astringent taste, its
alkaline extract ruby-red ; hard, heavy, compact, its specific gravity
about 0.96; of irregular grain, fine- textured, fairly easy to work,
takes a high polish, appears durable; strong but brittle. The usual
English name is "sea-grape." When cut, the bark yields an astrin-
gent red sap which is the source of West Indian kino. This product,
known also as gum kino, American kino, American extract of
rhatany, and false rhatany extract, was formerly an article of trade,
but the commerical kino now is obtained from West Africa and the
East Indies. The wood has been employed locally for cabinetwork
and is burned for charcoal. The juicy fruit is edible, having an
acidulous and somewhat astringent flavor. In the West Indies it has
been fermented with sugar to produce an alcoholic beverage. In
Florida it is much used for making jelly. Oviedo records that in
early colonial days the large stiff leaves were used by the Spaniards
as a substitute for writing paper, the characters being impressed
upon the surface with a pin or other sharp-pointed implement.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 119

GYMNOPODIUM Rolfe
Reference: S. F. Blake, Bull. Torrey Club 48: 83-84. 1921.

Shrubs or small trees with divaricate, usually flexuous or crooked branches;
leaves mostly fasciculate on short spurs, short-petiolate, membranaceous or sub-
coriaceous; flowers small, green, perfect, slender-pedicellate, fasciculate, the fasci-
cles forming short, simple or branched racemes; perianth segments 6, the 3 outer
ones larger, carinate, not winged, the 3 inner ones smaller, plane, erect; stamens 9,
inserted at the base of the perianth, the filaments filiform, the anthers ovate;
ovary glabrous, the styles short, filiform, the stigmas capitate; ovule erect, sub-
sessile; achene acutely trigonous, included in the accrescent and closed perianth;
seed trigonous, with a large embryo, the cotyledons orbicular.

One other species is known, on the north coast of Yucatan. So
far as known at present, the genus is confined to the Yucatan
Peninsula.

Gymnopodium floribundum Rolfe in Hook. Icon. 27: pi.
2699. 1901. Millspaughia leiophylla Blake, Contr. Gray Herb. 52: 62.
1917 (type from Manatee Lagoon, British Honduras, M. E. Peck
320). Crucito (British Honduras).

Thickets or wooded swamps, Pete*n; British Honduras (type from
Manatee, E. J. F. Campbell 60). Tabasco; Campeche.

A shrub or small tree 3 meters tall or more, the trunk to 8 cm. in diameter,
the bark brown, shredded, the branchlets sparsely pilose; leaves on very short
petioles, narrowly cuneate-oblong to oblong or elliptic-oblong, mostly 2-3.5 cm.
long, obtuse, glabrous or with a few hairs along the nerves, the veins prominent
and reticulate beneath; ocreae very small; racemes mostly terminal, sometimes
7.5 cm. long but usually shorter; outer sepals ovate or rounded-ovate, acute or
subacute, in fruit 1 cm. long, greenish, reticulate-veined; achene 6 mm. long.

Called "bastard logwood" in British Honduras.

Gymnopodium floribundum var. antigonoides (Robinson)
Standl. & Steyerm. Field Mus. Bot. 23: 5. 1943. Millspaughia
antigonoides Robinson, Bot. Jahrb. 36, Beibl. 80: 14. 1905.

At 500-800 meters; Huehuetenango (between Nenton and Mira-
mar, Steyermark 51459). Chiapas and Yucatan to British Hon-
duras; type from Progreso, Yucatan.

Differing from the typical form of the species in having the leaves sparsely
or densely pubescent beneath, at least when young.

Although maintained as a distinct species by Blake, this seems
to differ from G. floribundum only in amount of pubescence. Prob-
ably in no species of the genus are the leaves always and completely
glabrous, as indicated in his key. It is somewhat questionable

120 FIELD IANA: BOTANY, VOLUME 24

whether his third species, G. ovatifolium (Robinson) Blake, of Yuca-
tan, is more than a form of G. floribundum. Maya names reported
from Yucatan for the variety are "tzitzilche" and "zactzitzilche."
The wood is said to give a good quality of charcoal, and the flowers
to yield much honey of excellent flavor.

MUEHLENBECKIA Meisner

Plants suffrutescent or shrubby, often scandent; leaves alternate, petiolate,
sometimes small and orbicular, frequently cordate, deltoid, or sagittate, varying
to linear, sometimes none; ocreae small, often almost obsolete; flowers small,
fasciculate within the ocreolae, the fascicles axillary and solitary or in terminal
or axillary, simple or branched spikes or racemes, polygamo-subdioecious; perianth
deeply 5-fid, the lobes subequal or the 3 outer ones slightly larger, in fruit persistent
and usually fleshy; stamens 8, inserted at the base of the perianth, the filaments
filiform, the anthers ovate, or in the pistillate flowers reduced to small staminodia
or altogether absent; ovary trigonous; styles 3, short, the stigmas capitate, sublo-
bate, or fimbriate; achene obtusely or acutely trigonous, enclosed in the fleshy
and accrescent perianth, or its apex often exserted, the pericarp crustaceous or
coriaceous; seed usually 3-sulcate or subtrilobate, the embryo excentric or lateral,
the cotyledons narrow or oblong.

About 20 species, in Australia, New Zealand, the Pacific islands,
and the higher mountains of tropical America. Only the following
ones are known from North America.

Plants leafless, the stems flat, articulate, ribbon-like M. platydada.

Plants with normal green leaves, the stems subterete, continuous.

Flowers solitary or fasciculate in the leaf axils; plants small, prostrate or nearly

so; leaves acute at the base M. volcanica.

Flowers racemose or paniculate; plants scandent; leaves cordate at the base.

M. tamnifolia.

Muehlenbeckia complexa Meisn., sometimes called "wire vine"
by florists of the United States, a native of New Zealand, is planted
for ornament in Guatemala City and perhaps elsewhere. It has
long, much-branched and interlaced, scandent, woody stems and
small, orbicular or panduriform, green leaves 1-2 cm. broad.

Muehlenbeckia platyclada Meisn. Bot. Zeit. 22: 313. 1865.
Solitaria; Secretariat, (probably an accidental alteration of the first
name); Tenia.

Cultivated commonly for ornament or as a curiosity in gardens
at low and middle elevations; more or less naturalized about Coban
in thickets and hedges, and probably also in other parts of the
country. Native of the Solomon Islands.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 121

A small or large vine, often somewhat woody below, sometimes suberect,
glabrous, pale green; older stems subterete, the branches flat and ribbon-like,
mostly 1-1.5 cm. wide, many-nerved, conspicuously articulate and divided by
cross partitions into short joints; flowers small and greenish, inserted at the sides
of the nodes; fruits small, berry-like, bright red.

The succulent fruits are sometimes eaten. The plant is common
in many parts of Guatemala, where it thrives with little or no
attention and endures perfectly the long dry season. In Salvador
it is sometimes called "pie de muneco."

Muehlenbeckia tamnifolia (HBK.) Meisn. Gen. PI. 2: 227.
1840. Polygonum tamnifolium HBK. Nov. Gen. & Sp. 2: 180. 1817.
P. flexuosum Benth. PL Hartweg. 80. 1841 (type from Quezaltenango,
Hartweg 561).

Wet or damp thickets or forest, often on open banks, 1,800-3,500
meters; El Progreso; Zacapa (Sierra de las Minas); Guatemala;
Sacatepe*quez ; Chimaltenango; Quiche"; Huehuetenango; Quezalte-
nango; San Marcos. Southern Mexico; Costa Rica; Panama;
western South America.

A scandent glabrous herb or shrub, sometimes covering tall trees, the stems
as much as 2.5 cm. thick, the branches terete or angulate; leaves slender-petiolate,
oblong to elliptic-ovate, mostly 5-11 cm. long, acute to long-acuminate, usually
subhastate-cordate at the base, with a rather shallow, open sinus, sometimes with-
out basal lobes; flowers racemose, the racemes mostly shorter than the leaves but
sometimes longer and paniculate; flowers yellowish green; fruits small, subglobose
or ovoid, red, turning bluish black at maturity.

The fresh leaves are slightly succulent. The vine often makes
dense tangles over stumps and small trees. On Volcan de Acate-
nango it grows in open places in the Chiranthodendron forest.

Muehlenbeckia volcanica (Benth.) Endl. Gen. PI. Suppl. 4:
51. 1847. Polygonum volcanicum Benth. PI. Hartweg. 81. 1841.

Rocky mountain summits or in alpine meadows, 2,400-4,000
meters; Guatemala (cone of Volcan de Pacaya); Solola; Suchite-
pe"quez (Volcan de Atitlan); Huehuetenango (Sierra de los Cuchu-
matanes); Quezaltenango (Volcan de Santa Maria, where the type
was collected, Hartweg 562); San Marcos (Volcan de Tacana).
Chiapas; Ecuador to Bolivia.

A low, densely branched shrub, often forming dense clumps or wide mats, the
individual stems mostly 10-30 cm. long, densely leafy, angulate; ocreae deciduous;
leaves short-petiolate, thick and somewhat fleshy, rhombic-elliptic, 8-15 mm.
long, acute, cuneate-attenuate to the base; pedicels very short, solitary or fasci-

122 FIELDIANA: BOTANY, VOLUME 24

culate in the leaf axils, the flowers greenish white; fruits fleshy, black, the fruiting
calyx 3-4 mm. long; achene ovoid-trigonous, obtusely angulate.

One of the characteristic alpine species of Guatemala, confined
to the tops of the highest peaks, above timber line, and to the wide
alpine meadows of the Cuchumatanes.

NEOMILLSPAUGHIA Blake

Shrubs or small trees; leaves alternate, orbicular, cordate at the base, deeply
emarginate at the apex, on rather short petioles, membranaceous, the ocreae
deciduous; flowers small, perfect, in fascicles of 2-6 within the ocreolae, the fasci-
cles racemose and paniculate, the panicles large, rather dense, terminal; pedicels
filiform, 3-winged above, articulate below the middle; perianth petaloid in flower,
in fruit accrescent and dry, the tube very short, the 3 outer segments ovate or
oval-ovate, broadly winged along the keel, the wings decurrent upon the pedicel,
the 2 inner segments oval or oval-ovate, slightly shorter than the outer ones;
stamens 8-9, the filaments united at the base, pubescent below, the anthers
suborbicular; ovary trigonous, glabrous, the ovule erect, subsessile; styles 3,
slender, the stigmas capitate; achene trigonous-ovoid, subacute, with flat sides;
seed trigonous, the endosperm not ruminate; embryo subcentral, straight, the
radicle superior, shorter than the suborbicular cotyledons.

Two species, in Yucatan and Central America. The genus was
named for Dr. Charles F. Millspaugh, first Curator of the Depart-
ment of Botany of Chicago Natural History Museum.

Neomillspaughia paniculata (Donn. Smith) Blake, Bull.
Torrey Club 48: 85. 1921. Campderia paniculata Donn. Smith,
Bot. Gaz. 27:440. 1899.

Dry thickets of the Oriente, 300-450 meters; Zacapa; Chiquimula
(north of Chiquimula). Honduras, the type from Rio Chamelecon.

A large shrub or usually a tree of 6-11 meters, the branches widely spreading;
branchlets cinereous-puberulent; ocreae oval, 4 mm. long, caducous; petioles
stout, 1.5-3 cm. long; leaf blades orbicular, 12-22 cm. long, very deeply and nar-
rowly emarginate at the apex, shallowly and openly cordate at the base, green
above, puberulent and rough to the touch, puberulent or short-pilose beneath or
glabrate; panicles large and pyramidal, 20-25 cm. long, the pedicels mostly 3-4
mm. long, the flowers white or greenish white; fruiting perianth 5-6 mm. long,
the wings of the sepals about 1 mm. wide; achene about 3 mm. long.

Sometimes called "amarra-jabon" in Honduras. Apparently
of limited occurrence in Guatemala but conspicuous where it does
grow. It is plentiful in some of the thickets along the railroad
between Gualan and Zacapa but was not observed in the immediate
vicinity of Zacapa. It is abundant in the Comayagua desert region
of the Department of Comayagua, Honduras. Very closely related

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 123

is the only other species of the genus, N. emarginata (Gross) Blake,
of Yucatan. Leaf and pubescence characters used by Blake in
separating the two species do not hold, but the flowers of N. emargi-
nata are substantially smaller than those of N. paniculata and prob-
ably two distinct species are involved. The Yucatan species, which
may reach Pete"n, is said to be called in Maya "sacitsa" or "tsaitsa."

PODOPTERUS Humboldt & Bonpland
Reference: S. F. Blake, Bull. Torrey Club 48: 86-87. 1921.

Shrubs or small trees, the branches often flexuous but rigid, mostly spinose
at the tip; leaves clustered at the nodes, deciduous and the plants leafless for much
of the time, the blades membranaceous, the ocreae small; flowers perfect, geminate
or few in the axils of bracts, the fascicles racemose, the racemes subpaniculate at
the ends of the branches; perianth segments 6, the 3 outer ones larger, the keel
extended into a scarious wing that is decurrent upon the pedicel, the smaller
sepals plane, erect, enlarged in fruit; stamens 6, the filaments filiform, the anthers
ovate; ovary trigonous, the styles short, the stigmas capitate; ovule subsessile;
achene included in the broadly 3-winged perianth.

Three species, in Guatemala and Mexico.

Leaves rather densely pilosulous beneath on the surface as well as on the veins,
rounded or cordate at the base P. guatemalensis.

Leaves glabrous beneath or merely pilosulous at the base of the costa, acute at
the base P. mexicanus.

Podopterus guatemalensis Blake, Bull. Torrey Club 48: 87.
1921. Crucito.

Type from El Barranquillo, El Progreso, 550 meters, Wilson
Popenoe 973; collected also at El Rancho and elsewhere in the same
department; endemic.

A shrub or small tree, the branches somewhat zigzag, gray-barked, the branch-
lets spinose, densely puberulent; leaves on petioles 4-15 mm. long, broadly obovate
or oval-obovate, 2-4.5 cm. long, 2-3 cm. wide, broadly rounded or obtuse at the
apex, narrowed to a rounded or cordate base, densely short-pilose beneath; flowers
in many-flowered fascicles on usually leafless branches, the glabrous pedicels
12-17 mm. long, winged for half their length or more; calyx in fruit 8 mm. long,
glabrous, the sepal wings 2 mm. wide; stamens 8, the filaments glabrous; achene
trigonous-ellipsoid, subobtuse at each end, 5 mm. long, pale brownish.

Podopterus mexicanus Humb. & Bonpl. PI. Aequin. 2: 89.
pi. 107. 1812.

Zacapa, between Agua Blanca and Cumbre de Chiquimula.
350-500 meters, in shaded quebrada, Standley 74412. Southern
Mexico.

124 FIELDIANA: BOTANY, VOLUME 24

A tree of 6 meters, the branchlets stout, often spinose, dark ferruginous or
blackish; leaves mostly fascicled on short spurs, slender-petiolate, glabrous or
nearly so, deciduous, rounded-obovate, 4-6.5 cm. long, 2.5-4.5 cm. wide, broadly
rounded at the apex, cuneately narrowed to the often somewhat unequal base, the
veins rather prominently reticulate; flowers greenish, usually appearing before the
new leaves.

t
POLYGONUM L.

Reference: John Kunkel Small, A monograph of the North Ameri-
can species of the genus Polygonum, Mem. Dept. Bot. Columbia
Coll. vol. 1. 1895.

Annual or perennial herbs, glabrous or pubescent, often glandular, sometimes
scandent, the stems often enlarged at the nodes; leaves alternate, entire, mostly
membranaceous, often glandular-punctate; ocreae cylindric or funnelform, mem-
branous, hyaline, or rarely herbaceous, often ciliate or fringed with bristles at
the summit; flowers perfect, small, green, white, or red, fasciculate in the leaf axils
or often forming racemes or spikes; perianth herbaceous or membranaceous,
persistent, usually closely investing the achene, of 4-6 lobes or segments, these
subequal or the outer ones larger; stamens 3-9, the filaments subulate or filiform,
the anthers oblong or ovoid; ovule usually stipitate; style 2-3-cleft or 2-3-parted,
the stigmas capitate; achene lenticular or triquetrous, smooth or granular; seed
sessile, the endosperm corneous or farinose; embryo ex centric, the cotyledons
foliaceous, slender, accumbent or incumbent.

About 150 species, widely distributed in both hemispheres.
Only the following are known in Central America but 70 or more are
found in North America, mostly in the United States.

Flowers inserted in the leaf axils; leaf blades articulate with the petioles.

P. aviculare.
Flowers spicate or racemose; leaf blades not articulate.

Leaf blades rounded or subcordate at the base; plants often scandent; ocreae

oblique; racemes usually 1 cm. long or less P. Meisnerianum.

Leaf blades acute at the base; plants never scandent; ocreae truncate; racemes
all or mostly much more than 1 cm. long.

Ocreae with spreading green margins P. hispidum.

Ocreae appressed, the margins not spreading, thin and dry, not green.
Margins of the ocreae naked, not fringed with bristles.

Leaves white-tomentose beneath P. tomentosum.

Leaves neither white nor tomentose beneath.

Peduncles stipitate-glandular P. mexicanum.

Peduncles without stipitate glands.

Fruiting calyx scarcely 2 mm. long P. longiocreatum.

Fruiting calyx fully 3 mm. long P. portoricense.

Margins of the ocreae conspicuously fringed with long or sometimes very

short bristles.

Racemes very slender and interrupted; sepals punctate. . .P. punctatum.
Racemes dense, stout, not interrupted; sepals not punctate.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 125

Leaves densely strigose beneath over the whole surface.

P. acuminatum,

Leaves glabrous or glabrate beneath except sometimes on the costa.
Achenes lenticular.

Leaves lanceolate, mostly 3-5 cm. wide P. ferrugineum.

Leaves linear-lanceolate, mostly 1-2 cm. wide.

Racemes oblong, dense P. persicarioides.

Racemes linear or narrowly oblong, rather lax P. segetum.

Achenes trigonous.

Racemes oblong, dense P. persicarioides.

Racemes linear or narrowly oblong, rather lax . P. hydropiperoides.

Polygonum acuminatum HBK. Nov. Gen. & Sp. 2: 178. 1817.
P. guatemalense Gandoger, Bull. Bot. Soc. France 66: 225. 1919
(type from Alta Verapaz, Tuerckheim, probably No. 11.1399).
Chilillo; Chilillo de chucho.

Usually in marshes or stream borders, often in shallow water,
ascending from sea level to about 1,550 meters; Pete*n; Alta Verapaz;
Izabal; Jalapa; Santa Rosa. Southern Mexico; British Honduras;
Honduras; Panama; West Indies; South America.

A stout erect perennial, often a meter high or even taller, the stems glabrous
below, densely sericeous-strigose above; ocreae 2-4 cm. long, densely strigose, long-
fringed at the apex, membranaceous; leaves short-petiolate, lanceolate, 10-30
cm. long, long-attenuate, green above, somewhat paler beneath, strigose on both
sides, more densely so beneath; racemes few, paniculate, 4-10 cm. long, linear,
very dense; flowers greenish white or pinkish; achene lenticular, 2-2.5 mm. long,
black and lustrous.

A characteristic marsh plant of the Atlantic lowlands of Central
America.

Polygonum aviculare L. Sp. PI. 362. 1753. Tabaco (Quezalte-
nango; a questionable name, but the informant was insistent upon
it).

Roadsides and waste or cultivated ground, sometimes on sand-
bars along streams; central and western mountains, abundant in
some areas, 1,400-2,500 meters; Guatemala; Chimaltenango; Que-
zaltenango. Native of Europe and Asia, now naturalized as a weed
in many parts of North and South America.

A pale green annual, often bluish green, simple or much branched, procum-
bent or ascending, densely leafy; leaves almost sessile, oblong or obovate-oblong,
mostly 1-4 cm. long, acute or obtuse, narrowed and acute at the base; ocreae
membranous, white, becoming lacerate; flowers in axillary fascicles of 5 or fewer,
short-pedicellate; sepals green, the margins white or pink, 2-3 mm. long; achene
3-angulate, ovoid, acute, 3-4 mm. long.

126 FIELDIANA: BOTANY, VOLUME 24

The plant is plentiful in some parts of Quezaltenango and Chi-
maltenango, forming dense and large colonies in settlements. In
the United States it grows profusely in dooryards, where it with-
stands trampling better than almost any other plant except some
of the Juncus species. Elsewhere in Central America the species
has been found only in Costa Rica.

Polygonum ferrugineum Wedd. Ann. Sci. Nat. III. 13: 252.
1849.

In shallow water on lake shores of the Oriente, 500-1,600 meters;
Jalapa (near Jalapa) ; Jutiapa (Lago de Atescatempa). West Indies;
Brazil.

A coarse perennial a meter high or less with thick stems, glabrous or essentially
so; ocreae cylindric, 2-4 cm. long, sparsely ciliate when young; leaves on petioles
1-2 cm. long, lanceolate, mostly 9-25 cm. long and 2-5 cm. wide, long-attenuate,
acute or acuminate at the base, sparsely strigose beneath on the costa; inflores-
cence paniculate, the racemes spikelike, linear, 2-7 cm. long, dense, erect; ocreolae
3 mm. long, conspicuous, serrate and ciliate at the apex; flowers pinkish, the peri-
anth 3-4 mm. long; style biparted almost to the base; achene lenticular, 3-3.5
mm. long, orbicular, almost black, lustrous.

Polygonum hispidum HBK. Nov. Gen. & Sp. 2: 178. 1817.

Marshes, wet fields, often at the borders of streams or lakes,
sometimes on sandbars, ascending from sea level to about 1,800
meters, most common at low elevations; Izabal; Zacapa; Chiqui-
mula; Jalapa; Jutiapa; Guatemala; Quiche". Honduras; Panama;
West Indies; South America.

A coarse stout perennial, often a meter tall, glutinous, the stems hispid and
glandular; ocreae cylindric, 1-3 cm. long, often concealing almost all the stem,
densely hispid, with a conspicuous green herbaceous spreading border, fringed with
long bristles; leaves petiolate, ovate to broadly lanceolate, bright green, mostly
10-20 cm. long and 2-8 cm. wide, long-acuminate, abruptly contracted and decur-
rent at the base, strigose or hispid on the nerves or sometimes almost glabrous;
racemes paniculate, very dense, linear-oblong, 2-10 cm. long, erect, the flowers
white, greenish, or dark red, pedicellate; perianth 4.5 mm. long; style 2-parted
to below the middle; achene lenticular, 4.5 mm. long, rounded-obovoid or orbic-
ular-oblong, sometimes broader than long, black and lustrous.

Polygonum hydropiperoides Michx. Fl. Bor. Amer. 1: 239.
1803. Flor de chajutal (Quezaltenango).

Willow thickets and sandbars near streams, 1,200-1,900 meters;
Alta Verapaz; Huehuetenango; Quezaltenango. United States;
Mexico; Honduras; Panama; western and southern South America.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 127

Plants perennial, slender, glabrous or sparsely strigillose, erect, or the bases of
the stems decumbent and rooting, simple or branched; ocreae cylindric or funnel-
form, 1-2 cm. long, strigose, fringed with long bristles; leaves short-petiolate,
lanceolate to oblong-lanceolate or linear-lanceolate, mostly 5-10 cm. long and
5-15 mm. wide, attenuate at each end, ciliate; racemes narrowly cylindric or almost
linear, 3-6 cm. long, erect, more or less interrupted; ocreolae 2.5-3 mm. long,
ciliate; flowers green or white, sometimes pinkish, the perianth 2 mm. long, glandu-
lar; style 3-parted to below the middle; achene triquetrous, 3 mm. long, ovoid,
pointed, lustrous, black.

Polygonum longiocreatum Bartlett, Proc. Amer. Acad. 43:
51. 1907.

Marshy fields or stream borders, sometimes on sandbars, 400
meters or less; Zacapa (type from Gualan, C. C. Deam 374); Chi-
quimula. Atlantic coast of Honduras.

Plants perennial, erect or decumbent, the base often elongate and rooting,
70 cm. high or less, glabrous; ocreae cylindric, mostly 1.5-2 cm. long, naked at
the apex, glabrous or nearly so; leaves short-petiolate, narrowly lanceolate, 8-14
cm. long, 1.5-3 cm. wide, long-attenuate, acute at the base, glabrous; racemes very
slender, almost linear, 3-6 cm. long, dense or somewhat interrupted, the peduncles
glabrous; perianth rose-pink, in fruit about 2 mm. long; achene lenticular, 2 mm.
long, black, lustrous.

Polygonum Meisnerianum Cham. & Schlecht. Linnaea 3:
40. 1828. P. Beyrichianum Cham. & Schlecht. op. cit. 40. 1828.
P. Meisnerianum var. Beyrichianum Meisn. in Mart. Fl. Bras. 5, pt.
1: 19. 1855.

Usually in marshes or open swamps, 1,300-1,800 meters; Alta
Verapaz; Jalapa. Costa Rica; southeastern United States; southern
Mexico; West Indies; Brazil.

Plants perennial, very slender, often more or less scandent and with elongate
stems, these sparsely glandular-hispidulous and often with larger recurved prickle-
like hairs at the nodes; leaves sessile or short-petiolate, linear or lance-linear, 5-15
cm. long, 5-15 mm. wide, attenuate, subcordate at the base or sometimes hastate,
usually aculeolate beneath along the costa, elsewhere glabrous or nearly so; ocreae
oblique, not ciliate; racemes mostly 1 cm. long or less, few-flowered, the peduncles
dichotomous, few, the peduncles glandular; perianth greenish white or pink, 2-3
mm. long; achene triquetrous, dark brown, lustrous.

The plant is abundant in some of the bogs and marshes not far
from Coban, but during April, at least, it seems to be a shy bloomer.
Few of the plants at that time are well developed, and it is probable
that they attain their best development during the wet summer
months.

128 FIELDIANA: BOTANY, VOLUME 24

Polygonum mexicanum Small, Bull. Torrey Club 19: 356.
1892. P. segetum var. verrucosum Stanford, Rhodora 27: 181.
1925 (type from Coban, Alta Verapaz, Tuerckheim 11.1207). Lechuga
de agua; Chilillo.

In shallow lake margins or on moist banks, 450-1,300 meters;
Alta Verapaz; Jalapa; Jutiapa (Lago de Giiija); Santa Rosa;
Escuintla. Southern United States and Mexico. .»..- .......

Plants annual or perennial, slender, glabrous below the inflorescence or some-
times stipitate-glandular on the stems, usually 60 cm. tall or less; ocreae cylindric,
5-15 mm. long, sparsely hispidulous or almost glabrous, not ciliate; leaves petio-
late, narrowly lanceolate to linear-lanceolate, 5-12 cm. long, mostly 2 cm. wide or
less but sometimes as much as 3 cm. wide, obscurely, punctate, ciliate, sometimes
glandular or stipitate-glandular beneath; peduncles usually densely glandular,
the racemes oblong, 1.5-3.5 cm. long, erect, dense; ocreolae funnelform, 3 mm.
long, ciliate; calyx pale pink, 2-3 mm. long; style 2-parted to below the middle;
achene lenticular, 3-4 mm. long, ovoid or broadly ovoid, inconspicuously gibbous
on one side, dark brown or almost black, usually granular and dull.

P. segetum var. verrucosum seems to be satisfactorily referable
here. It certainly is a species altogether distinct from P. segetum,
to which Stanford referred it.

Polygonum persicarioides HBK. Nov. Gen. & Sp. 2: 197.
1817.

Stream and lake margins or wet thickets, often on sandbars,
ascending from near sea level to about 1,800 meters; Alta Verapaz;
Baja Verapaz; Zacapa; Chiquimula; Jalapa; Santa Rosa; Escuintla;
Guatemala; Huehuetenango. United States and Mexico; western
and southern South America.

Plants perennial, almost glabrous or strigillose, erect or decumbent and rooting
at the base, mostly 70 cm. tall or less; ocreae cylindric or funnelform, 1-2 cm. long,
glabrous or sparsely strigose, inconspicuously fringed with short bristles; leaves
short-petiolate, lanceolate or linear-lanceolate, mostly 4-10 cm. long and 4-15
mm. wide, acuminate or attenuate at each end, sometimes strigose beneath on the
costa, punctate; racemes erect, narrowly oblong or linear, 2-6 cm. long, rather
lax; ocreolae oblique, 3 mm. long, ciliate or naked; perianth 2-3 mm. long, pinkish
white or green and pink; style 2-3-parted to near the base; achenes lenticular or
triquetrous upon the same plant, 2.5-3 mm. long, black, lustrous.

Polygonum portoricense Bertero ex Meisn. in DC. Prodr. 14:
121. 1856, as syn.; Small, Mem. Dept. Bot. Columbia Coll. 1: 46.
pi. 10. 1895. Lechuga.

Wet meadows or stream borders, 500-2,500 meters; Baja
Verapaz; Jalapa; Escuintla; Quezaltenango. Southern United
States; West Indies; South America.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 129

A rather stout perennial, glabrous or nearly so, erect, a meter tall or less, often
much branched; ocreae cylindric, 1.5-4 cm. long, when young often ciliate but in
age without marginal bristles, sometimes hispid; leaves petiolate, lanceolate or
narrowly lanceolate, 5-25 cm. long, 1-4 cm. wide, acuminate or attenuate at each
end, very obscurely punctate; racemes linear, 2-10 cm. long, erect, dense; ocreolae
funnelform, 3 mm. long, narrow, obtuse or acute, with a membranous margin;
perianth white or pink, about 3 mm. long; style 2-3-parted to below the middle;
achenes lenticular or triquetrous, 2.5 mm. long, very broadly oblong or suborbic-
ular, sometimes broader than long, black, lustrous.

Polygonum punctatum Ell. Bot. S. C. & Ga. 1: 455. 1817.
P. acre HBK. Nov. Gen. & Sp. 2: 179. 1817. Chilitto; Canilla de
pava; Chilitto de perro.

Common through much of Guatemala, in marshes and bogs,
margins of streams and lakes, wet thickets, sandbars, and waste
ground, ascending from sea level to 1,800 or rarely to about 2,400
meters; Pete"n; Alta Verapaz; Baja Verapaz; Izabal; Zacapa; Jalapa;
Santa Rosa; Escuintla; Guatemala; Sacatepe"quez; Chimaltenango;
Quiche"; Retalhuleu; Quezaltenango; San Marcos; Huehuetenango.
United States and Mexico to British Honduras and Panama; West
Indies; South America.

A slender annual or perennial, often forming dense colonies, usually glabrous
throughout, a meter high or usually lower, the stems erect or often creeping and
rooting at the base, simple or much branched; ocreae cylindric at first, 1-1.5 cm.
long, glabrous or sparsely strigose, ciliate with rather long bristles; leaves short-
petiolate, lanceolate or narrowly lanceolate, mostly 4-15 cm. long and 3.5 cm.
wide or less, acuminate at each end, conspicuously punctate, often with a few short
hairs on the costa beneath; racemes linear, very slender and often much inter-
rupted, 1-6 cm. long, erect; ocreolae funnelform, 2.5-3 mm. long, ciliate; perianth
greenish or greenish white, 2 mm. long, the segments glandular-punctate; style
2-parted or sometimes 3-parted to the base; achene lenticular, occasionally tri-
quetrous, 2.5 mm. long, black, lustrous.

This is by far the most abundant Polygonum species in Central
America and the only common one of general distribution. In
habit it often is decidedly weedy. Poultices of the leaves are applied
in Guatemala to dogs suffering from jiote or mange. The English
name, "smartweed," is applied in the United States to this and
related species of Polygonum because of the acrid properties of the
foliage.

Polygonum segetum HBK. Nov. Gen. & Sp. 2: 177. 1817.

Chiefly along stream borders, at 500 meters or less; Pete"n;
Retalhuleu; San Marcos. Southeastern United States; Mexico;
Honduras (Atlantic coast); West Indies; Colombia.

130 FIELDIANA: BOTANY, VOLUME 24

An almost glabrous perennial, sometimes strigose or glandular about the inflo-
rescence, erect, mostly 75 cm. tall or less; ocreae cylindric or narrowly funnelform,
1-1.5 cm. long, the upper ones often strigose; leaves short-petiolate, narrowly
lanceolate or linear-lanceolate, mostly 6-15 cm. long and 7-15 mm. wide, long-
attenuate to each end, sometimes strigose beneath on the costa; racemes erect,
2-4 cm. long, cylindric, rather lax; ocreolae funnelform, 2-2.5 mm. long, coria-
ceous with a membranous margin, somewhat scurfy; perianth 2-2.5 mm. long,
pale pink; style 2-parted to below the middle; achene lenticular, 2.5 mm. long,
ovoid, dark brown, minutely glandular, dull.

Polygonum tomentosum Schrank, Baier. Fl. 1: 669. 1789.
P. incanum Schmidt, Fl. Boem. 4: 90. 1795. P. Persicaria
var. incanum Meisn. Monogr. 68. 1826. P. lapathifolium var.
incanum Koch, Syn. Fl. Germ. 711. 1837.

Sandbars or rocky stream beds, sometimes a weed in gardens,
rare, 1,350-1,800 meters; Guatemala; Chimaltenango; Huehuete-
nango. Probably native of Europe, but sparingly naturalized in
North America.

Plants annual (at least Guatemalan specimens), erect, 30-50 cm. tall, the
stems scurfy or glabrate; ocreae lax, membranous, glabrous or nearly so, the mar-
gin naked or sparsely ciliate; leaves lanceolate or narrowly lanceolate, petiolate,
acute or acuminate, the tip sometimes obtuse, glabrous or nearly so above, beneath
densely covered with a white or grayish tomentum; racemes 1.5-3.5 cm. long,
rather lax, narrowly oblong; perianth green or greenish white, 3 mm. long; achene
lenticular, dark brown, lustrous.

By Small this plant was treated as a variety of P. lapathifolium
L., but recent European writers, when not recognizing it as a dis-
tinct species, have mostly considered it a form or variety of P.
Persicaria L. It seems to us that it is more easily recognizable than
most Polygonum species of this relationship, and that it might well
be considered an independent species, as treated by Ascherson and
Graebner. (Their treatment, it must be admitted, is somewhat
equivocal.)

RHEUM L. Rhubarb

Stout perennial herbs with thick, somewhat woody rhizomes; leaves often
very large, palmately nerved, often sinuate-dentate or palmate-lobate; ocreae
membranous-scarious, lax, marcescent; flowers pedicellate, fasciculate, the fasci-
cles racemose, the narrow racemes paniculate; flowers perfect or by abortion
staminate, the perianth 6-parted, spreading, the segments subequal or the outer
ones somewhat smaller, not accrescent in fruit, marcescent; stamens usually 9,
the anthers ovate; ovary trigonous, the 3 styles short, recurved, stigmatose at the
apex; achene narrowly or broadly 3-winged; embryo straight, subcentral, the
cotyledons plane, cordate or ovate, the radicle short, superior.

About 20 species, natives of eastern Asia.

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 131

Rheum Rhaponticum L. Sp. PI. 371. 1753. Ruibarbo.

Cultivated as a food plant at middle elevations, and sold in the
markets of the central region and of Coban. Native of southern
Siberia, but cultivated in most temperate regions.

A coarse perennial with thick clustered roots; leaves mostly radical, the petioles
semicylindric, succulent; leaf blades suborbicular, often 50-80 cm. broad, deeply
cordate at the base, undulate-margined, about 5-nerved at the base, glabrous
above, pubescent beneath on the veins; inflorescence a tall narrow leafy panicle
of numerous small whitish flowers; achene oblong-oval.

Rhubarb or pieplant is not common in Guatemala and is very
rare or probably absent in other parts of Central America. So sour
a plant will never find favor among tropical people, who esteem
fruits in proportion to the amount of sugar they contain. The
plant was noted as thriving about Coban and near Tecpam, and
occasionally may be found in the larger markets, where it is sold
mostly to foreigners. It was served upon the table in a pension at
Coban, with a very ample amount of sugar. The stalks seen on
sale were of medium size and thickness. The rhubarb used in medi-
cine is derived from the root of a distinct species, Rheum officinale
Baillon, of Tibet and western China.

RUMEX L.

Reference: K. H. Rechinger, Die siid- und zentralamerikanischen
Arten der Gattung Rumex, Ark. Bot. 26A, No. 3. 1933.

Chiefly perennial herbs, rarely annuals, sometimes tall and shrub-like; leaves
often forming a basal cluster, sometimes mostly cauline and alternate, often cordate
or hastate, succulent, entire or dentate; ocreae membranaceous-scarious, often
hyaline, at first sheathing, later lacerate and withering; flowers perfect or unisexual,
fasciculate at the nodes of the branches, the clusters subtended by an ocreiform
bract, the pedicels not bracteolate, the fascicles of flowers usually forming terminal
racemes or panicles; perianth segments generally 6, in anthesis sometimes equal,
the outer ones unchanged in fruit, the inner ones somewhat accrescent and embrac-
ing the fruit, entire or fimbriate, the costa sometimes bearing on the outside a
granule-like tubercle; stamens 6, the filaments very short, the anthers oblong;
ovary trigonous, the styles 3, filiform, spreading or recurved, the stigmas fimbriate
or penicillate; achene included in the inner perianth segments, trigonous, the angles
usually acute; embryo lateral, incumbent-incurved or almost straight, the coty-
ledons linear or oblong.

Perhaps 100 species, mostly in the temperate regions of the
northern hemisphere, very few in tropical regions. Several Euro-
pean species have become widely established as weeds in the New
World. The only other species of Central America is R. costaricensis
Rechinger, endemic in the high mountains of Costa Rica. It is

132 FIELDIANA: BOTANY, VOLUME 24

remarkable for its extraordinary size, being a coarse herb 5-6 meters
tall, with a stem sometimes 10 cm. in diameter.

Leaves hastate-lobate at the base; flowers dioecious; inner sepals unchanged in

fruit R. Acetosella.

Leaves not hastate; flowers perfect or polygamo-dioecious; inner sepals enlarged
in fruit.

Inner sepals laciniate-dentate R. obtusifolius.

Inner sepals entire or nearly so.

Leaves flat, pale green R. mexicanus.

Leaves crispate, deep green.

Fruiting sepals about 3 mm. long; verticels of flowers remote. R. Berlandieri.
Fruiting sepals 4-5 mm. long; verticels of flowers crowded R. crispus.

Rumex Acetosella L. Sp. PI. 338. 1753.

Damp thickets or open fields, 2,000-2,600 meters; Chimalte-
nango (Las Calderas); Quezaltenango (above Palojunoj). Native
of Europe and Asia, widely naturalized as a weed in North and South
America.

Plants perennial, slender, usually 30 cm. high or less, glabrous, with long
slender rootstocks; leaves long-petiolate, oblong or narrowly oblong, 2-10 cm.
long, obtuse or acute, somewhat fleshy, hastate-lobate at the base, the basal
lobes small, the terminal lobe entire; flowers dioecious, 1.5 mm. long, green or
often dark or bright red, in slender racemes arranged in small terminal panicles.

The plant is rather common in mountain meadows and pastures
of Costa Rica, probably introduced with grass seed.

Rumex Berlandieri Meisn. in DC. Prodr. 14: 45. 1856.

In waste ground, 1,800-2,400 meters; El Progreso (Finca Pia-
monte); Solola (San Pedro, on shore of Lago de Atitlan). South-
western United States; Mexico; probably introduced in Guatemala.

An erect perennial herb, commonly about 30 cm. high, the stems usually
several, glabrous; leaves deep green, darkening when dried, glabrous, slender-
petiolate, somewhat crispate, oblong to oblong-lanceolate, rounded to attenuate
at the apex, rounded or obtuse at the base and often abruptly decurrent; inflores-
cence often much branched, the branches erect, leafy, the verticels of flowers
numerous, separated and often remote; flowers short-pedicellate, densely crowded;
inner perianth segments in fruit about 3 mm. long, rounded-ovate, strongly
venose, each bearing dorsally a rather large and conspicuous tubercle.

The plant is rare in Guatemala, and probably has been imported
from Mexico.

Rumex crispus L. Sp. PI. 335. 1753. Lengua de vaca; Lengua de
caballo; Lechugon (fide Aguilar).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 133

Frequent in many localities, along ditches or roadsides, wet
meadows, moist thickets, sometimes a weed in cultivated ground,
especially old gardens and cafetales, chiefly at 1,500-2,500 meters;
Alta Verapaz; Baja Verapaz; Jalapa; Guatemala; Sacatepe"quez ;
Chimaltenango; Huehuetenango. A native of Europe and Asia,
thoroughly naturalized as a weed in temperate North and South
America as well as in many other regions.

A glabrous perennial 50-100 cm. tall with thick yellow perpendicular roots;
leaves petiolate, often forming dense clusters at the base of the plant, oblong or
oblong-lanceolate, mostly 15-30 cm. long, rounded or cordate at the base, some-
what fleshy, undulate or crispate; flowers green, in small or large, narrow panicles,
long-pedicellate; outer calyx lobes broadly cordate, entire or nearly so, obtuse or
acute, each bearing on the costa a hard brown tubercle; achene dark brown,
lustrous.

Called "curled dock" or "yellow dock" in the United States,
where the young leaves often are used as a pot herb, cooked like
spinach. The Guatemalans seem not to have discovered the edible
qualities of the plant, although in some localities there is an ample
supply of it available. In some parts of the world the plant is used
in domestic medicine. It is worthy of note that the very large,
fleshy roots of one species of Rumex native in northern Mexico and
southwestern United States have been found excellent for tanning
skins and have been used commercially for the purpose.

Rumex mexicanus Meisn. in DC. Prodr. 14: 45. 1856.

Moist fields, often a weed in old gardens, in the Occidente, 2,200-
2,500 meters; Quezaltenango; San Marcos. Mexico and New Mexico.

A glabrous perennial, erect or decumbent, usually 30-50 cm. tall, with stout
stems; leaves pale green, short-petiolate, chiefly cauline, narrowly oblong-lanceo-
late to linear-lanceolate, long-attenuate, obtuse or narrowed at the base; panicles
mostly rather small and very dense, naked or nearly so, pale green; inner sepals
in fruit 4-5 mm. long, broadly ovate-triangular, entire or nearly so, obtuse or
subacute, reticulate- veined, each bearing a small tubercle; achene fuscous or
almost black, 2.5 mm. long, acuminate.

The habitats of this plant in western Guatemala are such that
it appears to be an introduction (from Mexico) and rather probably
is not native in Guatemala.

Rumex obtusifolius L. Sp. PI. 335. 1753. Barba herbata
(Quezaltenango) .

Widely distributed in the mountains and abundant in many
regions, moist fields, meadows, or thickets, often in waste or culti-

134 FIELDIANA: BOTANY, VOLUME 24

vated ground, 1,200-2,700 meters; Alta Verapaz (Coban and else-
where); Guatemala; Chimaltenango; Totonicapan; Huehuetenango;
Quezaltenango; San Marcos. Native of Europe and Asia, abund-
antly naturalized in various parts of North and South America.

A coarse erect glabrous perennial, a meter tall or less, often forming large
dense colonies; basal leaves often in dense clusters, long-petiolate, oblong or
oblong-lanceolate, mostly 15-40 cm. long, obtuse or acute, cordate or rounded at
the base, almost flat; panicles usually large and rather open, the flowers green, in
loose whorls in the long racemes, the pedicels long and slender; inner sepals in
fruit about 5 mm. long, the margins deeply laciniate, one of the sepals bearing a
tubercle; achene 2 mm. long, dark red, acute, lustrous.

The greatest abundance of this plant that we have noted is in
meadows along the Rio Samala near Olintepeque, where there are
wide areas almost exclusively covered with it. Evidently it is not
eaten by stock of any kind. It is plentiful almost anywhere in the
Quezaltenango region and also about Coban. In the United States
the leaves of this species are not eaten, or at most very rarely.

RUPRECHTIA C. A. Meyer

Trees, similar to Triplaris, the leaves alternate, penninerved; ocreae deciduous;
flowers small in anthesis, dioecious, pedicellate and fasciculate within the ocreolae,
racemose, the racemes simple or paniculate, the perianth usually becoming red
in age; staminate perianth 6-parted, the segments subequal or the 3 inner ones
somewhat smaller; stamens 9, inserted on a central disk, this commonly pilosulous
or lobulate, the filaments filiform, mostly exserted, the anthers ovate or oblong;
pistillate perianth deeply 6-parted, the 3 outer segments oblong or lanceolate,
erect, accrescent after anthesis, the 3 inner segments smaller and linear, sometimes
almost obsolete; ovary trigonous, the angles obtuse; stigmas erect, oblong or
lanceolate, sessile at the apex of the ovary or on a short style; ovule sessile; achene
obtusely trigonous, pyramidal, 3-6-sulcate, hidden by the perianth; seed 3-6-sul-
cate, the endosperm lobate and ruminate, the embryo subcentral, the cotyledons
broad, plane or somewhat convolute.

Perhaps 25 species, in tropical America. Only two are known
from Central America but several occur in Mexico.

Leaves usually copiously pubescent beneath, the veins very conspicuous and
reticulate on the lower surface; fruiting calyx usually 2-2.5 cm. long.

R. chiapensis.

Leaves glabrous beneath or nearly so, the veins inconspicuous; fruiting calyx
generally 3-4 cm. long R. costata.

Ruprechtia chiapensis Lundell, ined., sp. nov.

Coastal thickets, San Marcos (Ocos). Chiapas, the type from
Las Garzas.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 135

A tree about 9 meters tall, the young branches cinnamon-brown, glabrous;
leaves coriaceous, on very short petioles, elliptic, oblong-elliptic, or ovate-elliptic,
mostly 3.5-7.5 cm. long and 2-4 cm. wide, acute or short-acuminate, acute or
obtuse at the base, entire, almost completely glabrous, at least at maturity, the
costa and nerves very slender and inconspicuous, the veins mostly obscure;
racemes mostly short and dense; outer calyx lobes oblong-spatulate, obtuse or
rounded at the apex, red, short-pilose with ascending hairs.

Arbor, ramulis glabris; folia coriacea breviter petiolata elliptica vel ovato-
elliptica 3.5-7.5 cm. longa, acuta vel breviter acuminata, basi acuta vel obtusa,
fere omnino glabra; segmenta exteriora perianthii oblongo-spathulata apice
obtusa vel rotundata pilis adscendentibus breviter pilosa.

Mexico: Las Garzas, Chiapas, January, 1939, E. Matuda 2673
(type in Herb. Chicago Nat. Hist. Mus.).

Specimens from Veracruz, in young flower only, probably are
referable to R. chiapensis. Mexican and Central American material
has been confused in the past with R. Cumingii Meisn., a South
American species.

Ruprechtia costata Meisn. in DC. Prodr. 14: 180. 1856.
R. Deamii Robinson, Proc. Amer. Acad. 43: 51. 1907. R. Keller-
manii Bonn. Smith, Bot. Gaz. 47: 260. 1909. Carreto; Sangre de
toro.

Mostly on dry rocky slopes, sometimes on arroyo banks, in the
Oriente; Zacapa (type of R. Deamii from Gualan, C. C. Deam 231;
type of R. Kellermanii from the same locality, W. A. Kellerman
5985); Chiquimula; El Progreso. Nicaragua; probably also in
Salvador.

A tree 5-9 meters high with a dense spreading crown, the trunk 35 cm. or
more in diameter; leaves on very short petioles, mostly membranaceous, often
with undulate or shallowly crenate margins, acute or acuminate, obtuse or rounded
at the base, glabrous above or nearly so, finely pilosulous beneath, densely so at
first, sometimes glabrate in age, the lateral nerves stout and very prominent, the
veins closely reticulate and prominent; racemes usually shorter than the leaves,
often numerous and crowded, the flowers short-pedicellate; fruiting calyx red,
the outer segments 3-4 cm. long, linear-spatulate, obtuse, reticulate-veined, pilose
with subappressed hairs.

The type of this species is Friedrichsthal 1179. Like all of this
collector's plants, the label of this one is headed "Guatemala," but
the original label at Vienna bears a name that has been deciphered
as "Tinotepe," and probably should be interpreted as Jinotepe,
Nicaragua. This type collection was reported from Guatemala by
Hemsley as Ruprechtia Cumingii Meisn., and that species has been
reported from Salvador, Costa Rica, and Panama, and even from

136 FIELDIANA: BOTANY, VOLUME 24

Mexico. While we have seen no South American material of R.
Cumingii, it is improbable that that species extends so far north
as Guatemala or Mexico. R. costata is a showy tree when in fruit
because of the great abundance of red inflorescences. In gen-
eral appearance it is much like a Triplaris, but smaller in all its
parts. We have seen fragmentary material of the type of R. costata,
and it agrees well with Guatemalan specimens.

TRIPLARIS Loefling

Trees, the branches mostly hollow and septate; ocreae deciduous; leaves large,
short-petiolate, often with 3-6 longitudinal distant lines on each side of the costa,
these indicating the folds of the blade in bud; flowers dioecious, racemose, the
racemes paniculate or fasciculate, the bracts small, ovate, acute, the ocreolae
larger, long-acuminate, deeply slit on the anterior side; staminate perianth seg-
ments 6, subequal; stamens 9; segments of the pistillate perianth 6, the 3 outer
ones connate into a short or long tube, in fruit greatly enlarged and red, the 3
inner segments free or partially adnate to the tube, small and narrow, little if at
all exceeding the tube, usually shorter; achene trigonous, its angles usually acute;
seed ovoid-trigonous; endosperm more or less lobate and ruminate; embryo sub-
central, the cotyledons broad, plane or slightly convolute, the radicle short,
superior.

Probably about 20 species, mostly in South America, only two
in North America. One other species, T. surinamensis Cham., with
oblong glabrous leaves, grows in Costa Rica and Panama.

Triplaris melaenodendron (Bertol.) Standl. & Steyerm.
Field Mus. Bot. 23: 5. 1943. Vellasquezia melaenodendron Bertol.
Fl. Guat. 40. pi. 11. 1840. T. Macombii Bonn. Smith, Bot. Gaz. 19:
257. 1894 (type from Salvador). T. Macombii var. rufescens Donn.
Smith, Bot. Gaz. 20: 293. 1895. Mulato; Palo mulato; Hormigo
(Santa Rosa).

Thickets or forest of the Pacific plains and foothills, at 750
meters or less; Santa Rosa; Escuintla (type from Escuintla, Velas-
quez); Suchitepe"quez (type of T. Macombii var. rufescens from
Mazatenango, Heyde & Lux 6375); Retalhuleu; San Marcos.
Chiapas; Salvador; Nicaragua; Costa Rica; Panama.

A tree 6-12 meters tall or often larger, with rounded crown; ocreae thin and
loose, deciduous; leaves short-petiolate, mostly elliptic to oval-elliptic, about
17-35 cm. long and 8-16 cm. wide, thin, bright green, acute or abruptly short-
acuminate, rounded or even shallowly cordate at the base, strigose when young
or short-pilose, sometimes glabrate in age; flowers greenish at first, becoming red
in age, the racemes forming large terminal panicles; fruiting calyx about 5 cm.
long, the tube 1-1.5 cm. long, sericeous, the lobes oblong-spatulate, obtuse,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 137

reticulate- veined, the 3 interior lobes linear or subulate, about equaling the tube;
achene 1 cm. long, lustrous.

In Salvador sometimes called "canilla de mula" and "gallito."
The wood is yellowish, rather light and soft but firm, with straight
or fairly straight grain, of medium texture, easy to work, takes a good
polish, apparently is not durable. It is used locally for construction
purposes. The hollow branches are almost invariably inhabited by
savage ants that inflict • painful bites when the tree is molested.
The stumps send up sprouts after the tree has been felled. The tree
is abundant in many parts of the Pacific plains, and often affords
wide displays of color, especially in late January and February. It
is one of the most characteristic species of the Pacific coast of all
Central America. The name "mulato" refers to the coarsely mottled,
pale bark.

The nomenclature of the Triplaris species is confused and the
classification of the species is obscure and difficult. The species
have been based upon minor flower details which are found incon-
stant if more than a single specimen of each "species" is available.
It is probable that when the genus is carefully monographed with
the rather ample material now available, a large proportion of the
published names will be reduced -to synonymy. T. melaenodendron
may well be the same as one of the South American species, although
it antedates most of them. The earliest species names published by
Linnaeus, Aublet, and Jacquin have been treated by most authors
as undeterminable, but a sensible study of the genus probably will
result in their identification. T. melaenodendron has been referred
in most recent publications upon Central America to T. americana
L., the earliest species of the genus, whose identity is at present
uncertain.

CHENOPODIACEAE. Goosefoot Family
Reference: Standley, N. Amer. Fl. 21: 3-93. 1916.

Herbs in the Guatemalan groups, sometimes shrubs or small trees, glabrous
or pubescent, the pubescence often of inflated hairs; leaves opposite or alternate,
sessile or petiolate, often succulent, sometimes reduced to scales; flowers perfect,
polygamous, monoecious or dioecious, usually regular, small, and greenish; peri-
anth simple, sometimes wanting in pistillate flowers, herbaceous or membrana-
ceous, usually of 2-5 segments, these more or less united below, persistent after
anthesis; stamens equaling or fewer than the perianth segments and opposite
them, hypogynous or adnate to a disk or to the base of the perianth; filaments
linear, subulate, or filiform, the anthers dorsifixed, didymous, oblong, or sagittate,
2-4-celled, introrse, dehiscent by ventral or lateral fissures; ovary superior, free
or rarely adnate to the base of the perianth, 1-celled; style terminal, the stigma

138 FIELDIANA: BOTANY, VOLUME 24

capitate, or the styles 2-3, elongate, and introrsely papillose, the stigmas 2-5
and sessile; ovule solitary, campylotropous, erect on a short basal funicle or sus-
pended from the apex of an elongate funicle; fruit a utricle, usually included in
the perianth and often deciduous with it, indehiscent or rarely circumscissile; seed
erect, inverted, or horizontal, the endosperm farinaceous, fleshy, or none; embryo
annular or hippocrepiform and enclosing the endosperm, or sometimes dorsal
and conduplicate.

A large family, most abundantly represented in Asia and eastern
Europe but generally distributed in temperate regions, with but
few representatives in the tropics. In North America about 25
genera are represented, chiefly in the western United States. Only
one genus is represented by native species in Central America.

Flowers bracteate and bracteolate; perianth closed, indurate, and nutlike in fruit.

Beta.
.Flowers without bracts or bractlets; perianth unchanged in fruit. . .Chenopodium.

BETA L. Beet

Annual, biennial, or perennial herbs, the roots fleshy and often much thick-
ened; basal leaves rosulate, the cauline ones alternate, entire or sinuate; flowers
perfect, bracteate and bibracteolate, small, in glomerules of 3 or more, the glom-
erules solitary in the axils or in terminal, simple or paniculate spikes; perianth
urceolate, 5-lobate, adherent to the base of the ovary and to the other flowers of
the same glomerule, in fruit closed and indurate, costate; stamens 5, perigynous;
filaments subulate, the anthers oblong; stigmas 2-5, short, connate at the base;
pericarp free from the seed, attached below to the perianth; seed horizontal,
orbicular or reniform, smooth, the embryo annular or nearly so, surrounding the
copious endosperm.

About half a dozen species, natives of Europe, northern Africa,
and Asia.

Beta vulgaris L. Sp. PI. 222. 1753.

Nomenclature of the various forms of beets is greatly confused,
and European botanists are far from agreement as to the classifi-
cation and names of the cultivated or even the wild forms. It is
believed, however, that cultivated beets are derived from the wild
perennial beet (Beta vulgaris var. perennis L.) that grows along the
coasts of Europe from The Netherlands southward and eastward
along the Mediterranean shores. The common cultivated forms
known in America are the following:

Beta vulgaris var. crassa Alef. Landw. Fl. 280. 1866. Remo-
lacha; Acelga.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 139

A common vegetable almost throughout Guatemala, grown
most extensively in the mountains but also in the lowlands. The
roots are a common article of food and the leaves also are cooked
and eaten very generally. To this variety belongs also the sugar
beet, cultivated on a large scale in the United States and Europe
as a source of sugar. It, however, is not grown in Central America
unless it may have been planted experimentally.

Beta vulgaris var. Cicla L. Sp. PL 222. 1753.

To this variety belongs the chard or Swiss chard, which has
slender roots that are not eaten, and very large, pale, more or less
crisped and curled leaves with very thick and succulent midribs.
It was noted in cultivation at Guatemala, Coban, and Momoste-
nango, and is planted occasionally throughout the cooler regions.
The leaves are cooked and eaten. It is not a common vegetable,
but is sometimes seen in the markets.

CHENOPODIUM L.

Annual or perennial herbs, sometimes with a strong odor, usually either
glandular or covered with a farinose pubescence of small white inflated hairs;
leaves alternate, usually petiolate, entire, dentate, or pinnatifid; flowers mostly
perfect, without bracts, small, usually glomerate, the glomerules variously ar-
ranged; perianth usually 5-parted or 5-lobate, the segments often carinate or
corniculate-appendaged, herbaceous; stamens 5 or fewer, the filaments sometimes
connate at the base, the anthers didymous or oblong; style usually none, the
stigmas 2-5, subulate or filiform; utricle ovoid and erect, or depressed-globose,
the pericarp membranaceous or fleshy, free from the seed or adherent to it; seed
horizontal or vertical; embryo annular or incompletely annular, surrounding the
copious farinaceous endosperm.

Probably 80 species or more, about 50 being known from North
America, the rest distributed through the other continents, chiefly
in temperate regions. Only the following species are known from
Central America. Chenopodium Quinoa Willd. (C. Nuttalliae Saf-
ford) is an important food plant in the Andes of Peru and Ecuador,
where the whitish seeds or the whole inflorescences are cooked and
eaten. The plant was introduced into central Mexico and is grown
for food in some regions of the Mexican mountains.

Plants farinose, not glandular, not strong-scented.

Leaves lustrous on the upper surface, not lobate; inflorescences mostly shorter

than the leaves, not forming conspicuous terminal spikes C. murale.

Leaves dull, often hastate-lobate; inflorescences mostly longer than the leaves

and usually forming a conspicuous terminal spike C. Berlandieri.

140 FIELDIANA: BOTANY, VOLUME 24

Plants not farinose, gland-dotted, strong-scented.

Leaves pinnate-lobate; inflorescence loosely dichotomous, some of the flowers
pedicellate; calyx lobes with hornlike appendages; pericarp not gland-
dotted C. graveolens.

Leaves merely dentate or entire; inflorescence glomerate-spicate, the flowers
sessile; calyx lobes not appendaged; pericarp gland-dotted.

C. ambrosioides.

Chenopodium ambrosioides L. Sp. PI. 219. 1753. C. anthel-
minticum L. op. cit. 220. Apazote; Apazote de caballo; Apazote de
zorro; Epazote; Sicaj (Baja Verapaz, fide Tejada); Siquij (Chimalte-
nango, fide Tejada) ; Saqueen (Huehuetenango, fide Tejada) ; Uicqej
(Huehuetenango, fide Tejada) ; Achij (Huehuetenango, fide Tejada) ;
Rescaj (Quiche", fide Tejada); Sicajpar (Totonicapan, fide Tejada);
Eiskiij pur (Coban, Quecchi) ; Pazote.

Usually a weed in waste ground about houses, often in cultivated
fields, sometimes on sandbars, widely distributed, and ranging from
sea level to 2,700 meters or more; Pete'n; Alta Verapaz; Zacapa;
Chiquimula; Jalapa; Jutiapa; Santa Rosa; Escuintla; Guatemala;
Sacatepe'quez ; Chimaltenango; Quiche"; Huehuetenango; Totoni-
capan; Quezaltenango; San Marcos. United States to Mexico,
British Honduras, and Panama; West Indies and South America;
naturalized in many parts of the Old World.

Plants annual or perennial, erect or ascending, ill-scented, a meter high or
less, stems simple or branched, glandular-villous or tomentulose about the inflores-
cence; lower leaves petiolate, the blades 3-10 cm. long, 1.5-5.5 cm. wide, oblong
to ovate or lanceolate, coarsely and irregularly sinuate-dentate or sinuate-pinnati-
fid, obtuse to attenuate at the apex, cuneate at the base, copiously gland-dotted,
or the glands sometimes wanting, puberulent, short- villous, or glabrous; flowers
usually densely glomerate in dense or interrupted spikes, these leafy or naked;
calyx 1 mm. high, glabrous or short-villous, usually gland-dotted, the lobes com-
pletely enclosing the fruit; seed horizontal or vertical, 0.6-0.8 mm. broad, almost
black.

The plant has a very distinctive and nauseous odor. It has long
been known as an efficient agent for expelling intestinal parasites,
and is official in the pharmacopoeias of the United States and other
countries, the seeds being known in the United States as Mexican
wormseed. It is much used for this purpose in Guatemala, and small
bunches of the green shoots are offered in the markets. Strangely
enough, considering its vile odor, the plant is employed also for
flavoring food, especially frijoles negros and jutes (fresh-water snails),
to which it imparts an altogether agreeable taste. The plant finds
still further use in local medicine. There came to the attention of
the senior author a case in which fomentations of the plant and hot

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 141

poultices were applied to an inflamed and supposedly infected foot
by one of the best-known North American doctors practicing in
Guatemala. It is said that about Coban the plant is employed as a
"narcotic," the plant being placed beneath the pillow to induce sleep.
Considering how unpleasant the odor is, one would expect the effect
to be quite the opposite.

Chenopodium Berlandieri Moq. Chenop. Enum. 23. 1840.
C. Berlandieri subsp. yucatanum Aellen, Repert. Sp. Nov. 26: 59.
1929. Bledo.

Occasional as a weed in cultivated ground, in streets, on sand-
bars, or along roadsides, 1,300-2,200 meters; Jalapa; Guatemala;
Sacatepe"quez ; Chimaltenango; Quezaltenango. United States and
Mexico.

An erect annual, usually a meter high or less, often much branched, pale;
leaves slender-petiolate, rhombic-ovate to ovate or oblong, mostly 3-6 cm. long,
acute or obtuse, irregularly sinuate-serrate and often somewhat hastate-lobate,
especially the larger or lower leaves, often densely farinose when young, some-
times glabrate, the uppermost leaves smaller and narrower; glomerules of flowers
usually forming dense or lax, paniculate spikes; calyx densely farinose, the broad
lobes carinate, enclosing the fruit at maturity; pericarp more or less adherent
to the seed, this horizontal, 0.8-1 mm. broad, punctate, black, usually rather dull.

This species is closely similar to C. album L., with which it had
been generally confused until this group of the genus was intensively
studied by Aellen. At least during the dry months, it is a rare
plant in most parts of Guatemala but may be more plentiful during
the wet season. In the United States the leaves and young shoots
of this group of Chenopodium have been much used for food, treated
like spinach. It is quite probable that they are so utilized in Guate-
mala when available.

Chenopodium graveolens Lag. & Rodr. Anal. Cienc. Nat. 5:
70. 1802. C. incisum Poir. in Lam. Encycl. Suppl. 1: 392. 1811.
Epazote de zorro; P azote; Apazote de zorro.

Open rocky hillsides, often a weed in cornfields, 1,800-3,000
meters or even higher; Chimaltenango; Quiche"; Quezaltenango.
Southwestern United States and Mexico; South America; Africa.

A strong-scented erect annual 20-80 cm. tall, simple or branched, sparsely
puberulent or glabrate, often tinged with red; leaves slender-petiolate, deltoid-
ovate or oblong to narrowly oblong in outline, 2-6 cm. long, 1.5-3 cm. wide, obtuse
to acuminate, truncate or narrowed at the base, sinuate-pinnatifid or laciniate-
pinnatifid, the lobes obtuse to long-acuminate, bright green, glabrous or minutely
viscid- villous on the upper surface, covered beneath with yellow glands; inflores-

142 FIELDIANA: BOTANY, VOLUME 24

cence of numerous, loosely few-flowered, axillary cymes, these forming narrow
elongate naked panicles; flowers sessile in the forks of the branches and solitary
at the ends of the slender lateral branches, the pedicellate flowers usually abortive,
their pedicels spinose; calyx lobes corniculate-appendaged, covered with yellow
glands, incompletely enclosing the fruit; seed horizontal, 0.5-0.8 mm. broad, dark
brown, the pericarp adherent.

The names "epazote de toro," "hediondillo," and "quelite
hediondo" are reported from Mexico. During the rainy season
the plant is plentiful about Quezaltenango, especially in cornfields,
but it withers quickly after the rains cease, or perhaps after being
frosted. In North American Flora the name Chenopodium incisum
Poir. was used for this species, a usage followed also by Aellen, C.
graveolens being cited as a doubtful synonym. An excellent specimen
in the Herbarium of Chicago Natural History Museum of pre-
sumably authentic material of C. graveolens, received from the
Madrid herbarium, is certainly conspecific with C. incisum.

Chenopodium murale L. Sp. PI. 219. 1753. Hedionda;
Hediondilla; Paletilla.

A weed in gardens, waste ground, or old fields, sporadic or in
some localities plentiful, 800-2,500 meters; Baja Verapaz; Jalapa;
Escuintla; Guatemala; Sacatepe*quez; Chimaltenango; Quiche; Hue-
huetenango; Quezaltenango. Native of Europe, Asia, and Africa,
but widely naturalized in America as a weed.

An erect or ascending annual, succulent, usually 40-60 cm. high, simple or
usually much branched from the base, the branches glabrous or sparsely farinose;
leaves slender-petiolate, ovate or rhombic-ovate, 3-8 cm. long, 2.5-5 cm. wide,
acute or obtuse, cuneate to subtruncate at the base, irregularly sinuate-dentate
to laciniate-serrate with obtuse or very acute teeth, glabrous or often copiously
farinose, at least beneath; flowers sessile, more or less farinose, the small glomerules
arranged in lax or dense, axillary and terminal, mostly leafless cymes or panicles;
calyx lobes obscurely carinate, incompletely enclosing the fruit; pericarp green,
adherent; seed horizontal, 1.2-1.5 mm. broad, dull, finely puncticulate.

The plant is a rather frequent weed in the cafetales about
Antigua.

Kochia scoparia (L.) Schrad., Globo japones, is sometimes planted
in Guatemalan gardens but is rather infrequent. It is an annual a
meter high or less, of very dense and bushy growth, with linear,
somewhat sericeous leaves. The cultivated form of the species has
been given the name K. trichophylla Stapf but it differs only varie-
tally, if at all, from the wild form of the species, which is a native
of Asia and southern Europe.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 143

Spinacia oleracea L., Espinaca ("spinach"), probably is grown
occasionally in the mountains as a food plant, but all or most
of the espinaca we have seen in Guatemalan gardens is actually
Tetragonia expansa (Aizoaceae), which thrives much better in tropi-
cal regions than does true spinach. The latter is perhaps native of
southwestern Asia but has been cultivated in the Old World for
many centuries, and is grown on a large scale for market in the
United States.

AMARANTHACEAE. Amaranth Family
Reference: Standley, N. Amer. Fl. 21: 95-169. 1917.

Herbs or shrubs, rarely trees, sometimes scandent; leaves opposite or alter-
nate, without stipules, petiolate or sessile, almost always entire; flowers perfect,
polygamous, or dioecious, bracteate and bibracteolate, or rarely in clusters of 2-5
and each cluster subtended by a bract and 2 bractlets, small and usually green or
greenish, solitary, capitate, spicate, or racemose; bracts and bractlets usually
hyaline, never foliaceous; perianth regular or nearly so, rarely absent, the seg-
ments generally 5, scarious, hyaline, or chartaceous, very rarely herbaceous, free,
or united at the base, usually erect, equal or the inner ones smaller; stamens
usually as many as the perianth segments and opposite them, hypogynous or
perigynous; filaments free or united into a short or elongate, 4-10-lobate tube, the
antheriferous lobes linear, subulate, or ligulate, entire or variously cut, often with
intermediate lobes (pseudostaminodia) ; anthers dorsifixed, short or elongate, 2- or
4-celled, dehiscent by introrse slits; ovary ovoid to globose, superior, free or adnate
to the base of the perianth, often compressed, glabrous or pubescent, 1-celled;
styles 1 or 2 or wanting, the stigma capitate, penicillate, or the stigma branches
2 or 3 and short or elongate; ovules solitary or numerous, erect or suspended from
the apex of an elongate basal funicle; fruit a membranaceous or fleshy utricle,
evalvate, indehiscent, irregularly dehiscent, or circumscissile; seeds erect or
inverted, lenticular, oblong to reniform-orbicular, naked or arillate, the testa
crustaceous or coriaceous, usually lustrous and smooth or nearly so; endosperm
copious, farinaceous, the embryo annular or hippocrepiform, the cotyledons
incumbent, the radicle superior or inferior.

A large family of about 50 genera, widely distributed in both
hemispheres, in America best developed in South America and chiefly
in tropical areas. In North America 21 genera are known. All the
Central American genera (and most of the species) are known from
Guatemala.

Leaves alternate; anthers 4-celled.
Ovules and seeds 2 or more.

Fruit dry; perianth segments erect; plants herbaceous Celosia.

Fruit baccate; perianth segments spreading in fruit; plants woody.

Pleuropetalum.

144 FIELDIANA: BOTANY, VOLUME 24

Ovule 1.

Plants woody and often scandent; seed arillate; filaments united at the base.

Chamissoa.

Plants herbaceous, annual, never scandent; seed not arillate; filaments free.

Amaranthus.
Leaves opposite.

Anthers 4-celled; flowers or flower clusters deflexed in age, the bracts or segments
of the sterile flowers slender and spine-like.

Flowers all fertile, each subtended by a bract and 2 bractlets, the tips of the
perianth segments never uncinate Achyranthes.

Flowers partly sterile, glomerate in the axils of bractlets, the tips of the seg-
ments of the sterile flowers uncinate at the apex Cyathula.

Anthers 2-celled; flowers not deflexed in age, their segments not spine-like.
Perianth segments united to form a hard tube, this cristate or winged in

fruit. Perennial herb with white tomentose pubescence Froelichia.

Perianth segments usually free or nearly so, unchanged in fruit, not cristate

or winged.
Stigma capitate or shallowly bilobate.

Lobes of the stamen tube entire; plants various in habit. . . Alternanthera.
Lobes of the stamen tube 3-lobate, dentate, or laciniate; scandent shrubs.

Pfaffia.
Stigma 2-3-lobate, the lobes subulate or filiform.

Lobes of the stamen tube 3-lobate, dentate, or laciniate; pseudostami-

nodia none; herbs Gomphrena.

Lobes of the stamen tube entire; pseudostaminodia sometimes present;

shrubs or often herbs.

Flowers compressed, in few spikes about 1 cm. thick; herb of seacoasts.

Philoxerus.

Flowers not compressed, in very numerous paniculate spikes rarely
as much as 5 mm. thick; herbs or shrubs, not of seacoasts.

Iresine.

ACHYRANTHES L.

Annual or perennial herbs, erect or decumbent, glabrous or pubescent; leaves
opposite, petiolate, entire; flowers perfect, bracteate and bibracteate, deflexed in
age, green or whitish, in slender, elongate, simple or branched spikes; perianth
4-5-parted, indurate in age, the segments subequal, nerved, glabrous or pubescent;
stamens 5 or rarely 2 or 4, the filaments filiform-subulate, united at the base;
pseudostaminodia quadrate, erose, lacerate, or entire, often cristate dorsally;
anthers 4-celled; ovary oblong, subcompressed, glabrous; style filiform, the stigma
capitate; ovule 1, suspended from the apex of an elongate funicle; utricle included
in the perianth, rounded or areolate at the apex, membranaceous, indehiscent;
seed inverted, oblong, the embryo annular.

About 10 species in the tropics of both hemispheres, the American
plants probably adventive from the Old World. Only two species
are known in America.

Leaves oval to broadly ovate, acuminate; sepals 6-7 mm. long A. aspera.

Leaves orbicular to obovate-orbicular, rounded and sometimes very abruptly
short-acute at the apex; sepals 4 mm. long A. indica.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 145

Achyranthes aspera L. Sp. PI. 204. 1753. Centrostachys aspera
Standl. Journ. Wash. Acad. Sci. 5: 75. 1915. Cola de armado; Pene-
gato (Guatemala); Pije de gato; Chile de perro.

A weedy plant, common in wet or moist thickets of the Pacific
lowlands and some other regions, ascending to about 1,100 meters;
Alta Verapaz; Jutiapa; Santa Rosa; Escuintla; Guatemala; Chi-
maltenango; Suchitepe"quez; Retalhuleu; Quezaltenango; San Mar-
cos. Widely distributed in tropical and subtropical regions, Florida
and central Mexico to Panama, and southward through much of
South America; southern coast of Europe to Asia and Africa;
probably introduced in America.

A coarse, erect or decumbent annual or perennial with branched stems, often
a meter high, the stems quadrangular, pilose; leaves on petioles 5-25 mm. long,
oval to ovate, 5-20 cm. long, 2-9 cm. wide, rather abruptly acuminate or long-
acuminate, obtuse to abruptly acuminate at the base, thin, green and pilose-
strigose on the upper surface, paler beneath and pilose-sericeous, often densely so;
flower spikes terminal and axillary, 4-30 cm. long, 10-12 mm. thick, the rachis
densely white- villous; bracts and bractlets glabrous, ovate, long-aristate; sepals
lanceolate, 6-7 mm. long, acuminate, not nerved, glabrous; utricle truncate at
the apex, glabrous.

Called "zorrillo bianco" in Yucatan, "abrojo" in Salvador, and
"mozote" in Salvador and Honduras. The Maya name is reported
from Yucatan as "zacpaiche"." The plant is an annoying weed,
often abundant in waste places. The sharp tips of the sepals pene-
trate the skin easily if the plant is handled carelessly.

Achyranthes indica (L.) Mill. Gard. Diet. ed. 8. No. 2. 1768.
A. aspera var. indica L. Sp. PL 204. 1753. Centrostachys indica
Standl. Journ. Wash. Acad. Sci. 5: 75. 1915. Pegapega; Goncilla
(Zacapa); Mozotlexc (Pete"n; apparently a combination of Spanish
and Maya).

A weed in moist or dry fields or thickets, 400 meters or lower;
reported from Pete"n; Zacapa; El Progreso. Southeastern United
States; Honduras to Panama; West Indies; South America; Africa,
Asia, and the Pacific islands; in America doubtless imported from
the Old World tropics.

An erect or spreading annual, the stems 2 meters long or usually much shorter,
simple or branched, the stems terete or obscurely quadrangular, whitish-pilose;
leaves on petioles 3-15 mm. long, rhombic-orbicular or obovate-orbicular, 2-7
cm. long and nearly or quite as wide, rounded at the apex and often abruptly
acute or acutish, rounded to cuneate at the base, pilose-sericeous on both surfaces
or glabrate above; flowers green, the spikes terminal, 10-40 cm. long, 6-7 mm.
thick, the rachis pilose or villous; bracts broadly ovate or orbicular, the midnerve

146 FIELDIANA: BOTANY, VOLUME 24

indurate and extended into a rigid spine as long as the body of the bract or longer;
bractlets ovate, long-aristate, shorter than the perianth; sepals narrowly lanceo-
late, 4 mm. long, acuminate, not nerved; utricle oblong, truncate at the apex,
glabrous.

This species is scarce in Central America and known only from
the Atlantic coast, while A. aspera is widely distributed and often
abundant locally.

ALTERNANTHERA Forskal

Herbs or shrubs, prostrate, erect, or scandent; leaves opposite, petiolate or
sessile, entire or nearly so; flowers perfect, bracteate and bibracteolate, capitate
or spicate, usually compressed, the heads or spikes few or numerous, sessile or
pedunculate, axillary or terminal; perianth sessile or stipitate, the sepals unequal,
glabrous or pubescent; filaments united into a short or elongate tube, this with 3-5
antheriferous lobes and as many intervening entire or variously laciniate or dentate,
short or elongate staminodia, or the staminodia rarely absent; anthers short or
elongate, 2-celled; ovary globose to ovoid or obovoid; style short or elongate, the
stigma capitate; ovule 1, pendulous from an elongate funicle; utricle membrana-
ceous, indehiscent; seed inverted, smooth, the embryo annular.

About 100 species, in tropical America and Australia. About
30 are known from North America, and several species besides those
listed here occur in southern Central America.

Flower heads on elongate naked peduncles.

Sepals glabrous A. microcephala.

Sepals pilose or pubescent.

Flowers sessile or nearly so within the bractlets A. laguroides.

Flowers conspicuously pedicellate or stipitate within the bractlets, the stipe
articulate, 5-sulcate.

Bractlets conspicuously longer than the sepals A. dentata.

Bractlets much shorter than the sepals.

Stems strigillose or glabrate; bractlets not cristate A. ramosissima.

Stems pilose with ascending or spreading hairs; bractlets conspicuously

cristate A. brasiliana.

Flower heads sessile or nearly so.

Utricle equaling or exceeding the sepals, emarginate A. sessilis.

Utricle much shorter than the sepals, not emarginate.

Outer bracts usually laciniate-lobate; petioles equaling or at least half as
long as the blades; leaves usually tinged with red, purple, or yellow.

A. BeUzickiana.

Outer bracts not laciniate; petioles less than half as long as the blades; leaves
green.

Sepals glabrous A. obovata.

Sepals pilose or villous.

Young leaves grayish, pubescent with branched hairs; flower heads when
well developed at least twice as long as thick; plant of saline soil.

A. halimifolia.

Young leaves not pubescent with branched hairs; flower heads less than
twice as long as thick; plants not of saline soil.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 147

Leaves rounded to acute at the apex, mostly 2.5 cm. long or less.

Sepals merely acute, muticous A. polygonoides.

Sepals acuminate, usually conspicuously mucronate A. repens.

Leaves long-acuminate, mostly 10-15 cm. long A. megaphylla.

Alternanthera Bettzickiana (Regel) Standl. Field Mus. Bot.
3: 254. 1930. Telanthera Bettzickiana Regel, Gartenflora 11: 178.
1862. A. spathulata Lem. 111. Hort. 12: pi. 445. 1865. Telanthera
picta C. Koch, Wochenschr. Gartn. 9: 15. 1866. Achyranthes
Bettzickiana Standl. N. Amer. Fl. 21: 138. 1917. Hierba te; Adorno;
Hierbilla.

Planted commonly for ornament in all except the higher regions
of Guatemala and perhaps more or less naturalized in some localities.
Described from Brazil, but probably unknown in a wild state,
although cultivated in tropical regions.

Plants annual or perennial, usually erect, commonly less than 40 cm. high,
often densely branched, the stems swollen at the nodes, villous when young but
soon glabrate; petioles slender, equaling or shorter than the leaf blades; blades
rhombic, rhombic-ovate, or rhombic-obovate, 1-3.5 cm. long, 1-1.7 cm. wide,
acuminate or abruptly acute, abruptly long-attenuate at the base, undulate or
crispate, sparsely appressed-pilose when young but soon glabrate, green or usually
purplish red or yellowish, often variegated; heads axillary, sessile, ovoid or oblong,
whitish; bracts and bractlets broadly ovate, aristate-acuminate, at least the lower
bracts laciniate-lobate, glabrous, half as long as the sepals; sepals lance-oblong,
acute or acuminate and mucronate, 3-nerved, sparsely pilose; staminodia equaling
the filaments, laciniate at the apex.

Called "perico" in Salvador and "colchon de nirio" in Honduras.
Known by the name "coqueta" in British Honduras, where there is
a belief that leaf -cutting ants will not pass "through, under, or
over" the plant. This species is well known in the United States
where it often is grown in pots, or more frequently in outdoor beds
in making formal designs. It is used in the same manner in the
parks of Guatemala. The species probably is one of the American
plants that has been long in cultivation, and has arisen from A.
ficoidea, from which it differs but little.

Alternanthera brasiliana (L.) Kuntze, Rev. Gen. 537. 1891.
Gomphrena brasiliana L. Cent. PI. 2: 13. 1756. Telanthera brasiliana
Moq. in DC. Prodr. 13, pt. 2: 382. 1849. A. brasiliana var. sericea
Kuntze, Rev. Gen. 2: 538. 1891. Achyranthes brasiliana Standl.
Journ. Wash. Acad. Sci. 5: 74. 1915.

Collected in British Honduras and doubtless extending into
Pete*n or Izabal. Southern Mexico; Brazil.

148 FIELDIANA: BOTANY, VOLUME 24

A much branched perennial, probably clambering, the slender branches
pilose with ascending or spreading hairs, sometimes glabrate; leaves on petioles
3-10 mm. long, oblong-ovate to oval or oblong, 4-10 cm. long, 1-4 cm. wide,
acuminate, rounded to acute at the base, pilose or pilose-sericeous; peduncles
usually simple, pilose, 2-10 cm. long; spikes globose, 8-12 mm. thick, the flowers
stramineous or whitish; bracts nearly as long as the bractlets, oblong-ovate, long-
acuminate, glabrous; bractlets half as long as the sepals, ovate-oblong, long-acumi-
nate, often denticulate, usually narrowly cristate near the apex, the crest denticu-
late; sepals ovate-lanceolate or lance-oblong, 3-4 mm. long, rigid, prominently
nerved, acute, short-pilose; pedicels 1 mm. long; staminodia longer than the fila-
ments, laciniate at the apex.

The type of Kuntze's var. sericea was collected somewhere in
Guatemala, Keck 416.

Alternanthera dentata (Moench) Stuchl. ex Fries, Arkiv Bot.
16, no. 13: 11. 1921. Gomphrena brasiliensis Jacq. Coll. Bot. 2: 278.
1788, not L. 1756. G. dentata Moench, Meth. Suppl. 273. 1802.
Mogiphanes Jacquini Schrad. Ind. Sem. Goetting. 4. 1834.

Dry or moist, often rocky, brushy hillsides, 1,000-1,100 meters;
El Progreso (between San Geronimo and Morazan, near Baja
Verapaz boundary); Guatemala (Fiscal). Salvador; West Indies;
South America.

A suberect or straggling, perennial herb about a meter high, sometimes sub-
scandent, the stems appressed-pilose or glabrate; leaves slender-petiolate, oval or
ovate to oblong, mostly 4-10 cm. long, acute or acuminate, abruptly acute at the
base, thin, sparsely or densely appressed-pilose or sericeous, sometimes glabrate;
peduncles simple or trifid, elongate; flower heads globose or short-cylindric, 1-2:5
cm. long, about 1 cm. broad; bracts short, white, long-acuminate; bractlets usually
longer than the sepals, oblong, acute, villous, cristate dorsally, the crest serrulate;
sepals lance-oblong, rigid, 3-nerved, acute, appressed-pilose, 3-3.5 mm. long;
staminodia longer than the filaments, ligulate, lacerate at the apex.

Alternanthera halimifolia (Lam.) Standl. ex Pittier, PI.
Usual. Venez. 145. 1926. Achyranthes halimifolia Lam. Encycl. 1:
547. 1785. Alternanthera asterotricha Uline, Field Mus. Bot. 1 : 419.
1899. Telanthera halimifolia A. Stewart, Proc. Calif. Acad. Sci. IV.
1:58. 1911.

In saline soil, along or near beaches, Champerico, Retalhuleu,
and probably in other Pacific departments. Yucatan (whence the
type of A. asterotricha); Panama; West Indies; Venezuela to Colom-
bia and Chile.

A prostrate perennial, often suffrutescent at the base, the stems a meter long
or less, simple or branched, pubescent with short, closely appressed, grayish,
branched or hispidulous hairs; leaves on petioles 2-8 mm. long, oblong to oval

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 149

or obovate-oblong, 1.5-6 cm. long, 1-2 cm. wide, usually rounded at the apex,
rather thick and fleshy, soon glabrate above, beneath densely pubescent with short
hispidulous hairs, in age sometimes glabrate; heads mostly axillary, sessile, solitary
or glomerate, short-cylindric or ovoid, 2 cm. long or shorter, stramineous;
bracts and bractlets half as long as the sepals, ovate, acuminate and mucronate,
appressed-pilose; sepals 3-4 mm. long, ovate-oblong, acute, 3-5-nerved, densely
pubescent; staminodia ligulate, longer than the filaments, laciniate at the apex.

This species appears to be confined to sea beaches or to inland
localities where the soil is strongly alkaline.

Alternanthera laguroides Standl. in Standl. & Cald. Lista
PI. Salvador 74. 1925. Achyranthes laguroides Standl. Contr. U. S.
Nat. Herb. 18: 90. 1916. Botoncito.

Dry or moist thickets, 700-1,500 meters; Santa Rosa; Escuintla;
Guatemala. Thickets of the Pacific slope, Guatemala to Panama.

Plants perennial, slender, branched, often clambering over shrubs, the stems
pilose-strigose or glabrate; leaves on very short petioles, narrowly lanceolate to
oblong-lanceolate or linear-lanceolate, 4-8 cm. long and 3-15 mm. wide or some-
times larger, acuminate or attenuate at each end, pilose-sericeous, densely so
beneath; peduncles simple or branched, 1-3 cm. long, or some of the heads sessile
or subsessile; spikes ovoid or cylindric, 1-2 cm. long and almost 1 cm. thick,
whitish-stramineous; bracts and bractlets ovate-triangular, half as long as the
sepals, acuminate or long-acuminate, sparsely pilose or glabrate; sepals linear-
oblong, 4-5 mm. long, acuminate, membranaceous, 1-nerved, pilose near the base
with straight erect nodulose white hairs; staminodia ligulate, longer than the
anthers, laciniate at the apex.

Alternanthera megaphylla Standl. Field Mus. Bot. 8: 9. 1930.
Achyranthes megaphylla Standl. N. Amer. Fl. 21: 141. 1917.

Wet mixed forest, at or near sea level; Izabal (Rio Bonito, Cerro
San Gil, Steyermark 41690). Costa Rica.

An erect or decumbent, perennial herb, often forming colonies, the stems
mostly 50 cm. long or shorter, often geniculate at the base and rooting at the lower
nodes, generally simple, appressed-pilose when young; leaves on petioles 3-13
mm. long, oval to lance-oblong, 10-17 cm. long, 3.5-7.5 cm. wide, gradually or
abruptly long-acuminate, rounded to acute and long-decurrent at the base,
rather succulent, dark olivaceous when dry, glabrous above, sparsely appressed-
pilose beneath; flower spikes axillary and terminal, sessile, solitary, about 2 cm.
long and 1.5 cm. broad, the flowers brown; bracts and bractlets half as long as the
perianth, ovate, long-attenuate, short-pilose, with rigid tips; sepals lance-oblong,
6-7 mm. long, 3-5-nerved, short-pilose; stamen tube short, the staminodia ligulate,
exceeding the anthers, pectinate-laciniate nearly to the base; style elongate; seed
ovoid, 2.5 mm. long, lustrous, reddish brown.

The species has not been collected along the Atlantic coast of
Honduras and Nicaragua but is to be expected there.

150 FIELDIANA: BOTANY, VOLUME 24

Alternanthera microcephala (Moq.) Schinz in Engl. & Prantl,
Pflanzenfam. ed 2. 16C: 75. 1934. Brandesia mexicana Schlecht.
Linnaea 7: 392. 1832. Telanthera microcephala Moq. in DC. Prodr.
13, pt. 2: 371. 1849. Telanthera mexicana Moq. op. cit. 372. Alter-
nanthera mexicana Hieron. Bot. Jahrb. 20: Beibl. 49: 8. 1895, not
A. mexicana Moq. 1849. Achyranthes mexicana Standl. Journ.
Wash. Acad. Sci. 5: 74. 1915.

Dense wet mixed forest, region of Tactic, Alta Verapaz, and
below, 600-1,600 meters. Southern Mexico; Panama.

Plants herbaceous, probably perennial, erect and a meter high or less, usually
rooting at the lower nodes, branched, the stems pilose with spreading or retrorse
hairs; leaves on petioles 2 cm. long or less, thin, ovate or elliptic, 3-10 cm. long
and 1-5 cm. wide or somewhat larger, rather abruptly long-acuminate, at the base
acute or obtuse, appressed-pilose on both surfaces with long slender hairs; pedun-
cles axillary, simple, filiform, 2-6.5 cm. long, sparsely pilose or glabrate; spikes
short-cylindric or subglobose, 5-10 mm. long, 5-7 mm. thick; bracts broadly ovate,
acute, subscarious, glabrous; bractlets broadly ovate, half as long as the sepals,
long-aristate, villous along the nerves, greenish white or stramineous; sepals nar-
rowly oblong, 2.5-3.5 mm. long, acute or acutish, membranaceous, greenish white
or stramineous, 3-nerved, glabrous; staminodia longer than the anthers, laciniate
at the apex.

Alternanthera obovata (Mart. & Gal.) Millsp. Field Mus. Bot.
1: 360. 1898. Bucholzia obovata Mart. & Gal. Bull. Acad. Brux. 10,
pt. 1: 348. 1843. Telanthera obovata Moq. in DC. Prodr. 13, pt. 2:
370. 1849. Achyranthes obovata Standl. Journ. Wash. Acad. Sci. 5:
74. 1915.

Wet soil, in fields or ditches, most often at the edges of streams
or ponds, ascending to 1,400 meters but mostly at much lower
elevations; Pete"n; Izabal; Alta Verapaz; Escuintla; Suchitepe'quez ;
Retalhuleu. Mexico and British Honduras to Honduras.

Perennial, suberect or usually decumbent or prostrate, the stems rather stout,
simple or branched, a meter long or usually shorter, densely villous when young
but soon glabrate; leaves on very short petioles, rounded-obovate to oval or oblong,
1.5-4.5 cm. long, 1-2 cm. wide, usually rounded at the apex, cuneate to rounded
at the base, rather thick and bright green when dry, villous when young but in
age almost glabrous; spikes axillary and terminal, sessile, subglobose or cylindric,
1.2-3.5 cm. long, 1 cm. thick, white; bracts and bractlets broadly ovate, half as
long as the sepals, acuminate, mucronulate, glabrous; sepals oblong, 4 mm. long,
acute, 1-nerved, serrulate at the apex, glabrous; staminodia linear, acutish, entire,
longer than the filaments.

Alternanthera polygonoides (L.) R. Br. Prodr. 417. 1810.
Gomphrena polygonoides L. Sp. PI. 225. 1753. A. paronychioides St.
Hil. Voy. Distr. Diam. 2: 439. 1833. Telanthera polygonoides Moq.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 151

in DC. Prodr. 13, pt. 2: 364. 1849. Achyranthes polygonoides Lam.
Encycl. 1: 547. 1785.

Pete"n (Lake Zotz, Lundell). Mexico and British Honduras to
Panama, southward to Brazil; West Indies.

A prostrate perennial, often forming dense mats, the stems branched, mostly
10-20 cm. long, often rooting at the nodes, white-villous when young, glabrate in
age; leaves short-petiolate, oval to elliptic or ovate-rhombic, 1-2.5 cm. long, 3-11
mm. wide, acute or obtuse, at the base acuminate or attenuate, glabrous or nearly
so above, densely villous beneath when young but soon glabrate; heads axillary,
sessile, solitary or glomerate, white, usually as broad as long; bracts and bractlets
half as long as the sepals or shorter, ovate, acute and mucronate, glabrous; sepals
oblong-lanceolate, 4 mm. long, acutish, 3-nerved, glabrous or practically so, often
sparsely pilose below; staminodia much shorter than the filaments, ovate, denticu-
late; utricle orbicular, almost half as long as the sepals.

Alternanthera ramosissima (Mart.) Chodat, Bull. Herb.
Boiss. II. 3: 355. 1903. Mogiphanes ramosissima Mart. Nov. Gen.
& Sp. 2: 36. 1826. Telanthera ramosissima Moq. in DC. Prodr. 13,
pt. 2: 381. 1849. Achyranthes ramosissima Standl. Journ. Wash.
Acad. Sci. 5: 74. 1915.

Pete"n (Lago de Pete*n). Southeastern Mexico to British Hon-
duras; southern Florida; West Indies; Guianas and Brazil.

Plants perennial, slender, branched, often clambering over shrubs, much
branched, the branches sparsely strigose or glabrate; leaves on very short petioles,
lanceolate to ovate, 2-8 cm. long, 3 cm. wide or less, long-acuminate or acute,
rounded to acuminate at the base, sparsely strigose or glabrate; peduncles simple,
2-10 cm. long, strigillose above; spikes subglobose or short-cylindric, 1-2.5 cm.
long, 1 cm. thick; bracts broadly ovate, acute or subacute, shorter than the bract-
lets, glabrous; bractlets triangular-ovate, acuminate, one-third as long as the
sepals, glabrous; sepals narrowly oblong or lance-oblong, 4-5 mm. long, acute,
short-mucronate, short-pilose with appressed or spreading hairs; staminodia much
longer than the filaments, ligulate, laciniate at the apex.

Alternanthera repens (L.) Kuntze, Rev. Gen. 536. 1891.
Achyranthes repens L. Sp. PI. 205. 1753. Alternanthera Achyrantha
R. Br. Prodr. 417. 1810. Sanguinaria; Hierba de toro (Guatemala) ;
Sacachiquim (Colomba).

Most abundant among cobblestones in streets of cities and
villages but growing also in open grassy places, often in sandy
stream beds; Alta Verapaz; Pete"n; Jutiapa; Santa Rosa; Guate-
mala; Sacatepe"quez ; Chimaltenango; Huehuetenango; Quezalte-
nango; San Marcos. Mexico to Panama, southward to Argentina;
southeastern United States; West Indies; southern Europe, Asia,
East Indies.

152 FIELDIANA: BOTANY, VOLUME 24

Plants perennial, prostrate, often forming small mats, much branched, the
stems white- villous, sometimes glabrate in age; leaves of a pair usually unequal,
short-petiolate, often crowded, rhombic-ovate to elliptic or obovate, 5-25 mm.
long, 3-15 mm. wide, obtuse, at the base acute or acuminate, sparsely villous when
young but soon glabrate; heads ovoid or short-cylindric, dirty-white, 5-15 mm.
long, 5-8 mm. thick, axillary, sessile, often glomerate; bracts and bractlets shorter
than the sepals, ovate, pungent-mucronate, glabrous or pilose, the margins usually
ciliate-denticulate; sepals very unequal, the outer ones oval or broadly ovate,
3-5 mm. long, acute and short-aristate, 3-nerved, villous along the nerves, especially
near the base, the inner sepals linear-subulate; staminodia usually shorter than the
filaments, triangular or subulate, entire or rarely denticulate.

The Maya name is reported from Yucatan as "cabalxtez." In
almost any Guatemalan town this plant may be found growing
abundantly among the cobblestones with which most streets are
paved, and every year thousands of small boys spend weary days
digging it and other small weeds from the streets in preparation for
Holy Week and other fiestas. In spite of long-continued eradica-
tion, the plant continues to thrive, just as it doubtless has done
for two hundred years or more. Possibly it was introduced into
Central America from Spain.

Alternanthera sessilis (L.) R. Br. Prodr. 417. 1810. Gomphrena
sessilis L. Sp. PI. 225. 1753. Achyranthes sessilis Steud. ex Standl.
Journ. Wash. Acad. Sci. 5: 73. 1915.

A weed in wet or moist thickets, open pastures, and waste
ground, 400 meters or less; Izabal; Zacapa; Chiquimula; Santa Rosa;
Escuintla; Suchitepe"quez. British Honduras to Panama; West
Indies; Guianas and Brazil; widely distributed in Old World tropics.

Procumbent annual or perennial, the stems 20-60 cm. long, often rooting at
the nodes, simple or sparsely branched, puberulent in lines or glabrate; leaves
short-petiolate, elliptic to oblong-obovate or spatulate-obovate, 1-2.5 cm. long,
5-20 mm. wide, rounded to acuminate at the apex, cuneate at the base, bright
green, glabrous, or sparsely villous beneath along the nerves; heads axillary, sessile,
solitary or glomerate, subglobose, bright white; bracts and bractlets ovate,
mucronate, one-third to half as long as the sepals, glabrous; sepals broadly ovate,
1.5 mm. long, acute, hyaline, 1-nerved, glabrous; staminodia equaling the fila-
ments, subulate, entire.

This has smaller flowers and heads than any other Central Ameri-
can species. It is easily recognized also by the broad obcordate
utricle, which projects slightly beyond the calyx.

AMARANTHUS L.

Annual herbs, erect or prostrate, glabrous or pubescent, usually branched;
leaves alternate, petiolate, entire or undulate, often mucronate; flowers small,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 153

monoecious, dioecious, or polygamous, bracteate and bibracteolate, glomerate,
the glomerules axillary, spicate, or paniculate; sepals 5 or rarely 1-3, membrana-
ceous, equal or subequal, sometimes indurate at the base after anthesis, erect in
fruit; stamens normally 5, the filaments distinct, filiform or subulate; anthers
oblong or linear-oblong, 4-celled; ovary ovoid, compressed, circumscissile or open-
ing irregularly, membranaceous or coriaceous, sometimes 2-3-dentate at the apex;
seed erect, compressed, smooth, the embryo annular.

About 50 species, in temperate and tropical regions of the whole
earth. Probably no other species occur in Central America but
about 40 are known from all North America.

Plants armed with spines A. spinosus.

Plants unarmed.

Flowers all clustered in the leaf axils A. polygonoides.

Flowers in terminal, usually paniculate spikes.

Fruit indehiscent, rugose A. viridis.

Fruit dehiscent, smooth.

Sepals of the pistillate flowers spatulate, contracted below into a narrow

claw, more or less urceolate in age A. scariosus.

Sepals of the pistillate flowers oblong to obovate, not contracted into a

claw, not urceolate.
Bracts equaling or shorter than the flowers; pistillate sepals usually

obtuse, equaling or usually longer than the fruit A. dubius.

Bracts longer than the flowers; pistillate sepals sometimes shorter than

the fruit.
Inflorescence deep red or purple; bracts less than twice as long as the

sepals A. caudatus.

Inflorescence green or faintly tinged with red or purple; bracts usually
twice as long as the sepals or longer A. hybridus.

Amaranthus caudatus L. Sp. PI. 990. 1753. A. cruentus L.
Syst. Veg. ed. 10. 1269. 1759. A. paniculatus L. Sp. PL ed. 2. 1406.
1763. A. sanguineus L. Sp. PI. ed. 2. 1407. 1763. A. leucospermus
Wats. Proc. Amer. Acad. 22: 446. 1887. Moco de chumpe (Zacapa);
Cola de zorro; Bledo cimarron (Coban); Bledo extranjero (Coban);
Ses (Quecchi) ; Bledo rojo.

Commonly cultivated, in its various forms, in gardens for orna-
ment, also occurring as a weed in gardens, cornfields, and waste
places; Alta Verapaz; Izabal; Zacapa; Jutiapa; Sacatepe"quez ;
Chimaltenango; Quiche"; Retalhuleu; Quezaltenango; San Marcos;
Huehuetenango; probably found, at least in gardens, in all the
departments. Widely distributed in the tropics of both hemi-
spheres, but principally in cultivation, or escaping; probably native
of the American tropics, but the original habitat unknown.

Plants stout, erect, commonly 1-1.5 meters tall, simple or much branched,
often colored almost throughout with red-purple or deep red, usually pubescent,

154 FIELDIANA: BOTANY, VOLUME 24

villous about the inflorescence; leaves on slender petioles 2-20 cm. long, elliptic
to ovate-lanceolate or rhombic-ovate, 5-30 cm. long, 2-10 cm. wide, attenuate to
acute, at the base acute to attenuate, sparsely pubescent or glabrate; flowers
monoecious, in dense panicles, these composed of numerous slender spreading
lateral spikes 4-18 cm. long and usually 6-8 mm. thick, the terminal spikes usually
twice as long as the lateral ones, erect or more often recurved or pendent; bracts
lanceolate to ovate, equaling or half longer than the sepals; pistillate sepals oblong,
1.5 mm. long, obtuse or rounded at the apex; stamens 5; style branches 3; utricle
conspicuously exceeding the sepals, circumscissile at the middle; seeds 1 mm. in
diameter, black, yellowish white, or red.

Called "pison calaloo" in British Honduras, and "amaranto"
and "chichimeca" in Salvador. About Coban the seeds are used with
panela to make a dulce or sweetmeat called "boroco." In Mexico
this species is sometimes cultivated or at least the seeds are gathered,
perhaps from partly wild plants, and used as food in the form of
atol or mush. Various specific names have been applied to this
plant but all the names listed above, as well as a good many more,
seem to relate to a single major species, which is itself rather doubt-
fully distinct from A. hybridus. It seems preferable to treat all
these red or purple forms as belonging to a single species rather
than attempt to separate them by minute characters, as was done
in North American Flora and is often the practice among European
writers.

Amaranthus dubius Mart. PI. Hort. Erlang. 197. 1814. A.
tristis Willd. Hist. Amaranth. 21. 1790, at least in part, not A. tristis
L. 1753. Chic-ixtez, Acilixtez (Pete"n, Maya, fide Lundell).

Pete"n (Uaxactun); doubtless also in Izabal, although no speci-
mens have been seen. Yucatan and British Honduras to Panama,
southward through tropical South America; West Indies; adventive
in Europe.

Plants stout and succulent, usually about 60 cm. tall, simple or much branched,
glabrous or nearly so; leaves on petioles 2-9 cm. long, ovate or rhombic-ovate,
4-12 cm. long, 2-8 cm. wide, acute to rounded at the apex, the tip usually emargi-
nate, rounded to acutish at the base, glabrous or nearly so; flowers monoecious,
green or whitish, chiefly in paniculate, often drooping spikes 5-25 cm. long and
4-12 mm. thick; bracts ovate or oval, acute, scarious, pungent-tipped, usually
shorter than the sepals; pistillate sepals oblong to ovate, 1.5-2 mm. long, obtuse
or acutish, often emarginate, mucronate, scarious; stamens 5; style branches 3;
utricle usually exceeding the sepals, dehiscent at the middle; seed 1 mm. in diame-
ter, lustrous, dark reddish brown or black.

Called "bledo de Jamaica" on the north coast of Honduras, where
the plant is believed locally to have been introduced by immigrating
Jamaicans. Maya names reported from Yucatan are "xetz" and

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 155

"chactez." The leaves of this as well as those of other species often
are gathered and used as a pot herb.

Amaranthus hybridus L. Sp. PI. 990. 1753. A. hypocondria-
cus L. Sp. PI. 991. 1753. A. chlorostachys Willd. Hist. Amaranth.
34. 1790. Bledo (often corrupted to Blero) ; Ses (Quecchi of Coban) ;
Huisquelite; Huisquilete.

A common weed in cultivated or waste ground, often abundant
in cornfields, cafetales, or thickets, mostly at 400-2,500 meters and
probably ascending even higher; Alta Verapaz; Jalapa; Jutiapa;
Santa Rosa; Escuintla; Sacatepe'quez; Chimaltenango; Solola;
Suchitepe'quez ; Retalhuleu; Quezaltenango; Huehuetenango; prob-
ably also in all or most of the other departments. Generally dis-
tributed in temperate and tropical regions of the New World;
adventive in many regions of the Old World; probably native in
America.

Plants stout, erect, sometimes 2 meters tall but usually a meter or less, often
much branched, rough-puberulent or glabrous below, usually sparsely villous
above, the stems striate or sulcate; leaves on slender petioles 9 cm. long or less,
lanceolate to ovate or ovate-rhombic, 5-15 cm. long, 2-7 cm. wide, acute to rarely
rounded at the apex, pubescent beneath or glabrous, often slightly tinged with
red; flowers monoecious, spicate, the spikes paniculate, the terminal one twice as
long as the lateral ones or shorter, 6-12 mm. thick; bracts twice as long as the
sepals, lanceolate to ovate, with a spinose tip; pistillate sepals 5, oblong, 1.5-2
mm. long, acute, or the inner sometimes obtuse, equaling or shorter than the fruit;
stamens 5; style branches 3; utricle thin-walled, circumscissile at the middle; seeds
1 mm. in diameter, dark reddish brown or black, shining.

The Maya name used in Yucatan is "xtez." In that state, as
well as in other parts of Mexico, the plant is known by the name
"quelite," a word of Nahuatl origin, applied generally to leaves
cooked and used as food. Amaranthus hybridus is especially abund-
ant on the Pacific plains, where it often forms extensive and dense,
tall stands, especially in old cornfields. The Quiche* name of Guate-
mala is reported by Tejada as "quiec tes."

Amaranthus polygonoides L. PI. Jam. Pugill. 27. 1759.

Zacapa, about 200 meters, moist fields. British Honduras, and
probably to be found in the adjacent departments of Guatemala.
Mexico; West Indies and northern South America; Florida and
Texas.

Stems slender, ascending or spreading, sometimes erect, 10-50 cm. long,
much branched from the base, villous about the inflorescence; leaves on petioles
2.5 cm. long or shorter, rhombic-ovate to obovate or oval, 1-3 cm. long, obtuse to

156 FIELDIANA: BOTANY, VOLUME 24

subtruncate and usually emarginate at the apex, acute or cuneate at the base and
decurrent, sparsely pubescent beneath or glabrous; flowers monoecious, in dense
sessile several-flowered axillary clusters; bracts lanceolate, acuminate, half as
long as the sepals or less; pistillate sepals spatulate, erect, obtuse or rounded at the
apex, often apiculate, 3-nerved, scarious, united at the base; stamens 2-3; style
branches 2-3; utricle circumscissile; seed black or dark brown, lustrous, 0.6-0.9
mm. in diameter.

From Yucatan the Maya name is listed as "sacxtez."

Amaranthus scariosus Benth. Bot. Voy. Sulph. 158. pi. 51.
1844. Bledo; Huisquilete.

Weedy fields, 325 meters or less; Zacapa; Santa Rosa. South-
western Mexico along the Pacific lowlands to Costa Rica; type from
Tigre Island, Golfo de Fonseca, Honduras.

Plants stout, 1-1.5 meters high or even taller, often much branched, the stems
glabrous or sparingly pubescent above; leaves on slender petioles 10 cm. long or
less, ovate or oblong-ovate, 6-12 cm. long, acute, the tip rounded, at the base
acute or abruptly acute, glabrous; flowers monoecious, spicate, the spikes 8-20
cm. long, erect or drooping, forming e large panicle; bracts subulate-lanceolate,
pungent-tipped, slightly exceeding the flowers; pistillate sepals 5, spatulate, 3 mm.
long, rounded at the apex, often retuse, scarious, 1-nerved, united at the base;
stamens 5; style branches 3; utricle much shorter than the sepals, circumscissile;
seed black, 0.8 mm. in diameter.

This species is decidedly limited in distribution, being confined,
so far as known, to the region indicated above. It is quite as weedy
as other members of the genus.

Amaranthus spinosus L. Sp. PI. 991. 1753. Huisquelite (of
Nahuatl derivation, signifying "spiny quelite"); Bledo macho; Ixtez
(Pete"n, Maya); Tsetz, Labtzetz (Quiche"); Bledo; Nigua (Zacapa).

A common weed found in waste or cultivated ground, or often
in thickets, chiefly in the lowlands but ascending sometimes to
about 1,800 meters; Pete"n; Alta Verapaz; Izabal; Zacapa; Jalapa;
Jutiapa; Santa Rosa; Escuintla; Sacatepe"quez; Retalhuleu; Que-
zaltenango; San Marcos. Generally distributed in tropical America,
and in many parts of the United States; also in the Old World
tropics; probably native in America.

Plants stout and succulent, erect or ascending, commonly 50-70 cm. tall,
glabrous below, more or less pubescent above, each axil provided with 2 rigid
sharp-pointed spines 2.5 cm. long or less; leaves slender-petiolate, ovate to rhombic-
ovate or lanceolate, 3-12 cm. long, acute at the base, narrowed toward the apex,
but the tip obtuse or broadly rounded, glabrous or sparsely pubescent; flowers
monoecious, the pistillate in dense, globose, sessile, chiefly axillary clusters, the
staminate in slender, erect or drooping, terminal spikes 3-18 cm. long and 4-8 mm.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 157

thick; bracts lanceolate or subulate, often spinose, shorter than the sepals or often
2-3 times as long; pistillate sepals 5, oblong, obtuse or acute, 1.5 mm. long; stamens
5; style branches 3; utricle about equaling the sepals, irregularly and imperfectly
circumscissile; seed black, lustrous, 0.7-1 mm. in diameter.

The Maya name in Yucatan is "xtez" or "kix-xtez." The leaves
and young shoots of this species are cooked and eaten, but less com-
monly perhaps than those of A. hybridus. Bunches of the young
shoots of the various Amaranthus species are offered for sale com-
monly in Guatemalan markets.

Amaranthus viridis L. Sp. PI. ed. 2. 1405. 1763. A. gracilis
Desf. Tabl. Bot. 43. 1804.

A weed in moist ground about dwellings, at or little above sea
level; Zacapa; Retalhuleu. Florida; Mexico; British Honduras;
Honduras; West Indies; South America; widely distributed in tropi-
cal regions of both hemispheres.

Stems rather slender, erect or procumbent, usually 20-50 cm. long, often
much branched, glabrous; leaves slender-petiolate, ovate or rhombic-ovate, 2-8
cm. long, rounded or narrowed at the apex and emarginate, rounded to broadly
cuneate at the base, glabrous; flowers monoecious, in slender, terminal, often
paniculate spikes 4-12 cm. long and 4-8 mm. thick; bracts ovate or lanceolate,
acute, much shorter than the flowers; sepals 3, oblong or linear-oblong, acute or
obtuse, cuspidate, 1-1.5 mm. long, equaling or shorter than the fruit; stamens 3;
style branches 3; utricle globose, strongly rugose; seed 1 mm. in diameter, black
or dark reddish brown, dull.

Called "bledo" in Honduras, and doubtless the same name is
used in Guatemala if any name is given the plant.

CELOSIA L.

Annual or perennial herbs or shrubs, pubescent or glabrous, usually erect;
leaves alternate, generally petiolate, entire or rarely lobate; flowers perfect,
bracteate and bibracteolate, in dense, terminal or axillary spikes, or fasciculate
along the simple or branched flowering branches, sessile or pedicellate; perianth
5-parted, the segments scarious, striate-nerved; stamens 5, the filaments subulate
or filiform, connate at the base into a short cup; anthers 4-celled; ovary subglobose
to cylindric, the style elongate, short, or none; stigmas 2-3, subulate or capitate;
ovules 2 or more; utricle included in the perianth or exserted, sometimes indurate
at the apex, circumscissile, rarely indehiscent or rupturing irregularly; seeds 2 to
many, usually erect, lenticular, smooth and lustrous, the embryo annular.

About 40 species, chiefly in Asia and Africa. Eight are found
in North America but only the following are known to occur in
Central America.

158 FIELDIANA: BOTANY, VOLUME 24

Inflorescence of simple terminal spikes 15-20 mm. in diameter (much larger in
cultivated forms) ; sepals 6-9 mm. long, bright white, pink, or red, sometimes
yellow C. argentea.

Inflorescence of terminal or axillary panicles composed of few or numerous spikes
3-10 mm. in diameter; sepals 4-6 mm. long, at least in the dried state stra-
mineous to dark brown.

Seeds 5-8; leaf blades ovate to lanceolate, decurrent nearly to the base of the

petiole C. virgata.

Seeds about 20; leaf blades deltoid to triangular-lanceolate, short-decurrent.

C. nitida.

Celosia argentea L. Sp. PL 205. 1753. C. cristata L. loc. cit.
Abanico; Flor de memo; Amaranto; Mano de leon (Pete"n); Cresta de
gallo; Amor seco (British Honduras).

Cultivated commonly for ornament, and sometimes to be found
as an escape, as in Suchitepe"quez. Cultivated generally in temper-
ate and tropical regions.

An erect annual a meter high or less, simple or branched, the stems glabrous;
leaves slender-petiolate, linear to lanceolate or ovate, acute to attenuate or acumi-
nate, rounded and decurrent at the base or acute to attenuate, glabrous; flowers
subsessile, in dense spikes terminating the branches, the spikes oblong or elongate,
2-20 cm. long, 1.5-2 cm. thick; bracts lanceolate or ovate, half as long as the sepals
or shorter, acuminate, carinate, scarious; sepals 6-9 mm. long, acute, carinate,
thin, mostly white, pink, or red; utricle ovoid or subglobose, containing 3-8 seeds,
these 1.2 mm. in diameter, nearly black, lustrous.

The plant is believed to be a native of tropical America but it is
unknown in a truly wild state and seems unable to perpetuate itself
except under cultivation. Half -wild plants, as described above, have
relatively small inflorescences of simple spikes. The better-known
form is var. cristata (L.) Voss, the garden cockscomb, grown in most
parts of the earth for ornament. In this the inflorescence usually is
fasciate, broad, thick, and more or less ruffled. There are also forms
in which the panicles are rather feathery and dissected. The more
ordinary garden varieties of cockscomb are grown commonly in
Central American gardens and almost throughout Guatemala. In
some Guatemalan gardens improved varieties of the plant, imported
from Europe or North America, are cultivated. Other names
reported for this species are "mono" (Honduras); "San Jose,"
"terciopelo" (Salvador).

Celosia nitida Vahl, Symb. Bot. 3: 44. 1794.

Pete"n (Uaxactun, on Maya ruins). Eastern and southern
Mexico to Campeche and Yucatan; Florida keys and southwestern
Texas; West Indies; northern coast of South America.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 159

Stems slender, erect or clambering, as much as 1.5 meters long, sometimes
woody at the base, glabrous, green or glaucescent; petioles 5-20 mm. long, naked;
leaf blades deltoid to ovate or triangular-lanceolate, 2-7 cm. long, 1-4 cm. wide,
obtuse to acuminate, at the base obtuse or truncate, slightly decurrent, glabrous,
often with fascicles of smaller leaves in the axils; inflorescence lax, terminal, of
loosely flowered, sessile or pedunculate spikes 1-4 cm. long and 7-10 mm. thick;
bracts rounded-ovate, about one-fourth as long as the sepals, obtuse or acutish,
often ciliolate; sepals 5 mm. long, oblong or oval, acute or acutish, mucronulate,
firm, dark brown or yellowish when dry, prominently and finely nerved; utricle
equaling or shorter than the sepals; seeds about 20 and 1 mm. in diameter, black
and lustrous.

The Maya name "zabacpox" is reported from Yucatan.

Celosia virgata Jacq. Coll. Bot. 2: 279. 1788.

Dry or moist thickets of the lowlands of the Oriente, about
200-600 meters; Zacapa; Chiquimula; Escuintla. Mexico; Cuba
and Puerto Rico; northern South America.

Plants erect, a meter tall or less, usually herbaceous, sparsely branched, the
stems slender, glabrous, striate; petioles shorter than the blades, winged nearly
or quite to the base; leaf blades ovate to lanceolate or elliptic, 5-15 cm. long, 1.5-9
cm. wide, acuminate, at the base acute or abruptly acuminate, sparsely pubescent
beneath along the nerves or glabrous; panicles terminal and axillary, composed of
few sessile or pedunculate, dense spikes 1-5 cm. long and about 7 mm. thick;
bracts one-third to one-half as long as the sepals, lanceolate or ovate, abruptly
attenuate to a subulate tip, carinate, often ciliate; sepals 5-6 mm. long, lance-
elliptic, acuminate, dark brown when dry, green in the fresh state, prominently
nerved; utricle shorter than the sepals; seeds 5-8, nearly smooth, black, lustrous,
0.6 mm. in diameter.

Maya names recorded from Yucatan are "hatanal," "halalnal,"
and "hatalnal," all evidently intended to represent a single word.
From the same region the Spanish name of "zorrillo negro de
monte" is reported.

CHAMISSOA HBK.

Herbs or shrubs, erect or scandent, pubescent or glabrous; leaves alternate,
petiolate, rather broad; flowers perfect or sometimes monoecious, abortive stamens
present in the pistillate flowers and an abortive ovary in the staminate flowers,
each flower subtended by usually 3 bracts; inflorescence of few or many, axillary
or terminal, simple or paniculate, dense or lax spikes; sepals 5; stamens 5, connate
at the base into a cup; anthers ovoid, 4-celled; the filaments subulate; staminodia
none; ovary 1-ovulate, the style short or elongate, the stigmas 2, short or elongate;
utricle thin, dehiscent at or below the middle, circumscissile, surrounded by the
persistent calyx; seed vertical, reniform-lenticular, surrounded by a well-developed
aril, or the aril minute; embryo annular.

About 5 species, in tropical America. One other species (C.
Maximiliani Mart.) is known from Costa Rica and Panama.

160 FIELDIANA: BOTANY, VOLUME 24

Fruit truncate and emarginate at the apex, or conspicuously areolate.C. altissima.
Fruit rounded at the apex, not at all areolate C. macrocarpa.

Chamissoa altissima (Jacq.) HBK. Nov. Gen. & Sp. 2: 197.
pi 125. 1817. Celosia paniculata L. Sp. PI. ed. 2. 298. 1762, not
L. 1753. Achyranthes altissima Jacq. Enum. PI. Carib. 17. 1760.
Bejuco de sajan (fide Aguilar).

Common in the lowlands, sometimes ascending as high as 1,200
meters but mostly at lower elevations, usually in thickets,
especially in second growth; Pete*n; Alta Verapaz; doubtless in
Izabal; Jutiapa; Escuintla; Guatemala; Sacatepe'quez (Las Lajas);
Suchitepe'quez; Retalhuleu; San Marcos. Mexico to Panama, south
to Peru and Brazil; West Indies.

A suberect or arching shrub, or often high-scandent, branched, the stems
smooth or sulcate, glabrous or sparsely pilose; leaves on slender petioles 1-4 cm.
long, ovate to lanceolate, 6-18 cm. long, 2-8.5 cm. wide, abruptly acuminate or
acute, acute to truncate at the base, glabrous, or sparsely pilose beneath; flowers
in large, terminal or axillary, naked or leafy panicles composed of numerous stout
or slender, densely or laxly flowered spikes 2-20 cm. long and 0.6-2 cm. thick, the
sterile spikes more slender than the fertile ones, the rachises of the spikes usually
pubescent; flowers green or greenish white; bracts thin, about half as long as the
sepals, ovate or broadly ovate, mucronate; sepals 3-4 mm. long, oval to oblong
or ovate, acute or acuminate, sometimes mucronate, firm in age, carinate, promi-
nently and coarsely nerved; style shorter than the elongate stigmas; utricle globose
or oblong-ovoid, equaling or slightly exceeding the sepals, marginate and usually
depressed at the apex, circumscissile at or below the middle; aril bivalvate, enclos-
ing the seed; seed flat, 2-2.5 mm. in diameter, black and lustrous, punctulate.

One of the commonest plants of the wide thickets of the Pacific
plains.

Chamissoa macrocarpa HBK. Nov. Gen. & Sp. 2: 197. 1817.

Known in North America only from Lundell 4223, from Jones
Bank, Belize River, British Honduras; to be expected in Pete"n or
Izabal; reported from Mexico, but the report not confirmed by
recent collections; Colombia to Brazil and Peru.

In all respects similar to C. altissima, but easily distinguishable
by the fruit, as indicated in the key to the species. C. Maximiliani
Mart, of Costa Rica and Panama differs from both the Guatemalan
species in having a minute aril and an elongate style, longer than the
stigmatic branches.

CYATHULA Loureiro

Annual or perennial herbs, pubescent, branched; leaves opposite, petiolate,
entire; flowers fasciculate, each fascicle consisting of 1-2 perfect flowers and few

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 161

or many sterile ones, the fascicles bracteate and bracteolate, spicate or capitate,
reflexed in age; bracts concave, scarious, usually aristate; segments of the sterile
flowers finally produced into elongate bristles, these uncinate at the apex; perianth
of the perfect flower scarious, 5-parted, the subequal segments 1-nerved; stamens
5, the filaments united at the base; staminodia linear or quadrate and lacerate;
anthers oblong, 4-celled; ovary obovoid, the style filiform, the stigma capitate;
ovule 1, suspended on an elongate funicle; utricle included in the perianth, areolate
at the apex, membranaceous, indehiscent; seed inverted, oblong, the embryo
annular.

About 10 species in Asia, Africa, and tropical America. Only
the following are known from North America.

Sepals 3-4 mm. long; sterile segments 3-6, in age twice as long as the perianth.

C. achyranthoides.
Sepals 2 mm. long; sterile segments 12-20, equaling the perianth. . . .C. prostrata.

Cyathula achyranthoides (HBK.) Moq. in DC. Prodr. 13,
pt. 2: 326. 1849. Desmochaeta achyranthoides HBK. Nov. Gen. &
Sp. 2: 210. 1818. Cola de armado.

Found in thickets or waste ground, sometimes in mixed forest,
ranging from sea level to about 1,200 meters but chiefly at low
elevations; Alta Verapaz; Izabal; Santa Rosa; Escuintla; Guate-
mala; Sacatepe"quez; Retalhuleu; Huehuetenango; probably in
Pete"n. British Honduras; southern Mexico to Panama, southward
to Chile and Brazil; Greater Antilles.

Annual or perennial, the stems a meter long or less, erect to decumbent, often
rooting at the lower nodes, the stems strigose or glabrate; petioles 2-10 mm. long;
leaf blades oval to rhombic-elliptic, 5-15 cm. long, 2-6 cm. wide, acuminate,
cuneate at the base, thin, bright green, strigose or glabrate; spikes terminal and
axillary, 4-20 cm. long, 6-7 mm. thick, obtuse, the rachis short-villous, the flower
glomerules short-stipitate, 3-4 mm. long, each with 2 perfect flowers; bracts ovate-
lanceolate, long-attenuate, glabrous; bractlets ovate-oblong, long-acuminate or
attenuate, shorter than the flowers; sepals lance-oblong, acutish, 3-nerved, villous;
segments of the sterile flowers 3-6, twice as long as the perianth, at least in age;
seed 2 mm. long, oblong-ovate, dark brownish, lustrous.

Called "mozote" in Honduras. An annoying weed in many
localities, the hooked bristles of the sterile flowers clinging to cloth-
ing and also penetrating the skin painfully. This species is particu-
larly common on the low plains of Escuintla.

Cyathula prostrata (L.) Blume, Bijdr. Ned. Ind. 549. 1826.
Achyranthes prostrata L. Sp. PI. ed. 2. 296. 1762.

Izabal, near Virginia, at 500 meters or less. Native of tropical
Asia and Africa, naturalized from southern Mexico to Panama, and
in many regions of West Indies and South America.

162 FIELDIANA: BOTANY, VOLUME 24

Plants perennial, branched, the slender stems prostrate to suberect, a meter
long or less, rooting at the lower nodes, sparsely hirtellous or puberulent; petioles
2-10 mm. long; leaf blades rhombic-obovate to oval or broadly elliptic, 2-7 cm.
long, 1-4.5 cm. wide, acute, at the base rounded to acute, pilose-strigose or almost
glabrous, the lowest leaves sometimes broadly oval or suborbicular and rounded
at the apex; spikes terminal and axillary, 5-30 cm. long, 5-7 mm. thick, obtuse or
acutish, much interrupted below, the rachis puberulent; bracts oblong-ovate, long-
acuminate, glabrous, half as long as the flowers; bractlets triangular-ovate, half as
long as the perianth, aristate-mucronate, villous; fertile flowers 2, the perianth
segments 2 mm. long, lance-oblong, acuminate, prominently nerved, villous; seg-
ments of the sterile flowers 12-20, yellowish, about as long as the fertile flowers;
seed oblong, 1.2 mm. long, brownish or fuscous, lustrous.

This plant seems to be of limited and local occurrence in Central
America, along the Atlantic coast.

FROELICHIA Moench

Annual or perennial herbs, erect or procumbent, pubescent, with simple or
branched stems; leaves opposite, sessile or petiolate, entire; flowers perfect, sessile,
spicate, bracteate and bibracteolate, the spikes sessile or pedunculate; perianth
5-lobate, the lobes glabrous, the tube lanate, indurate in age and usually with
longitudinal wings, crests, or rows of spines; stamens 5, the filaments united to
form an elongate tube, this 5-lobate at the apex, the lobes short or elongate, obtuse;
anthers 2-celled, sessile in the sinuses between the lobes; ovary ovoid, the style
elongate, with a capitate stigma, or the stigma penicillate and sessile; ovule 1;
utricle ovoid, membranaceous, indehiscent, included in the calyx tube; seed
inverted, lenticular or obovoid, smooth, the embryo annular.

About 10 species, in temperate and tropical America. Seven
are reported from North America but only one has been collected
in Central America.

Froelichia interrupta (L.) Moq. in DC. Prodr. 13, pt. 2: 421.
1849. Gomphrena interrupta L. Sp. PI. 224. 1753.

Dry rocky hills and plains, 200-300 meters; Zacapa. Western
Texas and Mexico; Greater Antilles; Colombia to Paraguay and
Chile.

Perennial from a thick woody root, erect or procumbent, often branched from
the base, the stems slender, mostly 25-50 cm. long, white-tomentose or sericeous,
slightly viscid above; leaves petiolate, the petioles of the lower leaves sometimes
as long as the blades; leaf blades oval to ovate-orbicular, sometimes oblong to
narrowly lanceolate, 2.5-10 cm. long, 1-3.5 cm. wide, obtuse or acutish or some-
times acute, rounded to acute at the base, scaberulous or short-pilose on the upper
surface, beneath sericeous or floccose-tomentose; inflorescence interrupted, the
bracts acute or acuminate, stramineous or brown; calyx lobes lance-oblong, obtuse;
calyx tube deltoid in outline, nearly as broad as long, broadly winged laterally, the
thin wings entire or crenulate, the sides of the tube not appendaged; seed brown,
1.5 mm. long.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 163

GOMPHRENA L.

Annual or perennial herbs, erect or prostrate, branched, the stems often with
thickened nodes; leaves opposite, sessile or petiolate, entire; flowers perfect,
bracteate and bibracteolate, spicate or subcapitate, the heads solitary or glomerate,
terminal or axillary, naked or subtended by leaves, the flowers white, yellow, or
red; bractlets concave, carinate, often winged or cristate dorsally; perianth sessile,
terete or compressed, 5-lobate or 5-parted, usually lanate at the base; stamens 5,
the filaments united to form a tube, this included in the perianth or exserted, 5-
lobate at the apex, the lobes bifid or emarginate, the anthers sessile or stipitate
in the sinus of the lobe, oblong or linear, 1-celled; ovary turbinate or subglobose,
the style short or elongate; stigmas normally 2, subulate or filiform, or the stigma
bilobate; ovule 1; utricle compressed; seed inverted, sublenticular, smooth, the
embryo annular.

Some 90 species in both hemispheres, chiefly in tropical regions.
About 15 species are listed for North America, but no others are
known from Central America.

Bractlets not cristate; flower heads naked at the base G. Tuerckheimii.

Bractlets cristate along the keel, at least near the apex; heads subtended at the
base by leaves.

Flower heads 2-2.5 cm. broad G. globosa.

Flower heads 1-1.5 cm. broad.

Crests conspicuously widest at or near the apex of the bractlets, the flowers
thus appearing obtuse or only acutish ; bractlets equaling or shorter than

the flowers G. dispersa.

Crests widest below the apex of the bractlets, if perceptibly widest anywhere,

the flowers thus acuminate; bractlets much longer than the flowers.
Heads mostly solitary but sometimes glomerate, about 1 cm. wide, white

or pink; bractlets narrowly cristate G. decumbens.

Heads mostly glomerate, about 1.5 cm. wide, bright white; bractlets
broadly cristate G. nana.

Gomphrena decumbens Jacq. Hort. Schoenbr. 4: 41. pi. 482.
1804. G. perennis subsp. pseudodecumbens Stuchlik, Repert. Sp.
Nov. 11: 153. 1912 (based in part upon Guatemalan material).
G. decumbens var. carinata Suessenguth, Repert. Sp. Nov. 39: 8.
1935 (type from Chupadero, Santa Rosa, Heyde & Lux 4064).
Botoncillo; Sangrinaria (corruption of Sanguinaria)', Sanguinaria;
Siempreviva de monte.

A frequent weed found in waste or cultivated ground, in
gravelly or sandy stream beds, or in fields or thickets, chiefly at
low elevations but ascending to about 1,700 meters; probably in
Pete'n; Izabal; Zacapa; Chiquimula; Jutiapa; Escuintla; Sacate-
pe"quez; Quiche"; Huehuetenango. British Honduras; northward
through much of Mexico; South America.

164 FIELDIANA: BOTANY, VOLUME 24

Plants annual, usually 20-50 cm. high, erect or decumbent, generally branched,
the slender branches pilose-strigose; leaves short-petiolate, obovate, oblong, or
oval, 1.5-6 cm. long, 5-25 mm. wide, obtuse or rounded at the apex, acuminate or
attenuate at the base, appressed-pilose; flower heads subglobose or oblong, mostly
solitary and subtended by 2 leaves; bracts ovate-triangular, acuminate, white;
bractlets twice as long as the bracts, yellowish white or sometimes tinged with red
or pink, acuminate, cristate from below the apex nearly to the base, the crests
laciniate-dentate or almost entire; perianth much shorter than the bractlets,
copiously lanate, the lobes linear, long-attenuate; stamen tube usually included;
style elongate; seed 1.5 mm. long, brown.

Var. carinata is a form in which the crest of the bractlets is much
reduced and very narrow.

Gomphrena dispersa Standl. Contr. U. S. Nat. Herb. 18: 91.
1916. G. decumbens var. grandifolia Stuchlik, Repert. Sp. Nov. 11:
157. 1912, in part (based in part on Guatemalan material). Boton-
cillo; Siempreviva; Sanguinaria.

Waste or cultivated ground, on gravel or sand bars, or in thickets,
ascending to about 1,500 meters; Izabal; Zacapa; Jalapa; Santa Rosa;
Escuintla; Guatemala; Quiche"; Suchitepe'quez; Retalhuleu; San
Marcos. Central Mexico to Panama; West Indies.

Plants annual or perennial, erect to procumbent or prostrate, sometimes
forming dense mats, the stems a meter long or less, sparsely or densely appressed-
pilose; leaves short-petiolate, oval-obovate to oblong, 1.5-5 cm. long, 0.5-2 cm.
wide, obtuse or rounded at the apex, mucronate, acuminate to attenuate at the
base, sericeous-pilose, often glabrate on the upper surface; heads usually solitary,
terminal and axillary, subglobose or short-cylindric, 9-13 mm. broad, each sub-
tended by 2 acute sessile leaves; bracts rounded-ovate, acuminate, white; bractlets
5-6 mm. long, about 3 times as long as the bracts, acute to obtuse, white or rarely
purplish red, narrowly cristate at the apex, the crest extending along the keel for
only a short distance, denticulate or laciniate; perianth usually equaling the bract-
lets, densely lanate, the lobes oblong-linear, acuminate, white; stamen tube
usually included; style elongate, with slender stigmas; seed 1.5 mm. long, reddish
brown, lustrous. .

Maya names used in Yucatan are "chacmol" and "tmuul."
Names used in adjoining regions are "sanguinaria," "sangrinaria,"
and "secicante" in Honduras; and "amor seco" and "siempreviva"
in Yucatan. This species has been reported from Guatemala as
G. decumbens Jacq. It is very closely related to that and separated
by none too convincing characters, but the characters, such as they
are, hold and it is merely a matter of deciding whether they are of
specific importance or not. This plant is a common weed about
settlements all along the Atlantic coast of Central America and often
is especially plentiful on railway embankments.

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 165

Gomphrena globosa L. Sp. PI. 224. 1753. Amor seco; Inmortal;
Siempreviva; Boton.

Cultivated in Guatemala for ornament and rarely escaping to
waste or cultivated ground. Originally described from India and
reported to be a native of southern Asia, but the plant probably is
of American origin, derived through cultivation from such a species
as G. decumbens.

Plants annual, a meter high or less, often much branched, the stems swollen
at the nodes, pilose-strigose or rarely spreading-pilose; leaves short-petiolate,
oblong to oval, broadly ovate, or spatulate, 2-10 cm. long, acute and mucronate,
at the base rounded to acuminate, appressed-pilose; heads subtended each by
usually 2 leaves, long-pedunculate, globose or short-cylindric, white, yellow, red,
or purple, mostly 2-2.5 cm. broad; bracts triangular-ovate, long-acuminate;
bractlets 8-12 mm. long, 2-3 times as long as the bracts, broadly cristate along the
keel, the crest serrulate; perianth densely lanate; stamen tube longer or shorter
than the perianth; style elongate, slender, the stigmas linear.

The globe amaranth or bachelor's button so frequent in United
States gardens is one of the commonest garden flowers of Guatemala
and all Central America, where it is found in almost every garden of
rich or poor, and is in flower throughout the year. It is one of the
flowers most frequently offered in the markets, where it is in great
demand for house decoration, especially for altars, and above all
for making coronas or funeral wreaths, a purpose for which it is
utilized in many parts of the earth.

Gomphrena nana (Stuchlik) Standl. N. Amer. Fl. 21: 150.
1917. G. decumbens var. nana Stuchlik, Repert. Sp. Nov. 11: 158.
1912. G. Palmeri Standl. N. Amer. Fl. 21: 149. 1917.

Dry thickets or dry rocky slopes of the Oriente, 200-700 meters;
Zacapa; Chiquimula. Mexico.

Plants annual, usually prostrate and often forming mats, the stems appressed-
villous; basal leaves long-petiolate, the cauline short-petiolate, broadly oval to
oblong or oblanceolate, 2-5.5 cm. long, 1-1.3 cm. wide, rounded or obtuse at the
apex and mucronate, acute to attenuate at the base, sericeous beneath, usually
short-pilose above or glabrate; heads short-cylindric, 12-15 mm. broad, solitary
or glomerate, mostly terminal, each head or cluster of heads subtended by 4 or
more leaf -like bracts; floral bracts broadly ovate-triangular, acuminate, white;
bractlets 3 times as long as the bracts, long-acuminate, white, broadly cristate
above, narrowly cristate to the base, the crests denticulate above; flowers very
strongly compressed, the perianth much shorter than the bractlets, densely lanate,
the lobes linear, acute; stamen tube usually exserted; style elongate, with slender
stigmas; seed 1.5 mm. long, brown, lustrous.

166 FIELDIANA: BOTANY, VOLUME 24

Gomphrena Tuerckheimii (Vatke) Uline & Bray, Bot. Gaz.
20: 161. 1895. Telanthera Tuerckheimii Vatke ex Uline & Bray, loc.
cit. as synonym. Botoncillo; Maki (Huehuetenango).

Wet thickets or sometimes a weed in waste ground, 600-2,100
meters; type from Coban, Alta Verapaz, Tuerckheim 416; Alta
Verapaz; Jalapa; Jutiapa; Santa Rosa; Quiche"; Huehuetenango.
Honduras.

Plants perennial, erect or ascending, simple or branched, a meter high or less,
the stems densely pilose when young; petioles slender, 1-2 cm. long; leaf blades
oblong-ovate, 6-11 cm. long, 2.5-5.5 cm. wide, acute or acuminate, acute at the
base, thin, bright green and sparsely appressed-pilose, densely pilose-sericeous
beneath; peduncles axillary, very slender, naked, 5-8 cm. long; heads globose,
1 cm. in diameter; bracts and bractlets less than half as long as the sepals, ovate-
triangular, aristate-acuminate, stramineous when dry, white or whitish when
fresh, pilose or glabrate; sepals 3.5-4 mm. long, narrowly elliptic-oblong, acute,
3-nerved, densely long-pilose at the base; style very short, the stigmas subulate
and elongate.

In appearance this is very unlike other Central American mem-
bers of the genus and looks more like a species of Alternanthera.

IRESINE P. Browne

Shrubs or small trees or erect to decumbent or scandent herbs, pubescent or
glabrous; leaves opposite, petiolate, entire; flowers perfect, polygamous, or dioe-
cious, bracteate and bibracteolate, capitate or spicate, the spikes usually numerous
and paniculate; perianth terete, sessile, the 5 segments distinct, commonly lanate
or pilose; stamens united at the base into a short tube, the 5 filaments subulate,
entire, the pseudostaminodia usually short or wanting; anthers oblong, 2-celled;
ovary compressed, the style short or none, the stigmas 2-3, subulate or filiform
or in the staminate flowers sometimes capitate; ovule 1; utricle compressed, mem-
branaceous, indehiscent; seed inverted, smooth, the embryo annular.

Probably 45 species, chiefly in tropical America, a few in tropi-
cal Africa. From North America about 30 species are known.

Leaves variegated with red and yellow, retuse at the apex; cultivated plants.

/. Herbstii.
Leaves green, not retuse; native plants.

Flowers perfect or polygamous (staminate and pistillate upon the same plant).

Spikelets not more than 2 mm. in diameter; trees /. arbuscula.

Spikelets 3-5 mm. in diameter; shrubs or vines.

Bracts and bractlets rounded or obtuse at the apex; an erect shrub or a
woody vine /. nigra.

Bracts and bractlets acute or acuminate, cuspidate; erect or scandent herb.

/. angustifolia.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 167

Flowers dioecious.

Plants woody throughout, erect or scandent shrubs.

Branches of the inflorescence glabrous or nearly so /. inlerrupta.

Branches of the inflorescence copiously, often densely, pubescent.

Panicles very dense; bracts and sepals villous only at the base; staminate

sepals 2.5-3 mm. long /. grandis.

Panicles lax and open; bracts and sepals copiously villous; staminate

sepals 2 mm. long or less /. Calea.

Plants herbaceous.

Pubescence of the inflorescence and lower leaf surface consisting in part of
lustrous amber-colored hairs; bracts usually dentate near the base.

/. spiculigera.

Pubescence of whitish, neither lustrous nor amber-colored hairs, or wanting.

/. Celosia.

Iresine angustifolia Euphrase"n, Beskr. St. Barthel. 165. 1795.
I. elatior Rich, ex Willd. Sp. PL 4: 766. 1805.

Dry or damp thickets of the lowlands, abundant in places along
the Pacific coast; El Progreso (Morazan); Santa Rosa; Escuintla;
Retalhuleu. Mexico to Panama, southward to Ecuador and Brazil;
West Indies.

Plants usually erect and as much as 1.5 meters high, sometimes subscandent,
often much branched, often blackening when dried, the slender stems green, very
sparsely villous when young; leaves on slender petioles 5-25 mm. long, lance-ovate
to linear-lanceolate, 5-10 cm. long, 1-4 cm. wide, acuminate or long-attenuate,
at the base acute to long-acuminate, glabrous, or very sparsely villous beneath
along the veins; flowers perfect, loosely paniculate, the spikelets short or elongate,
usually pedunculate, the rachis lanate; bracts broadly ovate, acute, the bractlets
ovate, cuspidate-acuminate, twice as long as the bracts and equaling the calyx,
hyaline, when dry brown or brownish, glabrous or villous; sepals elliptic-oblong,
acute or acutish, 1.5 mm. long, 1-nerved, densely villous.

Iresine arbuscula Uline & Bray, Bot. Gaz. 21: 350. 1896.
Durazno de montana (Quezaltenango).

Moist or wet forest of the central and western mountains, 150-
2,000 meters; type from Volcan de Tecuamburro, Santa Rosa,
Heyde & Lux 4570; Pete"n; Santa Rosa; Escuintla; Sacatepe"quez
(near Las Lajas); Suchitepe"quez (near Patulul); Quezaltenango
(between Santa Maria de Jesus and Calahuache"). Chiapas and
Tabasco.

A shrub or tree 4.3-12 meters high, glabrous except in the inflorescence, with
slender terete branches; leaves on slender petioles 2-4 cm. long, elliptic to elliptic-
oblong, 14-20 cm. long, 4-7 cm. wide, acute or acuminate at each end, blackish
or bright green when dried; flowers polygamo-dioecious, arranged in a very large,
lax panicle, white, the branches of the panicle puberulent or glabrate; spikelets
mostly sessile, the rachis lanate; bracts and bractlets ovate-orbicular, scarious,

168 FIELDIANA: BOTANY, VOLUME 24

rounded at the apex, glabrous; sepals of the staminate flowers oblong-oval, 1.5
mm. long, obtuse, not nerved, very sparsely lanate at the base.

This is probably the only North American species that becomes
a true tree, and it has a distinct and often rather thick trunk. The
living leaves are handsome because of their fresh green coloring,
and the whole tree with its large panicles of flowers is a rather
ornamental one. It has been found in some abundance in the
barrancos of the Volcan de Fuego. The name "cenicero" has been
reported for this species from the North Coast region, but we have
not seen the material so determined.

Iresine Calea (Ibafiez) Standl. Contr. U. S. Nat. Herb. 18: 94.
1916. Gomphrena latifolia Mart. & Gal. Bull. Acad. Brux. 10, pt. 1:
349. 1843. Achyranthes Calea Ibafiez, Naturaleza 4: 79. 1879.
/. latifolia Benth. & Hook. Gen. PI. 3: 42. 1880, not D. Dietr. 1839.
Hebanthe mollis Hemsl. Biol. Centr. Amer. Bot. 3: 20. 1882. Hierba
de burro (Guatemala) ; Pata de gallina (Guatemala) ; Bejuco gitano
( Jutiapa) ; Flor de Maria (Jutiapa).

Moist or dry thickets, sometimes in dry open forest, often in
roadside hedges, from sea level to about 1,800 meters, most plentiful
at middle elevations; Baja Verapaz; Zacapa; Chiquimula; Jalapa;
Jutiapa; Santa Rosa; Escuintla; Guatemala; Sacatepe*quez ; Chi-
maltenango; Solola; Huehuetenango; Retalhuleu; Quezaltenango;
San Marcos. Mexico to Costa Rica; not known from British
Honduras,

An erect or scandent shrub, often climbing rather high over large shrubs,
much branched, the branches densely covered with pale appressed hairs, at least
when young; leaves on petioles 1.5 cm. long or less, broadly ovate to ovate-oblong,
5-10 cm. long, 1.5-7 cm. wide, acuminate or rarely obtuse, at the base rounded
or obtuse, thinly scaberulous or glabrate above, densely or sparsely pilose-sericeous
beneath; flowers dioecious, in broad open panicles, white or whitish, the panicles
leafy below, the spikelets dense, short or elongate, sessile or pedunculate; bracts
and bractlets of the staminate flowers one-third as long as the sepals, broadly
ovate, rounded to acutish at the apex, more or less villous; sepals narrowly oblong,
obtuse, 2 mm. long, pilose; staminodia very short, broad, dissected at the apex
into short filiform segments or rarely subentire; bracts and bractlets of the pistil-
late flower nearly as long as the sepals, these lanceolate, attenuate, 1.5 mm. long,
densely pilose with white or brownish hairs, very faintly nerved.

In Salvador the shrub is given a large number of names, among
them "siete pellejos," "flor de corona," "algodoncillo," "flor de
Jesus," "cola de chivo," "cola de cabra," "cometernero," "siete
cascaras," "coyontura," and "tacuquelite," the last being Nahuatl.
The leaves and branches are much eaten by stock during the dry

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 169

season. In the Pacific lowlands of Guatemala the plant begins to
bloom about the first of January, and is without inflorescences during
the early part of the verano. The inflorescences retain their form
and coloring when dried and on this account are much used for the
decoration of houses and churches, especially upon altars. Indian
cargadores on their return from the coast to the Guatemalan high-
lands often carry large bunches of the plant, which perhaps is
employed also in domestic medicine. The specific name commemo-
rates a friend of the original describer of the species, and should be
written with a capital letter. Specimens of /. Calea have been
reported from Guatemala under the name I. canescens Humb. &
Bonpl.

Iresine Celosia L. Syst. ed. 10. 1291. 1759. Celosia paniculata
L. Sp. PI. 206. 1753. /. celosioides L. Sp. PI. ed. 2. 1763. Pie de
paloma (Quezaltenango, San Marcos); Velo de princesa (Guate-
mala) ; Adorno de nino (Jutiapa) ; Chancanil (Alta Verapaz, Quecch') ;
Tabudo (Santa Rosa); Mosquito (Jalapa).

One of the commonest plants of Guatemala, generally distributed
except at high elevations, often a weed in cultivated ground or waste
places, in thickets, or often in dense wet mixed forest, ascending to
about 2,800 meters or perhaps even higher; Peteri; Izabal; Alta
Verapaz; El Progreso; Zacapa; Chiquimula; Jalapa; Jutiapa; Santa
Rosa; Escuintla; Guatemala; Sacatepdquez; Chimaltenango; Solola;
Quich4 ; Huehuetenango; Suchitepe*quez ; Retalhuleu; Quezalte-
nango; San Marcos. Southeastern United States to Mexico and
Panama, southward through most of South America; West Indies.

Plants annual, but sometimes persisting more than a single year, the stems
usually branched, erect to procumbent or sometimes elongate and clambering,
glabrous or sparsely villous, especially at the nodes; leaves thin, slender-petiolate,
broadly ovate to lanceolate, 5-14 cm. long, 2-7 cm. wide, acute or acuminate,
rounded to broadly cuneate at the base, glabrous or somewhat villous; panicles
usually large and much branched, the branches more or less villous, the spikelets
sessile or pedunculate, usually dense, 5-25 mm. long; flowers white or pink, the
pistillate with copious long wool at the base; bracts ovate or ovate-orbicular,
obtuse or acute; sepals 1-1.5 mm. long, obtuse or rounded at the apex, those of the
pistillate flowers conspicuously 3-nerved; seeds 0.5 mm. in diameter, obovoid or
orbicular, dark red, lustrous.

The Maya names of Yucatan are recorded as "zactezxiu" and
"zacxiu." In Honduras the plant sometimes is called "hierba de
gato"; in Salvador, "siete pellejos," "coyontura," "coyontura de
polio," and "taba de giiegiiecho." It is one of the commonest weeds
of Central America and abounds in many parts of Guatemala, partic-

170 FIELDIANA: BOTANY, VOLUME 24

ularly on the Pacific plains. In spite of its great abundance, the
plant has little if any practical importance; little attention is paid
to it, and there is no constant or well-fixed vernacular name for it.
About Coban the sap is applied to the skin as a remedy for erysipe-
las. Guatemalan material referred here exhibits little variation.
Most remarkable is a form sometimes found in forest in which the
flowers are purplish pink rather than white or greenish. This per-
haps deserves rank as a form by those who are interested in such
trivial things. Some Guatemalan material has been referred to
Iresine frutescens Moq., a name probably better reduced to the
synonymy of /. Celosia.

Iresine grandis Standl. N. Amer. Fl. 21: 163. 1917.

Damp thickets or in oak forest, central and western departments,
1,600-2,400 meters; Sacatepe'quez; Chimaltenango; Quezaltenango.
Southern Mexico.

An erect shrub as much as 4.5 meters high or a large woody vine, the branches
at first densely villous-tomentulose, sometimes glabrate in age; leaves large, on
petioles 1-2 cm. long, ovate-rhombic or ovate, 6-15 cm. long, 3-7 cm. wide, acute
or acuminate, at the base acute or obtuse and short-decurrent, glabrate above,
densely villous-tomentose beneath, sometimes glabrate in age; flowers dioecious,
in large dense panicles 15-30 cm. long and often as broad; spikelets sessile or
pedunculate; bracts and bractlets of the staminate flowers one-third as long as the
sepals, ovate-orbicular, glabrous, the sepals oblong, obtuse or acutish, 2.5-3 mm.
long, glabrous; staminodia rhombic or lanceolate, finely dissected; bracts of the
pistillate flowers as long as the sepals, round-ovate, the sepals oblong, obtuse, 1.5
mm. long, densely pilose; seed 1 mm. long, reddish brown.

This species has been reported from Guatemala under the name
Iresine canescens Humb. & Bonpl. In Guatemala it seems to be
most plentiful on the mountains about Antigua.

Iresine Herbstii Hook. Gard. Chron. 1864: 654. 1864. Achy-
ranthes Verschaffeltii Lem. 111. Hort. 11: pi. 409. 1864. /. Verschaf-
feltii Lem. 111. Hort. pi. 418. 1864.

Cultivated commonly for ornament in gardens of most parts of
Guatemala except perhaps in the highlands. Cultivated throughout
tropical America and in many other regions of the earth; in the north
often seen in hothouses or as a summer bedding plant; probably of
American origin but unknown in a wild state.

An erect or ascending annual, rather stout and succulent, usually branched,
sparsely short- villosulous, especially about the nodes; leaves slender-petiolate,
suborbicular or ovate-orbicular, 2.5-6.5 cm. long and of equal or greater breadth,
rounded to truncate at the base and usually short-decurrent, deeply retuse at the

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 171

apex or sometimes merely rounded, thick and succulent, purplish red, or green
and striped with yellow or pink along the veins, glabrous above or sparsely sca-
berulous, beneath rather sparsely furnished with short, appressed, often lustrous
and yellowish hairs; flowers dioecious, the panicles 10-20 cm. long, the branches
villous with usually lustrous hairs, the spikelets slender and loosely flowered,
sessile or short-pedicellate, the flowers white or stramineous; bracts and bractlets
ovate-orbicular, obtuse, glabrous, half as long as the sepals; sepals 1 mm. long,
ovate to oblong, obtuse or acutish, those of the pistillate flowers 3-nerved.

Called "mano de lagarto" in Honduras, "chorcha de gallo" in
Salvador. Probably this plant has been grown in American gardens
for centuries, although it may not have been long in Mexico and
Central America. Unknown in the wild state, it may well have as
its not too remote ancestor Iresine spiculigera Seub., to which it is
nearly allied in flower characters and pubescence.

Iresine interrupta Benth. Bot. Voy. Sulph. 156. 1844. Pie de
zanate (fide Aguilar).

In forest or thickets, 1,100-1,650 meters; Alta Verapaz; Santa
Rosa; Sacatepe"quez(?); Quiche". Mexico.

A shrub, usually clambering or scandent, the terete branches striate, glabrous,
at least in age, pale green; leaves on stout petioles 5 cm. long or less, ovate-rhombic
to ovate or lanceolate, 5-15 cm. long, 1-10 cm. wide, acute to attenuate, rounded
or obtuse at the base and short-decurrent, thick, glabrous, prominently nerved;
flowers dioecious, in broad or narrow, open, sparsely leafy panicles; spikelets short
or elongate, sessile or pedunculate; sepals of the staminate flowers 1.5-2 mm. long,
whitish, scarious, densely villous; staminodia denticulate at the apex or entire;
bracts of the pistillate flowers ovate-orbicular, nearly as long as the sepals, obtuse,
mucronulate, stramineous, glabrous; sepals oval, obtuse, 1.5 mm. long, 3-nerved,
villous; seed orbicular, 0.8 mm. long, black and lustrous.

Common in some parts of Mexico, but in Guatemala rare or
overlooked.

Iresine nigra Uline & Bray, Bot. Gaz. 21: 350. 1896. Canilla
(fide Aguilar).

Dry or wet or moist thickets and forest, ranging from sea level
to about 1,800 meters; Pete'n; Alta Verapaz; Zacapa; Chiquimula;
Santa Rosa; Escuintla; Guatemala; Sacatepe"quez; Chimaltenango;
Suchitepe"quez; Retalhuleu; Sa'n Marcos. Veracruz to British Hon-
duras, Honduras, and Salvador; type from San Pedro Sula,
Honduras.

A shrub or small tree, sometimes attaining a height or length of 9 meters,
usually smaller, often scandent, blackish when dried, the slender branches glabrous,
or sparsely puberulent in the inflorescence; leaves short-petiolate, ovate to oblong-

172 FIELDIANA: BOTANY, VOLUME 24

lanceolate, 5-15 cm. long, 2-4 cm. wide, acute to long-acuminate, acute or acumi-
nate at the base and short-decurrent, glabrous or nearly so; flowers polygamo-
dioecious or sometimes perfect, in mostly small and lax panicles; spikelets short,
mostly sessile; bracts and bractlets half as long as the sepals or shorter, rounded-
ovate, rounded at the apex, white or stramineous, glabrous; sepals ovate-oblong,
1.5 mm. long, obtuse, obscurely nerved, glabrous or sparsely pilose at the apex,
the basal hairs soft, whitish, equaling or exceeding the sepals; staminodia minute.

Iresine spiculigera Seub. in Mart. Fl. Bras. 5, pt. 1: 228. pi. 70.
1875. /. acicularis Standl. Contr. U. S. Nat. Herb. 18: 93. 1916.
Pie de paloma (Quezaltenango).

Damp or wet forest, 1,200-1,800 meters, or also at lower eleva-
tions; Alta Verapaz; Escuintla; Sacatepe"quez (type of /. acicularis
from Volcan de Fuego, 2,700 meters, Kellerman 4549); Suchitepe"-
quez; Quezaltenango. Costa Rica; south to Brazil and Argentina.

A slender herb 1-3 meters long, erect or reclining, the stems branched, very
sparsely pubescent with short slender hairs; leaves slender-petiolate, ovate or
broadly ovate, 7-20 cm. long, 4-10 cm. wide, acute to long-attenuate, rounded or
obtuse at the base and abruptly short-decurrent, thin, bright green, very sparsely
villous above with short soft pale hairs, similarly pubescent beneath and with
numerous appressed, lustrous, amber-colored or bright yellow, acicular hairs,
villous-ciliate; panicles large and dense, somewhat leafy, the rachises sparsely
villous and with acicular hairs like those of the leaves, these most abundant at
the base of the spikelets; spikelets sessile or pedunculate, densely flowered, 4-12
mm. long; bracts white, rounded-ovate or narrowly ovate, acute, equaling or
half as long as the sepals; sepals 1.5 mm. long, narrowly oblong, acute, those of the
pistillate flowers 3-nerved; seed 0.5 mm. broad, dark reddish brown, lustrous.

PFAFFIA Martius

Herbs or shrubs, pubescent or glabrate, branched, sometimes scandent;
leaves opposite, sessile or short-petiolate; flowers mostly perfect, bracteate and
bibracteolate, capitate or spicate, the spikes or heads pedunculate, often numerous
and paniculate; perianth sessile, terete, the 5 segments free, subequal, pilose or
lanate; filaments united to form a 5-lobate tube, the lobes fimbriate, dentate, or
3-lobate; staminodia none; anthers narrowly oblong, 2-celled; ovary ovoid, the
style very short or none, the stigma capitate or bilobate; ovule 1; utricle ovoid,
membranaceous, indehiscent; seed inverted, smooth, the embryo annular.

About 20 species, all but the following South American.

Pfaffia Hookeriana (Hemsl.) Greenm. Field Mus. Bot. 2: 330.
1912. Hebanthe Hookeriana Hemsl. Biol. Centr. Amer. Bot. 3: 19.
1882.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 173

Damp forest or thickets, chiefly at low or middle elevations but
ascending to about 1,800 meters; Alta Verapaz; Izabal; Escuintla;
Sacatepequez; Retalhuleu; San Marcos. Southern Mexico and
British Honduras to Panama.

A scandent shrub, often greatly elongate and climbing over trees, the branches
terete, the younger ones and those of the inflorescence densely pilose with short
ascending fulvous hairs; leaves short-petiolate, ovate-oblong to broadly ovate or
oval-oblong, 4-10 cm. long, 1.5-4 cm. wide, abruptly acute or long-acuminate,
obtuse or rounded at the base, thick and firm, drying blackish, pilose-strigose, or
sometimes glabrate on the upper surface; flowers spicate, the spikes 1.5-5 cm.
long, verticillately paniculate, the panicles short and narrow; bracts and bractlets
about one-fourth as long as the sepals, suborbicular, concave, short- villous; sepals
ovate-oblong, 2-2.5 mm. long, obtuse, the outer ones strigose, the inner ones
densely pilose, the soft white hairs twice as long as the sepals; filaments filiform;
style very short.

PHILOXERUS R. Brown

Perennial herbs, prostrate or procumbent, branched, somewhat fleshy, gla-
brous or pubescent, the stems terete or angulate; leaves opposite, narrow; flowers
perfect, bracteate and bibracteolate, imbricate in dense, white, sessile or peduncu-
late, short or elongate spikes; bracts chartaceous; perianth dorsally compressed,
thickened at the base and short-stipitate, 5-parted, the segments subequal, the
outer ones obtuse, the inner ones narrower and acute; stamens 5, the filaments
subulate, connate at the base; anthers oblong, 2-celled; utricle broadly ovoid,
compressed, coriaceous, indehiscent; seed inverted, lenticular, smooth, the embryo
annular.

Probably three or four species, chiefly on seashores, tropical
America and western Africa. Only the following species is known
in North America.

Philoxerus vermicularis (L.) R. Br. Prodr. 416. 1810. Gom-
phrena vermicularis L. Sp. PI. 224. 1753.

In saline flats near the sea beaches; Izabal; doubtless also along
the Pacific coast. Florida to Texas, Mexico, and Panama; West
Indies; Colombia to Brazil; west coast of Africa.

Plants much branched, glabrous outside the inflorescence except in the leaf
axils, there villous; branches stout and succulent, prostrate or procumbent and
rooting at the nodes, usually 30-75 cm. long; leaves sessile, linear to oblanceolate
or rarely oblong, 1.5-5 cm. long, 2-12 mm. wide, obtuse or acute, attenuate to the
base, thick and fleshy; spikes solitary or glomerate, sessile or short-pedunculate,
globose or usually cylindric in age, 1-3 cm. long, about 1 cm. wide, obtuse, the
rachis lanate, the flowers white; bracts broadly ovate, 1-nerved, acute or obtuse;
bractlets ovate-oblong, slightly shorter than the sepals, acute, glabrous; sepals
oblong, 3-5 mm. long, the outer ones glabrous, the inner usually lanate near the
base; seed orbicular, 1 mm. broad, dark brown, lustrous.

174 FIELDIANA: BOTANY, VOLUME 24

One of the characteristic plants of sea beaches or salt flats in
many parts of Central America, usually found just back of mangrove
thickets.

PLEUROPETALUM Hooker f.

Glabrous shrubs with branched stems; leaves alternate, petiolate; flowers
perfect, bracteate and bibracteolate, pedicellate, racemose or paniculate, green;
perianth segments 5, subcoriaceous, equal, obtuse, striate-nerved, spreading in
fruit; stamens 5-8, the filaments subulate-filiform, connate basally into a short
cup, the anthers 4-celled; ovary globose-ovoid, attenuate to a short style, the stig-
mas 2-4, short, subulate; ovules numerous, on capillary funicles; fruit baccate,
globose, rupturing irregularly; seeds numerous, reniform-orbicular, lenticular, erect,
the testa black and lustrous; embryo annular.

About five species are recognized, three others having been
described from Central America. P. calospermum Standl. of Salva-
dor is to be expected in the Oriente of Guatemala. It has longer
sepals than P. Sprucei.

Pleuropetalum Sprucei (Hook, f.) Standl. N. Amer. Fl. 21:
96. 1917. Melanocarpum Sprucei Hook. f. in Benth. & Hook. Gen.
PL 3: 24. 1880. P. costaricense Wendl. ex Hook. f. Bot. Mag. pi.
6674. 1883. P. tucurriquense Bonn. Smith, Bot. Gaz. 61 : 387. 1916.
Ichaj (fide Aguilar); Cinco negritos (Huehuetenango).

Moist or wet, mixed forest, ascending from sea level to about
1,800 meters; Izabal; Guatemala; Sacatepe"quez; Chimaltenango;
Huehuetenango; Quezaltenango; San Marcos. Jalisco to Veracruz,
southward to Panama, and extending to Peru.

A shrub 1-1.5 meters high; leaves on slender petioles 2-3 cm. long, often
blackish when dry, elliptic to lance-oblong, 10-18 cm. long, 4-6.5 cm. wide, rather
abruptly acuminate, at the base acute or abruptly acuminate; flowers in dense
terminal panicles 3.5-6 cm. long, the bracts at the bases of the pedicels ovate,
1.5-2 mm. long; bractlets ovate, 1 mm. long; sepals ovate-oval, 2.5 mm. long,
glabrous; stamens 5-8; fruit red at maturity, sometimes dark purple or orange,
juicy, globose, 5 mm. in diameter; seeds 1.5 mm. in diameter.

NYCTAGINACEAE. Four-o'clock Family

Reference: Standley, N. Amer. Fl. 21: 171-254. 1918.

Annual or perennial herbs, shrubs, or trees, sometimes scandent, dichoto-
mously or trichotomously branched, glabrous or pubescent, the stems often swollen
at the nodes, sometimes armed with spines; leaves simple, opposite, alternate, or
verticillate, without stipules, sessile or petiolate, entire or essentially so, often
marked with conspicuous raphids; flowers perfect or unisexual, in the latter case
dioecious, regular, variously arranged, usually bracteate or variously involucrate,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 175

the involucre of free or connate segments, often calyx-like and enclosing 1 or more
flowers, persistent or deciduous, green or brightly colored; perianth inferior,
simple, herbaceous or usually corolla-like, small or large, tubular to campanulate
or funnelform, persistent in fruit and often accrescent, closely enclosing the peri-
carp; limb of the perianth truncate or with 3-5 teeth or lobes, the segments usually
induplicate-valvate in bud; stamens 1-many, hypogynous, the filaments usually
united at the base, unequal, filiform, included or exserted, the anthers dorsifixed
near the base, didymous, the cells dehiscent by lateral slits; ovary sessile or stipi-
tate, 1-celled, the style short or elongate, sometimes wanting, filiform, the stigma
simple and capitate, peltate, or fimbriate; fruit an anthocarp, composed of the
persistent, coriaceous, fleshy, or indurate base of the perianth tube enclosing the
indehiscent utricle and adherent to it, costate, sulcate, or winged, often viscous
when wet, frequently bearing viscous glands; seed erect, with hyaline testa, the
endosperm scant or abundant, the embryo straight or curved.

A chiefly tropical family, best represented in the warmer parts
of America. Besides the following genera, at least two others are
represented in Central America (Panama): Cephalotomandra and
Colignonia.

Leaves alternate.

Plants herbaceous, erect; flowers free from the small and inconspicuous green
bracts • Boldoa.

Plants woody vines; flowers inserted singly on the inner face of a large colored

bract Bougainvillea.

Leaves opposite.
Plants trees or shrubs.

Stamens included; fruit without stipitate glands; plants unarmed Neea.

Stamens exserted; fruit sometimes with stipitate glands; plants often armed
with spines.

Fruit juicy, without glands; plants unarmed Torrubia.

Fruit dry, bearing numerous stipitate glands; plants often armed with

spines Pisonia.

Plants herbaceous, sometimes slightly ligneous near the base, never trees or

large shrubs.
Fruit lenticular, with dentate winglike margins; flowers in clusters of 3,

subtended by a 3-parted involucre Allionia.

Fruit terete or angulate, never lenticular or with dentate margins; flowers

never as above.

Flowers surrounded by a calyx-like involucre of united bracts . . . Mirabilis.
Flowers not involucrate, the bracts distinct.

Fruit with 5 or fewer angles, obpyramidal or clavate, without stipitate

glands; perianth campanulate Boerhaavia.

Fruit 10-costate, terete, with numerous stipitate glands; perianth fun-
nelform Commicarpus.

ALLIONIA L.

Prostrate, annual or perennial herbs, pubescent; leaves opposite, those of a
pair very unequal, petiolate, the blades broad, entire or sinuate; flowers perfect,

176 FIELDIANA: BOTANY, VOLUME 24

in axillary pedunculate clusters of 3, each flower subtended by a broad green
concave bract, the bracts subequal, slightly united at the base, thin, enclosing
the fruit; perianth corolla-like, purple-red, short-funnelform, the tube constricted
above the ovary, the limb oblique, 4-5-1 obate, induplicate-plicate; stamens 4-7,
the filaments unequal, capillary, exserted, the anthers didymous; ovary ovoid, the
style capillary, the stigma capitate; fruit coriaceous, obovoid or oval, strongly
compressed, 3-costate or cristate on the inner surface, the outer surface bearing
2 parallel longitudinal rows of stipitate glands, the thin margins dentate or entire,
inflexed; embryo uncinate.

A small genus of 3 species in North and South America, or perhaps
of a single polymorphic species. A single species, at any rate, is
known from Central America.

Allionia incarnata L. Syst. ed. 10. 890. 1759. Wedelia incarnata
Kuntze, Rev. Gen. 533. 1891. Wedeliella incarnata Cockerell,
Torreya 9: 167. 1909.

Dry plains and hillsides of Zacapa, about 200 meters. South-
western United States; Mexico; Venezuela; Argentina and Chile.

Perennial from a slender or thick, vertical, sometimes ligneous root; stems
numerous, prostrate and often forming mats, a meter l&ig or less, viscid-villous
or glandular-puberulent; leaves on petioles 2 cm. long or less, mostly oval to ovate,
1-6 cm. long, 1-4.5 cm. wide, rounded to acute at the apex, subcordate or rounded
and unequal at the base, somewhat succulent, paler beneath, glandular-puberulent
or viscid-villous, sometimes glabrate in age; involucres on slender peduncles 5 cm.
long or usually much shorter, the segments obovate-orbicular, 5-8 mm. long,
rounded or obtuse at the apex; perianth 1-1.5 cm. long, purple-red or rarely white,
villous or puberulent outside; fruit 3-4.5 mm. long, pale brown or olive, the inner
side 3-costate, the margins usually with 3-5 teeth on each side, these strongly
incurved.

The plant is plentiful on the plains of Zacapa but probably
disappears during the latter part of the dry season. It affords a
rather interesting example of discontinuous distribution in North
America, for it is not found, so far as known, from the southern
border of Guatemala to Venezuela, where it reappears, in spite of
the fact that in the intervening areas there are habitats in which it
might well be expected. There is, of course, the possibility that it
was introduced by man into eastern Guatemala, since the fruits are
well adapted to human dispersal.

BOERHAAVIA L.

Annual or perennial herbs, branched, erect to prostrate, pubescent, the stems
often with viscous areas in the internodes; leaves opposite, petiolate, those of a
pair often unequal, entire or sinuate; flowers perfect, very small, umbellate,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 177

cymose, capitate, racemose, or solitary, bracteate, the bracts small and often
minute; perianth corolla-like, campanulate to almost rotate, constricted above the
ovary, the limb shallowly 5-lobate; stamens 1-5, exserted or included, the filaments
capillary, unequal, connate at the base, the anthers didymous; ovary stipitate, the
style filiform, the stigma peltate; fruit obovoid or obpyramidal, 3-5-angulate,
rarely winged, glabrous or pubescent, symmetric; embryo uncinate.

Forty species of wide distribution in tropical and warm regions,
most numerous in America. About 25 species are known from North
America, but only those listed here are reported from Central
America.

Fruit glabrous; plants annual, erect B. erecta.

Fruit viscid-pubescent; plants perennial, usually prostrate or procumbent.

B. diffuse.

Boerhaavia diffusa L. Sp. PL 3. 1753. B. caribaea Jacq. Obs.
Bot. 4: 5. 1771. B. coccinea Mill. Card. Diet. ed. 8. no. 4. 1768.
B. paniculata L. Rich. Act. Soc. Hist. Nat. Paris 1: 105. 1792.
B. hirsuta Willd. Phytogr. 1: L 1794. B. viscosa Lag. & Rodr. Anal.
Cienc. Nat. 4: 256. 1801. Hierba de cabro; Moradilla (fide Aguilar);
Erisipela (Pete*n).

Sandy fields or dry or moist thickets, often in cultivated ground,
common about dwellings, chiefly in the tierra caliente but ascending
to about 1,400 meters; Pete"n; doubtless in Izabal; Zacapa; Jutiapa;
Santa Rosa; Escuintla; Guatemala; Sacatepe'quez; Solola; Suchi-
tepe"quez; Quiche"; Huehuetenango; San Marcos; doubtless in all the
departments having land at 1,200 meters or less. Southern United
States to Mexico, British Honduras, and Panama; West Indies;
South America; Old World tropics.

Perennial from a thick woody root, the stems often much branched, a meter
long or less, decumbent to prostrate, viscid-puberulent almost throughout and
often hirsute; leaves on petioles 4 cm. long or shorter, suborbicular to oval, oblong,
or ovate, 2-5 cm. long, broadly rounded to acute at the apex, truncate or rounded
at the base, pale beneath, often brown-punctate, glabrous, puberulent, or villous;
flowers capitate, sessile or nearly so, the heads long-pedunculate, in terminal and
axillary cymes; bracts minute, lanceolate; perianth purple-red, 2 mm. broad,
puberulent or glandular-puberulent outside; stamens 1-3, short-exserted; fruit
narrowly obovoid, 2.5-3 mm. long, densely glandular-puberulent, 5-sulcate, the
angles and sulci smooth.

A common weed in most parts of the lowlands, especially plenti-
ful about dwellings. The fruits adhere tenaciously to feathers and
hair of animals, and thus are spread abundantly. The plant is said
to be used in Pete*n as a remedy for erysipelas, hence the name
"erisipela" given it there. Maya names reported from Yucatan are
"uxiuam" and "chacilxiu."

178 FIELDIANA: BOTANY, VOLUME 24

Boerhaavia erecta L. Sp. PL 3. 1753. Maravillita; Anisillo.

A common weed in fields and waste ground, often in streets,
ascending from sea level to about 1,200 meters; Pete"n; Zacapa;
Santa Rosa; Escuintla; Guatemala; Solola; Suchitepe"quez ; Retal-
huleu; doubtless in all the lowland departments. Southern United
States to Mexico, British Honduras, and Panama; West Indies;
South America.

An annual, usually much branched and erect, sometimes decumbent, the
branches reddish, finely puberulent below, the middle internodes often with brown
viscous bands; leaves on slender petioles 4 cm. long or less, ovate-rhombic to
deltoid-ovate, oval, or oblong, 2-6 cm. long, 1-4.5 cm. wide, broadly rounded or
obtuse to rarely acute at the apex, truncate or rounded at the base, bright green
above, paler or glaucous beneath, usually brown-punctate, glabrous or sparsely
puberulent; inflorescence cymose, much branched, the branches slender, mostly
glabrous, the flowers irregularly umbellate-cymose or subracemose at the ends of
long slender peduncles, the pedicels 1-5 mm. long; bracts minute; perianth white
or pinkish, 1-1.5 mm. long, glabrous, sometimes glandular-punctate; stamens 2-3,
exserted; fruit narrowly obpyramidal, 3-3.5 mm. long, 1-1.5 mm. broad at the
truncate apex, green, glabrous, 5-angled, the angles obtuse or subacute, smooth,
the sulci coarsely transverse-rugulose.

A common weed about dwellings or in cultivated ground in most
of the tierra caliente of Guatemala as well as in Central America
generally. Often this and B. diffusa grow together in abundance,
but on the Pacific plains of Guatemala, where Boerhaavia, erecta is
plentiful, B. diffusa seems to be scarce or even absent in some
regions. Maya names reported from Yucatan are "xaacil," "zacxiu,"
"zaciuthul," and "xacilsacxiu"; the Spanish name used there is
"hierba blanca." In Salvador this species is sometimes called
"escorian" and "golondrina." The leaves of this plant are said to
be cooked and eaten like spinach in the American Virgin Islands.

BOLDOA Lagasca

Tall coarse herbs, more or less glandular-pubescent in the inflorescence;
leaves alternate, petiolate, decurrent upon the petioles, thin, entire; flowers
perfect, not involucrate, ebracteate, small, glomerate and cymose-paniculate,
green; perianth herbaceous, subglobose or urceolate, 4-5-dentate, glandular and
covered with short uncinate hairs, little exceeding the fruit; stamens 3, inserted
on one side of the perianth, the filaments filiform, exserted, unequal, the anthers
didymous, the cells globose; ovary sessile, narrowed to a filiform style, the stigma
acute; fruit utricular, somewhat compressed, subglobose, coriaceous, costate on
one side; embryo hippocrepiform.

The genus consists of a single species.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 179

Boldoa purpurascens Cav. ex Lag. Gen. & Sp. Nov. 10. 1816.
B. ovatifolia Lag. loc. cit. Cryptocarpus globosus HBK. Nov. Gen.
& Sp. 2: 187. 1817. Salpianthus purpurascens Hook. & Arn. Bot.
Beechey Voy. 308. 1837. Hoja galan.

Damp thickets or in hedges, 200-600 meters; Zacapa; Jutiapa;
Santa Rosa. Mexico; Nicaragua; Cuba; Venezuela.

Plants 1-2 meters tall, often shrublike but really herbaceous, much branched,
the branches slender, green, subangulate, sparsely puberulent or glabrate, the
branches of the inflorescence viscid and bearing numerous short uncinate hairs;
leaves on petioles 1-10 cm. long, broadly rhombic-ovate to ovate-deltoid, 5-20
cm. long, 3-18 cm. wide, acute to attenuate, abruptly acute or acuminate at the
base and often long-decurrent, glabrate in age; flowers sessile or subsessile, glom-
erate or in short dense racemes at the ends of the panicle branches; perianth 2.5-3
mm. long, green, the teeth ovate-triangular, obtuse; fruit 1.5 mm. in diameter;
seed black, smooth, lustrous.

BOUGAINVILLE A Commerson

Shrubs or small trees, most often woody vines, glabrous or pubescent, often
armed with spines; leaves alternate, petiolate; flowers perfect, exinvolu crate,
usually in a 3-flowered axillary inflorescence consisting of 3 large persistent
colored bracts, a flower being borne on the inner surface of each bract, its pedicel
confluent with the costa of the bract; perianth tubular, the limb small, 5-lobate,
the lobes induplicate-valvate, the tube terete or 5-angulate; stamens 5-10, the
filaments capillary, somewhat unequal, connate at the base, the anthers didymous;
ovary stipitate, fusiform, slightly compressed laterally, the style short, filiform,
straight or slightly curved, included, papillose for part of its length; anthocarp
fusiform, coriaceous, 5-costate; embryo uncinate.

About 14 species, natives of South America, from the Andes of
Ecuador and central Brazil southward. Three species are found in
cultivation in most tropical regions of the earth. The name was
published originally as Buginvillaea, an erroneous form that does
not deserve perpetuation, especially since the group is horticulturally
important and has long been known by the form of the name used
here.

Leaves truncate or broadly rounded at the base; bracts crimson or orange, usually
rounded or broadly obtuse at the apex B. Buttiana.

Leaves mostly acute or acutish at the base; bracts purplish or magenta, usually
pointed at the apex and often acute B. glabra.

Bougainvillea Buttiana Holttum & Standl. Field Mus. Bot.
23: 44. 1944. Bombilla; Bugambilla; Pompilla.

Cultivated frequently for ornament in Guatemala, much less
common, however, than B. glabra. Doubtless native of Brazil, but

180 FIELDIANA: BOTANY, VOLUME 24

unknown at present in the wild state, although widely introduced
into cultivation since 1910 and now found in most tropical regions
of the earth.

A large vine, similar to the following species, but the leaves often larger,
usually broader, ovate-rounded or broadly elliptic-ovate, truncate or broadly
rounded at the base, even the uppermost leaves relatively broad; bracts crimson
or orange, relatively broader than in B. glabra, glabrous or nearly so.

This crimson Bougainvillea, much handsomer than the magenta
one, has become frequent in Central America in recent years and
may be seen rather commonly in Guatemala in the Centro or espe-
cially in gardens of the Pacific foothills. It is believed to have been
introduced from Cartagena, Colombia, to Trinidad in 1910, and its
general dispersal is thought to have taken place since that time.
The form with orange or apricot-colored bracts is very rare in Guate-
mala. A form with pale pink bracts, very rare in Guatemala,
probably is referable to B. glabra.

Bougainvillea glabra Choisy in DC. Prodr. 13, pt. 2: 437. 1849.
Bombilla; Buguenvilia; Bombilia; Boganbilla; Napoledn; Bogambilla ;
Gutembilla (Coban).

Planted for ornament in all except the colder parts of Guatemala,
from sea level up to the altitude of Quezaltenango (2,400 meters),
although rather uncommon at higher elevations. Native of Brazil,
but long grown for ornament in most tropical regions of the earth.

A large vine, the branchlets puberulent or glabrate, the spines short, often
somewhat recurved; leaves petiolate, broadly ovate to ovate-lanceolate, 4-10 cm.
long, gradually or abruptly acute or acuminate; puberulent when young but soon
glabrate; bracts broadly ovate to oval, mostly 2.5-4.5 cm. long, sometimes acumi-
nate, sparsely puberulent or almost glabrous; fruit 7-13 mm. long, puberulent or
glabrate.

Called "Napoleona" in Honduras, and known in Salvador by
the names "buganvilla," "buganvilea," "manto de Jesus," and
"pomonce." In Guatemala the plant has a high reputation as a
remedy for coughs, especially whooping cough. The vine may be
seen about a vast number of dwellings in the warmer parts of Guate-
mala, and there are many fine displays of it on some of the larger
fincas. Sometimes it climbs high upon tall trees, but more often it
is trained over trellises or hedges, where it blooms for a great part
of the year. In the Parque Central of Guatemala there is a particu-
larly fine vine with a huge trunk, the branches trained far out on
every side and forming a vast arbor to shelter benches and walks.
Many other isolated vines of large size are found elsewhere in the

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 181

parks, and mention may be made of such a vine in the otherwise
uninteresting plaza of the village of El Choi (Baja Verapaz). At so
great an elevation as Quezaltenango the Bougainvillea is not common
but it seems to thrive, at least in protected places, and to flower
there throughout the year. Bougainvillea spectabilis Willd. has been
reported as planted in Guatemala but we have seen no Guatemalan
specimens. It is distinguished by having copious pubescence on
almost all parts of the plant.

COMMICARPUS Standley

Perennial herbs or shrubs, pubescent or glabrous, usually decumbent or reclin-
ing, the stems much branched, with enlarged nodes; leaves opposite, those of a pair
subequal, petiolate, broad, more or less succulent, entire or sinuate; flowers perfect,
umbellate or verticillate, pedicellate, each pedicel bracteate at the base, the bracts
forming an involucel; perianth funnelform or campanulate, corolla-like, white or
green, usually with a distinct tube, constricted above the ovary, the limb shall owly
5-lobate, induplicate-plicate; stamens 2-5, the filaments exserted, capillary,
unequal, connate at the base; anthers didymous; ovary stipitate, attenuate to a
filiform style, the stigma peltate; fruit cylindric-fusiform, symmetric, finely
costate vertically, pubescent or glabrous, bearing numerous wart-like glands;
embryo uncinate.

About 15 species, in the tropics of both hemispheres. Only two
species are found in North America and only the following one in
Central America.

Commicarpus scandens (L.) Standl. Contr. U. S. Nat. Herb.
12: 373. 1909. Boerhaavia scandens L. Sp. PI. 3. 1753.

Dry thickets, 500 meters or less; Zacapa; El Progreso. South-
western United States and Mexico; West Indies; Venezuela and
Colombia to Peru.

Plants usually clambering over shrubs, somewhat woody below, much
branched, the branches pale green, glabrous or obscurely puberulent about the
nodes; leaves on slender petioles 1-2 cm. long, broadly cordate-ovate to ovate-
deltoid, 2-6 cm. long, 1-4.5 cm. wide, attenuate to acute or rarely rounded at the
apex, deeply cordate to truncate at the base, rather succulent, slightly paler
beneath, glabrous or nearly so; umbels of flowers on peduncles 2-4.5 cm. long, the
pedicels 5-10 mm. long; bracts lanceolate or oblong, 2-3 mm. long, ciliolate; peri-
anth greenish yellow, 3-4 mm. long and broad, glabrous or rarely somewhat
puberulent; stamens usually 2, exserted; fruit 1 cm. long, 2 mm. thick, glabrous,
bearing few or numerous glands irregularly scattered along the costae.

MIRABILIS L.

Perennial herbs, rarely shrubs, sometimes annuals, erect or procumbent,
viscid-pubescent or glabrous, usually branched, the stems often swollen at the

182 FIELDIANA: BOTANY, VOLUME 24

nodes; leaves opposite, sessile or petiolate, entire or undulate, often asymmetric;
flowers perfect, involucrate, the involucre calyx-like, enclosing 1 to several flowers,
usually 5-lobate, often accrescent in age; perianth campanulate to funnelform or
salverform, the tube short or often greatly elongate, the limb 5-lobate, the lobes
induplicate-valvate, the perianth deciduous after anthesis; stamens 3-5, the
filaments capillary, unequal, circinnate, short-connate at the base, usually exserted,
the anthers didymous; ovary ovoid or subglobose, the style filiform, the stigma
long-papillose; anthocarp globose to obovoid, terete or 5-angulate or 5-sulcate,
often rugose or tuberculate, constricted at the base, glabrous or pubescent, muci-
laginous when wet; embryo uncinate.

About 60 species, one native of southeastern Asia, the others
American, in tropical and temperate regions.

Perianth 1 cm. long or shorter, the tube short.

Fruit conspicuously angulate; plants more or less woody, at least below.

M. pulchella.

Fruit terete; plants herbaceous M. violacea.

Perianth 1.5-5 cm. long, the tube much longer than the limb.
Limb of the perianth scarcely broader than the tube; stamens 3.

M. Watsoniana.

Limb of the perianth several times as broad as the tube; stamens 5.

Perianth usually 7-9 cm. long, white or tinged with lavender or purple.

M. longi flora.
Perianth 3-5 cm. long, generally purple-red but variable in color.

M. Jalapa.

Mirabilis Jalapa L. Sp. PL 177. 1753. Maravilla.

A weed in waste or cultivated ground, also much cultivated for
ornament, chiefly in the lowlands but cultivated and escaping up to
2,500 meters or more; Pete"n; Alta Verapaz; Zacapa; Jalapa; Jutiapa;
Santa Rosa; Escuintla; Sacatepe"quez; Chimaltenango; Huehuete-
nango; Quezaltenango; Retalhuleu. Native of tropical America but
unknown in a truly wild state; cultivated in most tropical regions
of the earth, and also grown as a summer garden flower in temperate
regions.

A stout erect perennial a meter high or less, blooming the first year from seed,
the root usually large and fleshy, much branched, the stems glabrous or puberulent
or rarely villous; leaves slender-petiolate, ovate-deltoid to broadly ovate or lance-
ovate, acute to attenuate at the apex, subcordate or rounded at the base and short-
decurrent, glabrous or rarely puberulent; peduncles mostly 1-2 mm. long, cymose-
glomerate at the ends of the branches; involucre campanulate, 7-15 mm. long,
glabrous, puberulent, or short-villous, the lobes longer than the tube, linear-lanceo-
late to lance-ovate, acute to attenuate, usually ciliate; perianth 3-5.5 cm. long,
variable in color, most often purple-red, glabrous or sparsely villous outside, the
tube 2-5 mm. thick, the limb 2-3.5 cm. broad; stamens 5, equaling or slightly
exceeding the perianth; fruit obovoid or oval, 7-9 mm. long, 5-angulate, verrucose
or rugose, dark brown or black, glabrous or puberulent.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 183

The usual name for the four-o'clock throughout Central America
is "mara villa," but in Honduras the name "clavellino" is sometimes
applied. The Maya name is reported from Yucatan as "tutsuixiu."
Presumably this plant has been common in gardens of tropical
America since long before the Conquest, and if it ever grew wild it
has now become extinct. At present it seldom or never is found
outside cultivation far from cultivated ground. Its place of origin
may well be Mexico, for the most closely related species is probably
Mirabilis longiflora L., a white-flowered plant that occurs wild in
Mexico. The plant attracted attention from early European
explorers because of its extraordinary habit of producing flowers of
different colors upon the same plant or even upon the same branch.
This characteristic gave it its usual name of "mara villa" (marvel),
equivalent more or less to one of its English names, "marvel of
Peru." The flowers vary from red-purple to pink, white, and yellow,
and very often have longitudinal stripes of different colors. While
the Guatemalan plants are rather uniform in most characters, some
specimens noticed about Coban were notable in having white flowers
of only half the normal size. Rarely, too, the stems are densely
villous. The tuberous roots are said to supply a good food for pigs.
About Coban the Indians have the belief that aradores (redbugs or
chiggers) are particularly abundant upon the plant. The English
name, "four-o'clock," refers to the fact that the flowers open in the
evening and remain so during the night, closing at some time during
the following forenoon. They are exceedingly fragrant, especially
at night. It is said that the root is much used as a purgative by the
country people of Guatemala. It is first dried and pulverized, then
administered in sweetened water.

Mirabilis longiflora L. Svensk. Vet. Akad. Handl. 176. 1755.

Along streams, about 2,800 meters; Huehuetenango (near
deserted ranch house below Calaveras, Steyermark 50340). South-
western United States; Mexico.

Plants erect, branched, a meter high or less, the stems densely viscid-puberu-
lent or short- villous; leaves short-petiolate or the upper ones sessile, the petioles
usually less than 1 cm. long; leaf blades deltoid-ovate to lance-ovate, 6-11 cm.
long, acute to long-attenuate, cordate at the base, usually densely viscid-puberu-
lent; inflorescence of numerous, dense, axillary or terminal, leafy glomerules, these
often subtended by linear bractlike leaves; involucres short-pedunculate, cam-
panulate, 10-15 mm. long, densely glandular- villous, the lobes equaling or slightly
exceeding the tube, triangular or triangular-lanceolate; perianth 7-10 cm. long or
sometimes longer, white tinged with lavender or purple-red, the tube very slender,
2 mm. in diameter, abruptly expanded into a 5-lobate limb 2-3 cm. broad ; stamens 5,

184 FIELDIANA: BOTANY, VOLUME 24

exserted; fruit ellipsoid, 8 mm. long, obtusely 5-angulate, tuberculate, densely
puberulent between the tubercles.

Mirabilis pulchella Standl. & Steyerm. Field Mus. Bot. 23: 5.
1943.

Damp thickets and brushy rocky slopes, 400-600 meters; Zacapa
(type collected near divide on road between Zacapa and Chiqui-
mula, Standley 73841); Chiquimula.

An erect herb or shrub 30-100 cm. tall, branched, the older branches more or
less glaucous, the young ones densely villous; leaves slender-petiolate, the upper
ones short-petiolate, broadly ovate to oblong-ovate or deltoid-ovate, 3-8.5 cm.
long, 2.5-5.5 cm. wide, acute or acuminate, more or less unequal at the base and
truncate to obtuse, thick, sparsely villous on both surfaces; peduncles axillary and
arranged in small terminal cymes or racemes; involucre campanulate or in age
almost rotate, 7-10 mm. long, green, villous, 3-flowered, deeply 5-lobate, the lobes
triangular-ovate, acute or acuminate, ciliate; perianth rose-purple, about 1 cm.
long, funnelform-campanulate; stamens 3; fruit obovoid, about 5 mm. long and
2.5 mm. wide, obtusely 5-costate, narrowed at the base, pilosulous.

A rather handsome and showy plant, at least in the early morn-
ing, because of the abundant bright-colored flowers. These close
about noon, as is the case in most other members of this genus.

Mirabilis violacea (L.) Heimerl, Beitr. Syst. Nyctag. 23. 1897.
Allionia violacea L. Syst. ed. 10. 890. 1759. Oxybaphus violaceus
Choisy in DC. Prodr. 13, pt. 2: 432. 1849.

Moist thickets, 200-1,500 meters; Pete"n; Zacapa; Santa Rosa;
Guatemala. Southern Mexico; Honduras to Costa Rica; Colombia
and Venezuela.

Plants ascending or procumbent, lax, branched, the stems slender, a meter
long or less, bifariously puberulent or glabrate; leaves on slender petioles 1-6 cm.
long, usually broadly ovate-deltoid, sometimes ovate-oblong or elongate-deltoid,
2-8 cm. long, 1.5-5 cm. wide, usually attenuate to long-attenuate at the apex,
subcordate or truncate at the base, thin, bright green, puberulent, short-pilose,
or glabrate; inflorescence cymose, the cymes usually small and congested or in age
open, the branches viscid-pilose; involucres about 3 mm. long in anthesis, in fruit
5-6 mm. long, green, viscid-pilose, the lobes triangular-ovate, usually acute or
acuminate; perianth 5-6 mm. long, red-purple, viscid-pilose; stamens usually 3,
short-exserted ; fruit obovoid, 3.5-4 mm. long, terete, dark brown or blackish,
short-pilose, sparsely and irregularly tuberculate.

Names reported from Yucatan are "xpacumpac" (Maya) and
"hierba del golpe." This and M. pulchella are referable to the sub-
genus Oxybaphus which often has been treated as a distinct genus.
The group is connected by so many intermediate forms with typical

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 185

Mirabilis, as represented by M. Jalapa, that it can not be main-
tained as a distinct genus.

Mirabilis Watsoniana Heimerl, Bot. Jahrb. 11: 84. pi. 2, /.
2a-2h. 1889. Maravilla.

Usually at the base of cliffs, 1,350-2,200 meters; Solola (type
from Cuesta de Solola, Bernoulli & Carlo 2616); Huehuetenango;
El Progreso; endemic.

Plants erect or procumbent, much branched, densely leafy, the stems puberu-
lent above, glabrous or glabrate below; lower leaves long-petiolate, the upper ones
sessile or subsessile, ovate-deltoid to rounded-ovate, about 3.5 cm. long and 2.5
cm. wide or smaller, acuminate to obtuse or narrowly rounded at the apex, truncate
or cordate at the base and usually short-decurrent, thin, sparsely short-villous or
nearly glabrous, ciliate; peduncles slender, short-villous, arranged in dense leafy
clusters at the ends of the branches; involucre tubular-campanulate, unequally
5-lobate, short-villous, slightly accrescent in age and 8-9 mm. long, the lobes
lance-oblong, long-ciliate; perianth red-purple, 1.5-2 cm. long, tubular, short-
villous, slightly dilated upward, the limb scarcely broader than the tube, 5-lobate;
stamens 5, subequal, long-exserted ; fruit obovoid-pyramidal, dark brown, con-
stricted at each end, 5-angulate, the angles subtuberculate, the sides smooth,
puberulent.

Apparently a rare plant. The type locality is presumably the
steep descent along the road from Solola to Lake Atitlan, where
neither of the writers has collected.

NEEA Ruiz & Pavon

Shrubs or trees, glabrous or pubescent; leaves opposite or verticillate, usually
petiolate, membranaceous or subcoriaceous; flowers unisexual, dioecious, com-
monly with abortive organs of the other sex, small, white, green, or reddish, sessile
or pedicellate, usually 3-bracteolate, in axillary or terminal cymes; staminate
perianth urceolate, globose, or elongate, shortly 4-5-dentate; stamens 5-10,
included, the filaments unequal, the anthers oblong; pistillate perianth urceolate
or tubular, constricted above the ovary, 4-5-dentate and often contracted at the
mouth; ovary narrowly ovoid, the style terminal, filiform, often exserted, the
stigma penicillate; fruit ellipsoid, usually crowned by the persistent free portion
of the perianth, the stone hard, usually striate or costate; embryo straight.

A genus of about 70 poorly marked species, in tropical America.
A few besides those listed here are known from southern Central
America, and about 18 are known from all North America.

Upper surface of the leaves puberulent or pilose, the lower surface densely pilose
or villous N. fagifolia.

Upper surface of the leaves glabrous, rarely somewhat puberulent along the veins,
the lower surface usually glabrous or nearly so.

186 FIELDIANA: BOTANY, VOLUME 24

Leaves linear-lanceolate to lance-oblong, gradually long-attenuate or long-
acuminate, mostly 3-6 times as long as wide.
Leaves 1-4 cm. wide, often lustrous on the upper surface. . . .N. stenophylla.

Leaves mostly 6-10 cm. wide, not lustrous N. acuminatissima.

Leaves oblong to elliptic or obovate, obtuse to abruptly acuminate, mostly less

than 3 times as long as broad.
Leaves thin, mostly obtuse or acute, sometimes short-acuminate; pubescence

of the inflorescence chiefly grayish AT. psychotrioides.

Leaves coriaceous or subcoriaceous, usually abruptly acuminate; pubescence

of the inflorescence chiefly or in large part ferruginous.
Inflorescences usually abruptly recurved; fruit about 6 mm. long.

N. choriophylla.
Inflorescences mostly erect; fruit 10-12 mm. long or larger. . .N. belizensis.

Neea acuminatissima Standl. Field Mus. Bot. 4: 304. 1929,

Dense wet mixed forest, 150 meters or less; Izabal. British
Honduras; Atlantic lowlands of Honduras (type collected near Tela).

A shrub of 1-2 meters or sometimes a tree of 6 meters with a trunk 7 cm. in
diameter, sparsely branched, the branches glabrous; leaves opposite, on petioles
1-1.5 cm. long, rather thick, oblong or lance-oblong, 20-38 cm. long, 6-12 cm.
wide, narrowly long-acuminate, gradually narrowed to the unequal base, glabrous,
the lateral nerves about 18 on each side; pistillate inflorescence terminal, cymose-
paniculate, lax and few-flowered, 2.5-5 cm. long and broad, the branches glabrous
or sparsely and minutely puberulent, the stout pedicels 3-4 mm. long; fruit lance-
oblong, 12-16 mm. long, glabrous, narrowed to the apex, rounded at the base.

Neea belizensis Lundell, Contr. Univ. Mich. Herb. 7: 9. 1942.
Cerezo (Izabal).

Wet or rather dry forest or thickets, sometimes in pine forest,
most often on limestone, Atlantic watershed, ascending from sea
level to about 360 meters; Pete"n; Alta Verapaz; Izabal. Campeche
and British Honduras, along the Atlantic coast to Honduras and
Nicaragua; type from El Cayo, British Honduras, H. H. Bartlett
11445.

A shrub of 2 meters or sometimes a tree of 7 meters, the branches pale, terete,
rufous-puberulent at first but soon glabrate; leaves chiefly or all opposite, often
very unequal, thick-membranaceous, darkening when dried, on petioles 5-10
mm. long, oblong to elliptic-oblong or obovate-oblong, broadest at or above the
middle, 8-20 cm. long, 3.5-8 cm. wide, abruptly acuminate, with a short or elon-
gate acumen, unequal at the base and acute or cuneate, glabrous, at least in age,
the lateral nerves 7-8 pairs; inflorescences pedunculate, erect, laxly branched,
sparsely or densely rufous-puberulent, the staminate flowers usually slender-
pedicellate; staminate perianth tubular-campanulate, 5-6 mm. long, acute at the
base, sparsely rufous-puberulent or almost glabrous; pistillate perianth tubular,
2.5-3 mm. long, rufous-puberulent; fruit lance-oblong or elliptic-oblong, red or
blackish, 10-12 mm. long or even larger.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 187

There is some variation in the Guatemalan collections referred
here, and it is quite possible that when an ampler series of specimens
is available additional species will have to be recognized in this group.
The material here called N. belizensis has been referred heretofore
to N. psychotrioides, a species to which it is closely related. A shrub
probably of this species was planted and flowering in the park at
Puerto Barrios in 1939. The plant, however, is an inconspicuous
one and scarcely worthy of cultivation anywhere.

Neea choriophylla Standl. Contr. U, S. Nat. Herb. 13: 384.
1911. N. sphaerantha Standl. loc. cit.

Pete"n. Yucatan and northern British Honduras.

A slender shrub, commonly a meter high, or taller, the branches terete, pale,
glabrous or when young sparsely rufous-puberulent; leaves on petioles 5-10 mm.
long, oval to oval-oblong or oblong-obovate, mostly 5-7.5 cm. long and 2-3.5 cm.
wide, broadest at or above the middle, abruptly short-acuminate, usually broadly
cuneate at the base, subcoriaceous at maturity, sparsely puberulent beneath when
young but soon glabrate, the lateral nerves 5-8 on each side; peduncles usually
abruptly reflexed, at least after anthesis, 1.5-3 cm. long, the pistillate cymes 1.5-3
cm. broad or in fruit larger, the branches rufous-puberulent, the flowers sessile or
short-pedicellate, the perianth tubular-funnelform, 3 mm. long, minutely and
sparsely puberulent; staminate perianth urceolate or subglobose, 4-5 mm. long
and nearly as broad; stamens 6; fruit ellipsoid, when dry about 6 mm. long or
slightly larger, in the fresh state probably much larger.

The Maya name is reported from Yucatan as "xtadzi."

Neea fagifolia Heimerl, Beitr. Syst. Nyctag. 39. 1897.

Chiquimula, rocky outcrops along Rio Chiquimula between
Santa Barbara and Petapilla, 400 meters, Steyermark 30264- Type
from Granada, Nicaragua.

A tree 6 meters tall, the branches sparsely tomentulose at first, glabrate,
densely leafy; leaves opposite, on stout petioles 6-10 mm. long, elliptic-lanceolate
to oblong-elliptic, 4-8 cm. long, 2.5-4 cm. wide, subacute at the apex and often
apiculate, acute or subobtuse at the base, rather thick, lustrous above, glabrate
or permanently pilose, densely short-pilose beneath, the lateral nerves 5-7 on each
side; peduncles of the staminate inflorescence 2-3 cm. long, the cymes short-
pyramidal, the flowers sessile or short-pedicellate, the perianth ellipsoid, 5 mm.
long, somewhat narrowed at each end, glabrous; stamens 6; pistillate perianth
tubular, 4-5 mm. long, puberulent.

Neea psychotrioides Donn. Smith, Bot. Gaz. 16: 199. 1891.
Palo de sangre (fide Aguilar).

Dry or wet thickets or forest of the Pacific lowlands, often
extending upon the plains, 400 meters or less; Escuintla (type from

188 FIELDIANA: BOTANY, VOLUME 24

Escuintla, J. D. Smith 2069); Guatemala; Suchitepe"quez ; Retal-
huleu; San Marcos. Chiapas to Salvador, and perhaps southward
to Costa Rica and Panama.

A shrub 2-3 meters high, or sometimes a tree of 8 meters or more with thick
trunk and low dense crown, the branches mostly ochraceous, grayish-puberulent
when young but soon glabrate; leaves opposite or the upper ones verticillate, on
petioles 1 cm. long or less, oblong to elliptic-oblong, mostly 4-14 cm. long and 2-4.5
cm. wide, usually acute or obtuse, narrowly or broadly cuneate at the base, rather
thin, glabrous or nearly so, the lateral nerves about 10 on each side; staminate
cymes erect, pedunculate, terminal and axillary, lax and many-flowered, 5-10 cm.
broad, the branches slender, usually grayish-puberulent, the pedicels 1-5 mm. long,
the perianth tubular or suburceolate, 5-8 mm. long, minutely puberulent; stamens
5; pistillate perianth tubular, 3-4 mm. long, puberulent; fruit ellipsoid-oblong,
7-9 mm. long or even larger, red or dark red.

Known in Salvador by the names "frutilla," "sangre de chucho,"
"puruma," and "tenidor." The tree is a rather inconspicuous one,
common in some places along the Pacific plains, with no outstanding
characters by which it may be recognized easily.

Neea stenophylla Standl. Proc. Biol. Soc. Wash. 37: 51. 1924.

Moist or wet, mixed forest, 500 meters or less; Alta Verapaz(?);
Izabal (type from Puerto Barrios, Standley 25059) ; endemic.

A shrub or tree 1-7 meters high, the branches slender, glabrous or when young
sparsely and obscurely puberulent; leaves opposite or ternate, on stout petioles
2-6 mm. long, linear-lanceolate or narrowly lance-oblong, 5-19 cm. long, 1-4 cm.
wide, long-attenuate, acute or obtuse at the base, firm, glabrous, lustrous above,
the lateral nerves obscure; pistillate cymes axillary, on short slender peduncles,
8-15-flowered, lax, sparsely ferruginous-puberulent, the flowers red-brown, sessile
or on pedicels 1 mm. long or less, the bracts minute; pistillate perianth tubular,
3 mm. long, sparsely and minutely ferruginous-puberulent or almost glabrous.

This species is well marked by its extremely narrow and relatively
small leaves.

PISONIA L.

Trees or shrubs, often woody vines, glabrous or pubescent, often armed with
spines; leaves opposite, usually petiolate, entire; flowers dioecious, small, greenish,
in sessile or pedunculate cymes, not involucrate, 2-3-bracteolate; staminate
perianth obconic-campanulate, the limb 5-dentate; stamens 6-8, exserted, the
filaments unequal, filiform, short-connate at the base; pistillate perianth tubular,
the limb 5-dentate; ovary elongate-ovoid, sessile, attenuate to the slender short-
exserted style, the stigma penicillate; fruit coriaceous, clavate or oblong, terete
and costate or 5-angulate, the angles or costae furnished with one or more rows of
viscid stipitate glands; embryo straight.

A group of 10-15 species, in tropical regions of both hemispheres.
One other Central American species is known, in Costa Rica.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 189

Plants usually unarmed P. Donnell-Smithii.

Plants armed with spines.

Staminate inflorescences mostly fasciculate, with 10 or fewer flowers; spines

straight P. fasciculata.

Staminate inflorescences solitary, many-flowered; spines straight or curved.
Mature fruit 7-10 mm. thick, the glands of the angles in 2 or more rows;

spines straight P. macranthocarpa.

Mature fruit 3-4 mm. thick, the glands of the angles mostly in a single row;
spines recurved P. aculeata.

Pisonia aculeata L. Sp. PL 1026. 1753. P. grandifolia Standl.
Contr. U. S. Nat. Herb. 13: 391. 1911, not Warb. 1891 (type from
Cubilgiiitz, Alta Verapaz, Tuerckheim 7954). Una de gato; Huele
de noche.

Dry or moist thickets, chiefly in the tierra caliente, on the Pacific
slope ascending to about 1,400 meters; most plentiful on the Pacific
plains; Pete"n; Alta Verapaz; Baja Verapaz; El Progreso; Izabal;
Zacapa; Escuintla; Solola; Quezaltenango; San Marcos. Southern
Florida and Mexico to British Honduras and Panama; West Indies;

South America; Asia.

'

A densely branched shrub or tree, often with a thick trunk, the branches often
elongate and usually recurved or more or less scandent, usually armed with short
stout recurved spines; branchlets densely puberulent or short-villous; leaves on
short or elongate, slender or stout petioles, very variable in outline, mostly elliptic-
oval to ovate-oblong, obovate-orbicular, or even suborbicular, 5-15 cm. long,
usually acute or subacute, narrowly cuneate to rounded at the base, glabrous or
puberulent above, beneath glabrous, puberulent, or short-villous; peduncles 1-5
cm. long or in fruit longer, the inflorescence loosely or densely cymose, 2-6 cm.
broad, many-flowered, the pedicels short, usually with viscid pubescence; stami-
nate perianth broadly campanulate, 2-4 mm. long, densely puberulent or tomentu-
lose, yellowish green; stamens usually 6, twice as long as the perianth; pistillate
perianth tubular, 2-3 mm. long; fruit clavate, 9-12 mm. long, 3-4 mm. in diameter,
rounded at the apex, narrowed to the base, the sides glabrate or puberulent.

Names recorded from adjacent regions are "crucito" and "caga-
lero negro" (Salvador); "beeb" (Yucatan, Maya); "cargalera"
(Honduras) . The glands of the fruit are exceedingly viscid and retain
their viscosity in the herbarium indefinitely, or at least for numerous
decades. The fruits often adhere to the feathers of birds, and it is
said that they sometimes cause the death of small birds that become
entangled among the branches. This shrub or vine often constitutes
a large part of the undergrowth on the Pacific plains, as in Escuintla
and Retalhuleu, and in such places progress is difficult because the
hooked spines catch in one's clothing or even in the flesh.

190 FIELDIANA: BOTANY, VOLUME 24

Pisonia Donnell-Smithii Heimerl ex Standl. Contr. U. S. Nat.
Herb. 13: 387. 1911.

Damp thickets or forest, 1,000-1,800 meters, Pacific bocacosta,
endemic; type from Los Verdes, Guatemala, 1,050 meters, Heyde &
Lux 6301; Escuintla; Guatemala; Sacatepe"quez.

A shrub or small tree of 3-5 meters, the branches stout, unarmed, or some-
times armed with short straight spines, when young appressed-pilose with short
hairs, soon glabrate, densely leafy; leaves often crowded on short lateral branches,
bright green when dried, on petioles 4-8 mm. long, obovate to obovate-oblong or
elliptic-oblong, 3-6 cm. long, 1.5-3 cm. wide, obtuse, at the base acute or attenuate,
sparsely puberulent or glabrate above, short- villous beneath along the costa;
staminate peduncles solitary, 2-3 cm. long, the inflorescence capitate-cymose,
2 cm. broad or less, the flowers short-pedicellate; staminate perianth narrowly
campanulate, 5 mm. long, pale green, minutely puberulent; pistillate flowers and
fruit unknown.

Apparently a rare species. It has been observed by the writers
only along the road between Antigua and Escuintla.

Pisonia fasciculata Standl. Contr. U. S. Nat. Herb. 13: 388.
1911. Crucito.

Dry rocky hillsides or plains, 100-200 meters; Zacapa. Nicara-
gua, whence the type.

A shrub or small tree 4.5 meters high, the branchlets sparsely puberulent when
young; spines few, stout, straight, 3-4 mm. long; leaves on petioles 4-5 mm. long,
oblong-elliptic to oval-elliptic, 3.5-4 cm. long, 1.5-2 cm. wide, acute or abruptly
acute, acute at the base, bright green, sparsely puberulent beneath along the costa,
at least when young, elsewhere glabrous; staminate peduncles in clusters of 2-5,
10-12 mm. long, viscid-villous, the cymes headlike, about 1 cm. in diameter, 5-10-
flowered, the flowers short-pedicellate; staminate perianth campanulate, 2-3 mm.
long, glandular-puberulent; stamens 6, almost twice as long as the perianth;
pisitllate flowers and fruit unknown.

Pisonia macranthocarpa Donn. Smith, Bot. Gaz. 20: 293.
1895. P. aculeata var. macranthocarpa Donn. Smith, Bot. Gaz. 16:
198. 1891. Clavo; Crucito; Palo caribe (fide Aguilar).

Dry thickets or forest, sometimes on rocky stream banks, 250-
800 meters; Zacapa; El Progreso; Jutiapa; Santa Rosa; Escuintla;
Guatemala; Retalhuleu; type from Escuintla, J. D. Smith 2091.
Chiapas to Costa Rica; Cuba; Venezuela.

A large shrub or small tree, sometimes 6 meters tall, the branchlets puberulent
at first, soon glabrate; spines few, often wanting on the branchlets, stout, usually
straight, 5-8 mm. long; leaves on petioles 5-25 mm. long, elliptic to broadly oval
or rarely obovate, acute to attenuate at each end, glabrous above, puberulent or
short-villous beneath along the costa; staminate peduncles 1.5-3 cm. long, the

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 191

cymes dense and many-flowered, 2-3.5 cm. broad, the flowers short-pedicellate;
perianth broadly campanulate, 3-4 mm. broad, yellowish green, puberulent;
stamens usually 8, twice as long as the perianth; pistillate perianth 3 mm. long,
funnelform; fruit ligneous, oblong or obovoid, 1-2 cm. long, 7-10 mm. thick,
truncate or depressed at the apex, acute at the base, the sides densely tomentulose.

Called "espuela del diablo" in Salvador and "cagalera prieta" in
Honduras.

TORRUBIA Vellozo

Unarmed shrubs or trees, glabrous or pubescent; leaves opposite, usually
petiolate, entire, often coriaceous; flowers dioecious, small, reddish, greenish, or
whitish, not involucrate, 2-3-bracteolate, sessile or pedicellate, in lateral or termi-
nal, pedunculate cymes; staminate perianth obconic-campanulate, the limb
5-dentate, the short teeth induplicate-valvate; stamens 6-10, exserted, the fila-
ments unequal, filiform, short-connate at the base, the anthers oblong; pistillate
perianth tubular, the narrow limb shallowly 5-dentate; ovary elongate-ovoid,
sessile, attenuate to the usually short-exserted style, the stigma penicillate; fruit
drupaceous, the exocarp fleshy and juicy, red or black, the stone elongate, coria-
ceous, striate; embryo straight.

Probably 40-50 species, mostly South American, about 18 in
North America. Two Central American species are known from
Costa Rica and Panama.

Leaves glabrous or nearly so T. linearibracteata.

Leaves densely pilose beneath T. petenensis.

Torrubia linearibracteata (Heimerl) Standl. Contr. U. S.
Nat. Herb. 18: 100. 1916. Pisonia linearibracteata Heimerl, Repert.
Sp. Nov. 12: 221. 1913.

Wet to dry thickets or forest, at or little above sea level; Pete*n;
Izabal. Yucatan (type from Chichen Itza) and British Honduras.

Usually a shrub of 2-3 meters, sometimes a small tree, the branchlets rufous-
puberulent at first, soon glabrate; leaves opposite or the uppermost verticillate,
on petioles 1 cm. long or less, rhombic-elliptic to lance-elliptic, rarely obovate or
oblanceolate-elliptic, about 7.5 cm. long and 4.5 cm. wide, often smaller, usually
short-acuminate at each end, glabrous or practically so; staminate peduncles 3-6
cm. long, glabrous or sparsely hirtellous, the inflorescence corymbose-paniculate,
3.5-6 cm. broad, many-flowered, usually lax, the branches rufous-puberulent, the
bracts linear, 1.5-3.5 mm. long, the pedicels 1-1.5 mm. long; perianth funnel-
form, 4-4.5 mm. long, puberulent; stamens 7-8; mature fruit purple or blackish,
with red juice, oval or broadly oblong, about 1 cm. long.

The flowers are whitish or dirty yellow. The Maya name is
recorded from Yucatan as "xtabdzi."

Torrubia petenensis Lundell, Carnegie Inst. Wash. Publ. 478:
208. 1937.

192 FIELDIANA: BOTANY, VOLUME 24

Known only from the type, in forest on top of limestone hill, La
Libertad, Pete"n, C. L. Lundell 3518.

A tree of 5 meters, the branches villous-tomentose; leaves firm-membrana-
ceous, blackish when dried, on slender petioles 1-2 cm. long, oblong-elliptic to
narrowly obovate, 7-9.5 cm. long, 2.5-4.5 cm. wide, obtuse or subacute with an
obtuse tip, acute or obtuse at the base, at first thinly villous above but soon gla-
brate, densely villous-tomentose beneath; staminate panicles appearing with the
leaves, broad and much branched, pedunculate, thinly villosulous, the flowers
short-pedicellate; perianth turbinate-campanulate, 4-5 mm. long, puberulent;
stamens 8, the slender filaments exserted.

PHYTOLACCACEAE. Pokeweed Family

References: H. Walter, Pflanzenreich IV. 83. 1909; Percy Wilson,
N. Amer. Fl. 21: 257-266. 1932.

Herbs, shrubs, or trees, sometimes scandent, occasionally armed with spines;
leaves alternate, usually entire, the stipules minute or usually absent; flowers
perfect or unisexual, in terminal or axillary racemes, rarely paniculate; sepals 4-5,
equal or unequal, often persistent in fruit; petals usually none, sometimes 5;
stamens 3-many, the filaments free or united at the base, the anthers 2-celled;
disk present or absent; gynoecium of 1-many carpels, these free or connate; ovary
superior or partly inferior; styles as many as the carpels, free or rarely connate
or almost none, the stigmas capitate, penicillate or papillose; ovules solitary in
each carpel, campylotropous; fruit drupe-like, berry-like, achene-like, or capsular;
seed erect, often compressed, the testa membranaceous or crustaceous; aril some-
times present and surrounding the seed; embryo annular, semi-annular, or erect;
cotyledons incumbent, foliaceous and plicate-convolute or linear and semi-
cylindric.

A small family, with 10 genera in North America. The only one
of these not represented in Central America is Phaulothamnus, of
Texas and northern Mexico.

Ovary partly inferior; a glabrous herbaceous vine; leaves cordate at the base.

Agdestis.
Ovary superior; leaves not cordate at the base.

Flowers with petals; ovary 1-celled, with 3-5 ovules; fruit capsular. Unarmed
shrubs Stegnosperma.

Flowers without petals; ovary 1-16-celled, with a single ovule in each cell;
fruit not capsular.

Gynoecium of 5-16 carpels Phytolacca.

Gynoecium 1-2-carpellate.
Flowers unisexual; trees, usually armed with spines. Fruit drupe-like.

Achatocarpus.
Flowers perfect; plants unarmed.

Fruit drupe-like.

Stamens 4; plants usually herbaceous throughout, sometimes woody
below; fruit red Rivina.

Stamens 8 or more; shrubs, usually scandent; fruit black . Trichostigma.
Fruit dry.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 193

Stamens 12; trees Ledenbergia.

Stamens 3-9; herbs, sometimes slightly woody at the base.

Fruit elongate and narrow, bearing 3-6 uncinate bristles at the apex;

sepals 4; stigma 1, penicillate Petiveria.

Fruit subglobose, echinate; sepals usually 5; stigmas 2, linear.

Microtea.

ACHATOCARPUS Triana

Shrubs or trees, often armed with spines; leaves alternate, entire, usually
blackening when dried; flowers dioecious, in simple or paniculate racemes, small
and greenish; sepals 5, persistent in fruit; staminate flowers with 10-15 stamens,
the filaments inserted at the base of the perianth segments, the anthers basifixed;
pistillate flowers without stamens or staminodia, the ovary somewhat compressed,
1-celled; stigmas 2 or rarely 3, linear or filiform, reflexed, papillose or fimbriate;
ovule 1, campylotropous; fruit drupe-like; seed 1, erect, black, with crustaceous
testa, the embryo annular, the endosperm farinaceous; cotyledons linear.

A dozen species are known, most of them South American. Only
one occurs in Central America.

Achatocarpus nigricans Triana, Ann. Sci. Nat. IV. 9: 46. 1858.
A. mexicanus H. Walt. Pflanzenreich IV. 83: 139. 1909. Ampelocera
hondurensis Bonn. Smith, Bot. Gaz. 54: 244. 1912 (type from San
Pedro Sula, Honduras).

Dense thickets near the coast or on low plains, 325 meters or
lower; El Progreso; Santa Rosa; Escuintla; Retalhuleu. Southern
Mexico; Salvador; Honduras; northern South America.

A dense tree 5-8 meters high, with a low trunk and spreading crown, the
branches often armed with stout sharp spines 7-10 mm. long, pale; leaves short-
petiolate, rather fleshy when green, coriaceous when dry, elliptic to elliptic-
lanceolate, 5-13 cm. long, 2.5-6 cm. wide, acute to obtuse or sometimes rounded
at the apex, acute at the base, glabrous; racemes simple or branched, mostly at
naked nodes, 3-6 cm. long, the flowers short-pedicellate, green; sepals elliptic or
obovate, 2.5-3 mm. long; stamens 12-16; pistillate sepals elliptic or oval, 2.5
mm. long; fruit subglobose, bluish black; seed compressed, 3.5 mm. in diameter.

Names given to this species in other regions are "limoncillo"
(Salvador); "huasicuco" (Michoacan); "palo dulce" (Veracruz,
Oaxaca); "limon-che"" (Campeche). The bark is described (in
Veracruz and Oaxaca) 'as medium brown, the inner bark reddish
brown; sapwood white to pale yellowish brown, the heartwood pale
greenish brown. It is said to be used for railroad ties in southern
Mexico. The flowers are described as fragrant.

AGDESTIS Mocino & Sesse

Herbaceous vines arising from large tuberous roots, glabrous; leaves alternate,
on long slender petioles, broad, membranaceous; flowers perfect, white, in axillary,

194 FIELDIANA: BOTANY, VOLUME 24

many-flowered, rather lax panicles; sepals usually 4; stamens 15-20, the filaments
filiform, the anthers dorsifixed; ovary partly inferior, 3-4-celled, the style cone-like,
the 3-4 stigmas subterete, erect in bud, recurved in an thesis, papillose; ovule
solitary, campylotropous; fruit small, turbinate, dry, surrounded by the persistent
sepals; seed lenticular, the testa crustaceous, the embryo annular; endosperm
sparse, farinaceous; cotyledons oblong.

The genus consists of a single species.

Agdestis clematidea Mocino & Sess£ ex DC. Syst. 1: 543. 1818.
Bejuco de ajo.

Moist or dry thickets, at low elevations; Izabal; El Progreso;
Santa Rosa; Suchitepe"quez; Retalhuleu; San Marcos; probably in
all the Pacific coast departments. Southern Mexico; Honduras.

A large branched vine, climbing high over bushes and small trees, the stems
very slender, the foliage ill-scented; leaves on very long and slender petioles, the
blades broadly ovate or suborbicular, 3-7 cm. long, 2-6 cm. wide, acute to broadly
rounded at the apex, shallowly or deeply cordate at the base, pale green; panicles
often 8-15 cm. long, the flowers fragrant, pedicellate; sepals white, oblong to
obovate, 4.5-6.5 mm. long, rounded or obtuse at the apex, reticulate- veined;
anthers 1.3 mm. long, oblong, cordate at the base.

The plant is an ornamental one and for that reason is sometimes
cultivated in distant regions, as in South America and the West
Indies. In some regions of the Pacific plains it forms dense tangles
over thickets, but the plants soon wither after the advent of the dry
season and are conspicuous only during the wet months. The leaves
are paler on the lower surface. When crushed they have a slight
garlic odor, or one somewhat suggestive of cabbage.

LEDENBERGIA Klotzsch

Shrubs or small trees; leaves membranaceous, slender-petiolate, alternate;
flowers perfect, in long racemes; sepals typically 4, rarely 5, accrescent and per-
sistent in fruit; stamens 12, the filaments filiform, the anthers dorsifixed; ovary
1-carpellate, 1-celled, the style subterminal, short, the stigma penicillate; ovule 1;
fruit dry, subglobose, somewhat compressed; seed erect, lenticular, the testa
crustaceous; embryo annular; endosperm farinaceous; cotyledons oblong.

Only two species are known, the other found in Venezuela and
Martinique.

Ledenbergia macrantha Standl. Journ. Wash. Acad. Sci. 13:
350. 1923. Flueckigera macrantha P. Wilson, N. Amer. Fl. 21: 260.
1932. Siete camisas.

Dry forests, about 1,300 meters; Guatemala (Lago de Ama-
titlan). Salvador, the type collected at Puerta de la Laguna.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 195

A tree about 6 meters high, sometimes 24 meters high, the branches more or
less pendulous, the young branchlets sparsely pubescent at first; petioles slender,
2-4.5 cm. long, sparsely pilose; leaf blades elliptic to broadly ovate, 4-8 cm. long,
2.5-4.5 cm. wide, acute, acuminate, or obtuse, at the base acute or obtuse, glabrous
above, pilose beneath along the costa, paler beneath; racemes pendent, 10-15 cm.
long or more; sepals oblong-oblanceolate, in fruit 8-13 mm. long, glabrate, con-
spicuously veined; fruit ellipsoid, 3 mm. long.

The name "nevado" is said to be given the tree in Salvador.

MICROTEA Swartz

Small annuals, erect or decumbent, rather succulent; leaves alternate, sessile
or petiolate, small; flowers minute, green, racemose or paniculate; sepals 5, rarely
4; stamens 3-9, the filaments linear, the anthers didymous; ovary 1-celled, with
2 linear stigmas; ovule 1, campylotropous; fruit subglobose, fleshy, smooth,
tuber culate, or echinate; seed erect, the testa crustaceous; embryo semi-annular;
endosperm fleshy; cotyledons elongate, concave.

About 9 species, in tropical America. Only one is known from
continental North America.

Microtea debilis Swartz, Prodr. Veg. Ind. Occ. 53. 1788.

Occasional in moist thickets or moist open ground, sometimes on
gravel bars or in waste ground about dwellings, lowlands of both
coasts, at or little above sea level; Izabal; San Marcos (near Mala-
catan). British Honduras to Panama; West Indies and South
America.

A glabrous annual, often much branched, prostrate or ascending; leaves
slender-petiolate, bright green, thin when dried, somewhat succulent when fresh,
spatulate to obovate, or the upper ones lanceolate to ovate, 5-9 cm. long and 1.5-3
cm. wide, or often smaller, acute to rounded at the apex, cuneate at the apex or
often abruptly contracted and long-decurrent; flowers small, green or greenish
white, in slender few-many-flowered axillary pedunculate racemes, short-pedicel-
late; sepals 5, elliptic, 0.5 mm. long, acutish; stamens 5, with minute anthers;
stigmas 2, ovate-triangular or triangular-lanceolate; fruit subglobose, green, about
1 mm. long, spinose-tuberculate and reticulate; seed black.

A small and inconspicuous weed, not common in Guatemala.

PETIVERIA L.

Plants erect, with an odor of garlic, herbaceous or somewhat woody, branched;
leaves alternate, petiolate, membranaceous, with minute stipules; flowers small,
greenish, perfect, in terminal and axillary, spikelike racemes; sepals 4, spreading
in an thesis, persistent and erect in fruit; stamens 4-9, the filaments subulate, the
anthers linear, 2-cleft at base and apex; ovary 1-celled, 1-carpellate, with 3-6
deflexed uncinate processes at the apex; stigma sessile, penicillate on the ventral

196 FIELDIANA: BOTANY, VOLUME 24

side; fruit achene-like, long and narrow, carinate on both sides, bilobate at the
apex and bearing 3-6 uncinate bristles; seed erect, linear, with scant endosperm,
the embryo erect, the cotyledons foliaceous, unequal.

One other species is known, in Brazil.

Petiveria alliacea L. Sp. PL 342. 1753. Apacina; Hierba de
zorrillo; Zorrillo; Apazote de zorro; Epacina; Ipacina; Apacin; Epacin;
Hierba de zorro.

Moist or dry fields, thickets, or even forest, frequent about
dwellings, especially in hedges and waste ground, chiefly in the tierra
caliente, but ascending to about 1,500 meters (at Antigua and
perhaps elsewhere) ; Alta Verapaz; Zacapa; Chiquimula; Santa Rosa;
Escuintla; Guatemala; Sacatepe"quez; Retalhuleu; San Marcos;
doubtless in all the lowland departments. Florida and Texas to
Mexico, British Honduras, and Panama; West Indies and South
America.

Plants usually stiffly erect and about a meter high or lower, often woody
below, the young branches puberulent or glabrate; petioles 1.5 cm. long or less, the
leaf blades oblong to elliptic or obovate, 5-15 cm. long, 2-6 cm. wide, acuminate
to rounded at the apex, narrowed to the acute or cuneate base, bright green, thin,
glabrous or sparsely pubescent; racemes slender, 10-35 cm. long, rather remotely
flowered, the flowers subsessile or on very short pedicels; sepals greenish white,
oblong-linear, 3.5-4 mm. long; fruit appressed to the rachis of the raceme, narrowly
cuneate, about 8 mm. long.

The Maya name used in Yucatan is "payche"" (skunk plant).
Called also "hierba de las gallinitas" in Yucatan, and in British
Honduras "guinea-hen root" and "skunkweed." The whole plant
has a most disagreeable odor suggestive of garlic, and it is said to
impart this odor and an unpleasant flavor to the milk of cows that
eat the foliage. The plant is much used in domestic medicine
throughout the American tropics. In the Jocotan region it is
reported to be administered to induce menstruation. The most
current name in Guatemala is "apacina." It is an unpleasant and
offensive plant because the hooked spines of the fruit cling tena-
ciously to clothing and also penetrate the skin readily if one brushes
against the branches. Because of the hooked tips of the bristles,
they can be withdrawn only with difficulty from the skin. The
plant is particularly plentiful in many of the dry thickets of the
Pacific plains.

PHYTOLACCA L. Pokeweed; Pokeberry

Coarse perennial herbs with thick roots, sometimes shrubs or trees, the stems
erect or sometimes weak and subscandent, glabrous or somewhat pubescent; leaves

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 197

often large, alternate, petiolate or sessile; stipules none; flowers small, perfect or
dioecious, white, greenish, or reddish, in simple or paniculate racemes or spikes,
terminal or axillary; pedicels bracted at the base and often bearing 1-2 bractlets
above the base; sepals 5, equal or unequal; stamens 6-33, 1-2-seriate, the fila-
ments subulate or filiform, free or somewhat connate at the base, the anthers
oblong or elliptic, dorsifixed; ovary subglobose, composed of 5-16 distinct or some-
what united carpels; fruit depressed-globose, 5-16-celled, fleshy and juicy; seeds
1 in each carpel, erect, compressed; embryo annular, the endosperm farinaceous;
cotyledons semiterete.

About 25 species in tropical and warmer regions of America,
Africa, and Asia. Only the following species are found in Central
America. Four additional ones are known from the United States,
Mexico, and West Indies, one of them extending northward to
southern Canada.

Sepals 5-6 mm. long P. Meziana.

Sepals 2.5-3.5 mm. long.

Pedicels 5-10 mm. long, much longer than the bracts; racemes mostly 10-50

cm. long, usually several timeb longer than the leaves P. rivinoides.

Pedicels mostly less than 5 mm. long, usually equaled or exceeded by the bracts;
racemes short, mostly 15 cm. long or less, often shorter than the leaves,
usually about equaling them, or but slightly longer.

Carpels of the ovary free at the apex in anthesis P. rugosa.

Carpels united to the apex in anthesis P. icosandra.

Phytolacca dioica L. Sp. PI. ed. 2. 632. 1762.

A species of southern Brazil, Paraguay, Uruguay, and Argentina,
the famous "ombu" of the last country. It is planted in the Jardin
Botanico and Finca La Aurora in Guatemala, and probably else-
where about the city. It differs from all the native species in being
a tree or large shrub.

Phytolacca icosandra L. Syst. Nat. ed. 10. 1040. 1759. P.
octandra L. Sp. PL ed. 2. 631. 1762. P. sessiliflora Kunth & Bouch^,
Ind. Sem. Hort. Berol. 1848: 15. 1849. P. octandra var. angusti-
folia Moq. in DC. Prodr. 13, pt. 2: 32. 1849. P. purpurascens
Braun & Bouche", Ind. Sem. Hort. Berol. 1851: 13. 1852 (type col-
lected in Guatemala by Warscewicz). P. icosandra var. sessili-
flora H. Walt. Pflanzenreich IV. 83: 61. 1909. Jaboncillo; Almorsaca;
Mazorquilla; Uaxit (fide Mrs. Osborne); Ixmaxin (Quezaltenango) ;
Amorzacate.

Moist fields or thickets or open slopes, sometimes in pine forest,
often in waste or cultivated ground, widely distributed, ascending
to 2,900 meters, most plentiful at middle or rather high elevations,

198 FIELDIANA: BOTANY, VOLUME 24

and seldom found in the tierra caliente; Pete*n; Alta Verapaz; El
Progreso; Zacapa; Jalapa; Escuintla; Guatemala; Sacatepe'quez ;
Chimaltenango; Solola; Quiche"; Huehuetenango; Quezaltenango;
Totonicapan. Mexico and British Honduras to Panama; West
Indies and South America; naturalized in some parts of the Old
World tropics.

A coarse, somewhat succulent herb 1-2 meters tall, branched; leaves slender-
petiolate, thin, narrowly elliptic to ovate-elliptic, 7-20 cm. long and 3-10 cm. wide
or even larger, acute or acuminate, attenuate or acuminate at the base, glabrous;
racemes terminal and axillary, numerous, mostly 8-15 cm. long, the rachis some-
what pubescent; pedicels 2-5 mm. long or sometimes none, the bracts subulate,
equaling or usually longer than the pedicels; sepals greenish white or red-purple,
2.5-3 mm. long, persistent; stamens 8-20; ovary of 6-10 carpels, these united to the
apex in flower; styles recurved; fruit depressed-globose, about 8 mm. in diameter,
green and red, turning purple-black; seeds black and lustrous, about 2.5 mm. long.

Called "calaloo" and "scorpion-tail" in British Honduras, and
"quilete" in Honduras. The Maya name used in Yucatan is "telcox"
or "telcocox." This plant and the other local species are of great
economic importance in Guatemala as a soap substitute. Through-
out the highlands, but especially in San Marcos, Quezaltenango, and
Totonicapan, great quantities of the green berries are gathered by
Indian women and children and used at home or sold in the markets.
The ripe fruits are not gathered, because they would leave stains,
but the green ones when macerated in water give a copious suds that
is found satisfactory for cleaning clothes. Along the Atlantic coast
of Central America, especially by the people of African origin, the
young shoots and leaves are much used as a pot herb, but we have
not seen them so used in the mountains of Guatemala. The juice
of the ripe fruits gives a red-purple color that is sometimes used for
ink or for coloring various small articles. There is a popular belief
in some regions that the fruits are poisonous, but they are some-
times eaten in at least small amounts by children, in both Central
America and the United States. The roots are known to be poison-
ous. Dried, they are employed in the United States as a remedy for
garget (caked udder) in cows, and formerly at least they were sold
commonly in drug stores for this purpose. The plants have been
much used in domestic or even official medicine in both America
and Europe. In Guatemala the fruits are said to be a favorite food
of the sensontles, the local mockingbirds, and are fed to those kept
in cages. In the Totonicapan region there were noted some plants,
probably of this species, that had white flowers, pale green leaves,
and pale green fruit, probably an albino form. The name "calalu,"
often applied to Phytolacca species in the Atlantic coast of Central

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 199

America, is believed to be of African origin. Phytolacca octandra is
maintained as distinct by Walter, on the basis of complicated stamen
characters, but Wilson is probably right in reducing it to synonymy
and thus greatly simplifying the taxonomy of the North American
species.

Phytolacca Meziana H. Walt. Pflanzenreich IV. 83: 57. 1909.
P. icosandra var. octogyna Bonn. Smith, Bot. Gaz. 18: 210. 1893.
Pinta cashorro (fide Aguilar).

Moist or wet forest or thickets, 2,000-2,600 meters; endemic; El
Progreso; Quiche* (type from San Miguel Uspantan, Heyde & Lux
3031).

A tall herb with glabrous branches; leaves slender-petiolate, elliptic-oblong or
ovate-oblong, 10-13 cm. long, 3-4.5 cm. wide, acute to long-acuminate, acute at the
base; racemes many-flowered, 15-18 cm. long and 2.5 cm. broad, the rachis
pubescent, the pedicels 7-8 mm. long; bracts subulate, equaling or longer than the
pedicels; sepals oblong-elliptic, 5-6 mm. long, rounded at the apex; stamens 12-25,
shorter than the sepals; carpels 7-8, the styles erect, recurved at the apex.

Herbarium specimens of this species are easily recognized because
they seem always to blacken in drying, those of other species usually
remaining green.

Phytolacca rivinoides Kunth & Douche", Ind. Sem. Hort. Berol.
1848: 15. 1849. Jaboncillo; Calalu (North Coast); Pinta-machete ;
Sacachdn (Huehuetenango) ; Yakl (Tactic, Alta Verapaz).

Damp or wet thickets or forest, ascending from sea level to about
2,600 meters (on the Pacific slope); Alta Verapaz; Izabal; Retal-
huleu; Quezaltenango; Huehuetenango. Southern Mexico and
British Honduras to Panama; West Indies and South America.

Plants erect and 1-1.5 meters tall, or often more elongate, as much as 3 meters
long, and supported on other vegetation, glabrous or practically so; leaves thin,
bright green, slender-petiolate, elliptic to ovate-lanceolate, mostly 10-18 cm. long
and 3-9 cm. wide, acuminate or long-acuminate, acute or cuneate at the base;
racemes pedunculate, many-flowered, 20-70 cm. long, rather lax, often recurved
or pendent, the pedicels divaricate, 5-10 mm. long, the subulate bracts shorter
than the pedicels; sepals pink, elliptic or oval, 2.5 mm. long, usually early decidu-
ous; stamens 9-22, shorter than the sepals; ovary depressed-globose, 10-16-car-
pellate, the styles cylindric, recurved; fruits black or purple-black, 7 mm. broad;
seeds suborbicular, 2 mm. long, scarcely lustrous.

Called "quilete" and "cola de ardilla" in Honduras. The rachis
of the inflorescence is usually bright carmine. The whole plant is
rather showy and, because of its habit and long inflorescences, much
handsomer than in the other species. The young shoots are cooked

200 FIELDIANA: BOTANY, VOLUME 24

and eaten in Huehuetenango and probably in other parts of the
country.

Phytolacca rugosa Braun & Douche", Ind. Sem. Hort. Berol.
1851: 13. 1852. Jabon; Sacchen (San Antonio de San Marcos);
Mazorquilla; Jaboncillo.

Damp or wet forest and thickets, mostly at 1,800-2,800 meters;
type collected in Guatemala by Warscewicz; Guatemala; Sacate-
pe"quez; Quiche"; Quezaltenango; San Marcos; Huehuetenango.
Southern Mexico; Honduras; Costa Rica; Panama; Colombia and
Venezuela.

A coarse herb 1-2 meters tall, often densely branched, almost glabrous; leaves
slender-petiolate, elliptic to elliptic-lanceolate, 6-17 cm. long and 3-6 cm. wide or
larger, thin, acuminate at each end; racemes mostly short but sometimes longer
than the leaves, the pedicels 3-4 mm. long, the bracts usually equaling the pedicels;
sepals pink or purplish red, oblong-elliptic, 2.5-3 mm. long, rounded at the apex,
usually persistent and often recurved in fruit; stamens 8-10; ovary usually 8-car-
pellate, the styles cylindric, recurved; fruit depressed-globose, about 6 mm. in
diameter; seeds subreniform, 2.5 mm. long.

This species is particularly abundant in the highlands of San
Marcos, where large quantities of its fruit are gathered. It is too
closely related to P. icosandra, and separated from it sometimes only
with difficulty.

RIVINA L.

Plants annual or perennial, erect, herbaceous or somewhat woody at the base,
glabrous or pubescent; leaves alternate, slender-petiolate, membranaceous, without
stipules; flowers perfect, racemose, small, the racemes terminal or pseudolateral,
the pedicels bracteate at the base and also bearing bractlets above; perianth
corolla-like, the 4 segments subequal, elliptic or obovate-oblong, rounded or
pointed at the apex, persistent and slightly accrescent in fruit, becoming recurved;
stamens 4, inserted on a small hypogynous disk, shorter than the sepals, the fila-
ments cylindric-filiform, the anthers linear, dorsifixed, deeply cleft at each end;
ovary 1-carpellate, ovoid, compressed, 1-celled, the style subterminal, shorter
than the ovary, slightly curved; stigma 1, papillose; ovule 1, basifixed, campylo-
tropous; fruit globose, red, juicy; seed lenticular, smooth or minutely scabrous;
embryo annular, the endosperm farinaceous.

A single species, of wide distribution in the tropics of both hemi-
spheres. Walter in Pflanzenreich recognizes three species, but the
characters by which he attempts to separate them are neither con-
stant nor significant.

Rivina humilis L. Sp. PL 121. 1753. R. humilis var. glabra L.
op. cit. 122. R. laevis L. Mant. 41. 1767. R. humilis var. laevis

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 201

Millsp. Field Mus. Bot. 2: 41. 1900. Coxubcanu (PetSn, Maya, fide
Lundell); Chile de raton; Chile (Alta Verapaz); Coralillo; Tomatillo;
Cusucdn (British Honduras, Maya).

Moist or dry thickets and forest, sometimes a weed in cafetales
or other cultivated places, chiefly at low elevations but ascending
to about 1,800 meters; Pete*n; Alta Verapaz; Izabal; Zacapa; Chi-
quimula; Jalapa; Jutiapa; Santa Rosa; Escuintla; Guatemala;
Sacatepequez; Suchitepe'quez ; Retalhuleu; San Marcos; Huehue-
tenango. Southern United States to Mexico, British Honduras,
and Panama; West Indies; South America; Old World tropics.

Plants usually herbaceous and 75 cm. high or less, sometimes becoming woody
and as much as 1.5 meters tall, glabrous or pubescent; leaves slender-petiolate,
ovate to oblong or lanceolate, 3-12 cm. long, 2-6 cm. wide, acute or acuminate,
acute to truncate at the base; racemes slender, 4-20 cm. long, lax, usually many-
flowered, the slender pedicels 3-5 mm. long; sepals green to pink or purple, about
2 mm. long; fruit bright red, 4 mm. in diameter; seed 2.5-3 mm. long.

The plant is rather scarce in most regions of Guatemala where
it is found, but it is rather frequent about Antigua, and particularly
plentiful on the Pacific plains. The brilliantly colored berries are
showy and rather handsome. Their red juice is sometimes used for
dyeing small articles or even as ink. Local names applied to the
plant are "cuxubcan" (Yucatan, Maya), "coral" (Yucatan), and
"achotillo" (Honduras).

STEGNOSPERMA Bentham

An erect or scandent shrub, glabrous; leaves alternate, petiolate, rather
succulent, often coriaceous when dried, without stipules; flowers perfect, in ter-
minal many-flowered racemes, the pedicels bracteate and bracteolate; sepals 5,
herbaceous, with pale membranaceous margins, persistent and somewhat enlarged
in fruit; petals 5, membranaceous, shorter than the sepals, white, deciduous,
imbricate in bud; stamens 10, the filaments subulate, dilated at the base and form-
ing a perigynous annulus, the anthers dorsifixed, sagittate at the base, rounded at
the apex; ovary superior, 3-5-carpellate, the styles as many as the carpels, curved,
papillose within; ovules 1 in each carpel, basifixed; capsule globose, coriaceous,
3-5-angulate, dehiscent from apex to base, 1-5-seeded, the styles persistent; seeds
erect, surrounded by a white or yellowish aril, globose, smooth, black and lustrous;
embryo slightly curved, the cotyledons flattened, equal.

The genus consists of a single species.

Stegnosperma scandens (Lunan) Standl. Field Mus. Bot. 23:
6. 1943. Trichilia scandens Lunan, Hort. Jam. 2: 320. 1814. S.
halimifolium Benth. Bot. Voy. Sulph. 17. pi. 12. 1844.

202 FIELDIANA: BOTANY, VOLUME 24

Dry or moist thickets of the lowlands, 200 meters or less, some-
times on dunes along the coast; Zacapa; Suchitepe"quez ; Retalhuleu;
San Marcos; probably in all the Pacific coast departments. Mexico;
Salvador; Greater Antilles.

An erect shrub 1.5-2 meters tall, or more often scandent over shrubs or trees
and often several meters long; leaves bright green, on petioles 9 mm. long or less,
obovate to elliptic or almost orbicular, 2-7 cm. long, 1-3.5 cm. wide, usually
rounded or obtuse at the apex, obtuse to acuminate at the base; racemes erect,
5-15 cm. long, lax or dense, the pedicels 5-9 mm. long; sepals 5 mm. long and 3 mm.
wide; petals elliptic or oval, 4 mm. long, white; anthers 2 mm. long; capsule sub-
globose, 7-9 mm. long, often tinged with red, the aril also often red; seeds 4 mm.
long.

The shrub apparently has properties similar to those of Phy-
tolacca, for it is reported that in Mexico the roots sometimes are
used as a substitute for soap.

TRIGHOSTIGMA A. Richard

Erect or scandent, glabrous shrubs; leaves alternate, slender-petiolate;
flowers perfect, usually greenish, in lax, terminal and axillary, many-flowered
racemes; bracts deciduous, the bractlets borne near the apex of the pedicel, per-
sistent; sepals 4, concave, spreading or reflexed in fruit; stamens 8-25, the filaments
cylindric-filiform or sometimes very short, the anthers dorsifixed; ovary 1-carpel-
late, 1-celled, the style short, the stigma sessile, penicillate; ovule 1; fruit drupe-
like, subglobose; seed with a crustaceous testa, the embryo annular, the endo-
sperm farinaceous, the cotyledons curved.

Three species are known, one Peruvian, the other, T. polyandrum
(Loes.) H. Walt., with 20-25 stamens, ranging from Nicaragua to
Panama.

Trichostigma octandrum (L.) H. Walt. Pflanzenreich IV. 83:
109. 1909. Rivina octandra L. Cent. PI. 2: 9. 1756. Vittamitta
octandra Hook. f. in Benth. & Hook. Gen. PI. 3: 81. 1880. Ldtigo
(fide Aguilar).

Dry thickets, Escuintla, 900-1,200 meters. Florida; Mexico to
Panama; West Indies; Venezuela to Argentina.

A suberect shrub or a vine as much as 10 meters long; leaves on petioles 1-3.5
cm. long, oblong to elliptic or ovate, 5-15 cm. long, 2-6 cm. wide, acute to acumi-
nate or rarely obtuse, acute to rounded at the base; racemes as long as the leaves
or longer, the pedicels 3-9 mm. long, the lanceolate bracts 2 mm. long; sepals
greenish white, ovate, obtuse, 3.5-4 mm. long, reflexed in age; stamens 8-12;
fruit subglobose, black, 6 mm. in diameter; seed 4-5 mm. long, black, shining.

Only a few collections of this shrub have been made in Guate-
mala, and neither of the authors has collected it. It is to be expected
in all the Pacific coast departments.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 203

AIZOAGEAE

Reference: Percy Wilson, N. Amer. Fl. 21: 267-277. 1932.

Annuals or perennials, usually herbaceous, sometimes suffrutescent, generally
succulent; leaves opposite, alternate, or verticillate, entire; stipules none or
scarious; flowers perfect, polygamo-dioecious, or unisexual, small or large and
showy; calyx usually with 4-5 lobes or sepals; petals none or present and numerous;
stamens few or many, the anthers oblong or linear, 2-celled; disk none or annular;
ovary superior or partly or wholly inferior, 1-many-celled; styles as many as the
ovary cells, the ovules few or many in each cell or sometimes solitary; fruit capsular
and loculicidally dehiscent or circumscissile, or rarely indehiscent and baccate or
nutlike; embryo more or less curved, the cotyledons narrow.

A rather large family, most abundantly represented in Africa.
Only seven genera are represented in North America by native
species, and only the following genera and species are native in
Central America.

Calyx tube partly or wholly adnate to the ovary; petals present.

Mesembryanthemum.
Calyx tube free from the ovary; petals none.

Leaves opposite, very succulent.

Stipules present; ovary 1-2-celled; leaves obovate to suborbicular.

Trianthema.

Stipules none; ovary 3-5-celled; leaves linear or oblanceolate . . . .Sesuvium.
Leaves verticillate, in whorls of 3 or more.

Seeds strophiolate; plants densely pubescent Glinus.

Seeds not strophiolate; plants glabrous or nearly so Mollugo.

GLINUS L.

Plants usually annual, procumbent or ascending, commonly much branched;
leaves mostly verticillate, those of a whorl unequal; flowers perfect, densely
glomerate in the leaf axils, small and inconspicuous; calyx 5-lobate; petals none;
stamens 3-5 or more, the filaments filiform, the anthers small, 2-celled; ovary
3-5-celled, the ovules numerous in each cell; style short, with 3-5 stigmas; capsule
loculicidally 3-5-valvate; seeds numerous, smooth or tuberculate, strophiolate,
borne on a long slender funicle; embryo curved, the cotyledons oblong.

About 10 species in tropical and subtropical regions of both
hemispheres. Only one species is native in North America.

Glinus radiatus (Ruiz & Pavon) Rohrb. in Mart. Fl. Bras. 14,
pt. 2: 238. pi. 55, /. 1. 1872. Mollugo verticillata Ruiz & Pavon, Fl.
Peruv. 1 : 48. 1798.

Damp thickets, or usually on drying or dried mud, at or little
above sea level, Pacific plains; Escuintla; Suchitepe"quez ; Retal-
huleu; probably in all the Pacific coast departments. Western

204 FIELDIANA: BOTANY, VOLUME 24

Texas and Mexico; Honduras; Salvador; Nicaragua; Greater
Antilles; South America.

Plants annual, erect or prostrate and sometimes forming small dense mats,
much branched, the stems mostly 10-30 cm. long, the whole plant grayish or
whitish and densely stellate-tomentose with very slender hairs; leaves small,
verticillate, slender-petiolate, obovate to rounded-spatulate or elliptic, 5-20 mm.
long, acute to broadly rounded at the apex, acute at the base or contracted and
decurrent, entire; flowers in clusters of 3-8; calyx lobes oblong or lanceolate,
2.5-3 mm. long; stamens shorter than the calyx; capsule ellipsoid, 3-3.5 mm. long;
seeds brown or red-brown, numerous, lustrous, smooth.

The plant is seldom found during the dry season but is plentiful
in many localities during the wet months.

MESEMBRYANTHEMUM L.

Succulent annuals or perennials, prostrate or erect, sometimes low shrubs,
very diverse in habit and foliage; leaves usually opposite, 3-angulate, terete, or
flat; flowers white, red, or yellow, mostly terminal, usually opening in sunshine;
calyx 5-parted, the lobes usually foliaceous and unequal; petals very numerous,
linear, in 1 to many rows, united at the base; stamens very numerous, in numerous
series, united at the base; ovary generally 5-celled; fruit a capsule, with 5 to many
cells stellately dehiscent at the apex, becoming somewhat baccate; seeds very
numerous.

A group of 300 or more species, almost all in South Africa. Two
are probably native in California. They are well known in cultiva-
tion because of the bizarre forms of many of the species. In recent
years the genus has been divided into very numerous small ones but
the name is used here in its collective sense.

Mesembryanthemum blandum Haworth, Suppl. PI. Succ.
95. 1819. Una de gato; Portugueses.

Planted abundantly in the highlands of the Occidente, especially
in Quezaltenango and San Marcos, also seen occasionally in the
central departments. Native of South Africa.

A stiff stout branched shrub 60-90 cm. tall, glabrous; leaves very fleshy,
obtusely trigonous, 2-4 cm. long, acute; flowers about 5 cm. broad, rose or rose-
red, the very numerous linear petals usually toothed at the apex.

This is one of the commonest ornamental plants in the Indian
gardens of the dry cold regions of Quezaltenango and especially
San Marcos. The name "una de gato" is said to refer to the reflexed
leaves, but such leaves are not in evidence in the plants we have
observed.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 205

Some other species of Mesembryanthemum are grown as pot or
garden plants in Guatemala, but only sporadically. One is M. cordi-
folium L. (called "siempreviva"), with elongate, sometimes scandent
stems, broadly ovate or cordate leaves, and small, deep rose-red
flowers. Probably M. crystallinum L., the ice-plant of the United
States, also is in cultivation.

MOLLUGO L. Carpet- weed

Slender annuals or perennials, usually much branched, often prostrate,
scarcely succulent; leaves verticillate, narrow or broad, a basal rosette often
present; flowers almost minute, perfect; calyx 5-parted, persistent; petals none;
stamens 3-10; ovary 3-5-celled, superior, the styles 2-5; ovules numerous in each
cell; fruit capsular, membranaceous, 3-5-celled, loculicidally 3-5-valvate; seeds
few or numerous, reticulate, granular, or variously sculptured; embryo curved,
the cotyledons narrowly oblong.

A group of about 15 species, in temperate and tropical regions
of both hemispheres. Nine species are recorded from North America,,
seven of them West Indian, only one in Central America.

Mollugo verticillata L. Sp. PI. 89. 1753.

Cultivated ground, roadsides, moist thickets, or sandbars, chiefly
in the lowlands at or little above sea level, but ascending to about
1,400 meters; Izabal; Zacapa; Chiquimula; Santa Rosa; Escuintla;
Guatemala; Suchitepe*quez ; Retalhuleu; San Marcos. United
States and Mexico to Panama; West Indies and South America;
Old World, where probably introduced.

A slender annual, dichotomously much branched, erect to prostrate; leaves
in whorls of 3-6, obovate to linear, unequal, 1-3.5 cm. long, 1-11 mm. wide,
rounded to subacute at the apex, long-attenuate to the base, short-petiolate,
entire; flowers 2-5 at each node, on pedicels 3-10 mm. long; sepals oblong or
elliptic, 2-2.5 mm. long; stamens usually 3 or 4; capsule ovoid or ellipsoid, 2.5-3
mm. long, 20-30-seeded; seeds dark brown, reniform, 0.6 mm. long, costulate on
the dorsal and lateral surfaces.

An inconspicuous and weedy plant that does not survive long
unless supplied with moisture. Called "clavellina months" in Salva-
dor, and "anisillo" and "culantrillo" in Oaxaca.

SESUVIUM L.

Annual or perennial herbs, usually prostrate and often rooting at the nodes,
very succulent; leaves opposite, the bases often dilated and connate; flowers
axillary, sessile or pedicellate; calyx lobes 5, usually with horn-like dorsal append-
ages below the apex; petals none; stamens 5-many, inserted on the calyx tube, the

206 FIELDIANA: BOTANY, VOLUME 24

filaments filiform; ovary 3-5-celled, the ovules numerous in each cell; styles 3-5,
papillose on the inner side; fruit a membranaceous capsule, 3-5-celled, circum-
scissile; seeds several or many in each cell; embryo annular, the cotyledons oblong.

About 5 species, widely distributed in both hemispheres, espe-
cially in saline or alkaline soil. Five species are listed for North
America, but only one is known from Central America.

Sesuvium Portulacastrum L. Syst. Nat. ed. 10. 1058. 1759.

Salt flats near the seashore, Pacific coast, probably also on the
Atlantic coast; Escuintla; Retalhuleu; doubtless in all the Pacific
coast departments. Southern United States and Mexico to British
Honduras and Panama; West Indies; South America; Old World
tropics.

A glabrous fleshy perennial, the stems branched, often greatly elongate and
rooting at the nodes; leaves oblong to linear or oblanceolate, 2-6 cm. long, 3-15
mm. wide, acute or acutish, clasping at the base; flowers solitary in the leaf axils,
pedicellate; calyx lobes lanceolate, fleshy, 5.5-7 mm. long, cucullate, purplish
within, appendaged dorsally; stamens numerous; styles sometimes distinct;
capsule conic, 9-11 mm. long, 5-6 mm. broad; seeds black, lustrous, smooth,
1.2-1.5 mm. in diameter.

Called "verdolaga de la playa" and "tsaycan" (Maya) in Yuca-
tan. The plant is confined in Central America (and probably else-
where) to the immediate vicinity of mangrove thickets.

Tetragonia expansa L., New Zealand spinach, native of eastern
Asia and New Zealand, is sometimes cultivated in Guatemala as a
pot herb. In general appearance as well as in esculent properties
the plant has much resemblance to the common spinach, Spinacia.
It thrives better in warm climates than does the latter.

TRIANTHEMA L.

Annual or perennial herbs, somewhat succulent, with branching, erect to
prostrate stems; leaves opposite, entire, those of a pair unequal, petiolate, the base
of the petiole sheathing; flowers small, axillary, sessile or pedicellate, solitary
or glomerate; calyx lobes 5, often appendaged dorsally below the apex; petals
none; stamens 5-10 or more numerous, inserted near the top of the calyx tube;
ovary 1-2-celled, few-ovulate, the styles 1-2; capsule membranaceous or coria-
ceous, bearing at apex or on one side a short, fleshy, sometimes lobate appendage,
at length circumscissile; seeds reniform, the embryo annular, the cotyledons oblong.

About 15 species, in the tropics or warmer regions of both hemi-
spheres. A single species occurs in North America.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 207

Trianthema Portulacastrum L. Sp. PI. 223. 1753. Verdolaga.

Moist or dry fields or thickets, often on salt flats along the sea-
shore, chiefly in the lowlands but ascending to 1,200 meters; Zacapa;
Jutiapa; Escuintla; Guatemala (Lago de Amatitlan) ; Suchitepe"quez;
Retalhuleu; San Marcos; doubtless in all the Pacific coast depart-
ments. Southern United States and Mexico to British Honduras
and Panama; West Indies; South America; Old World tropics.

A succulent annual, erect to prostrate, often tinged with red or purple, the
branches a meter long or usually much shorter; leaves petiolate, obovate to sub-
orbicular or elliptic, 1-4 cm. long, rounded and apiculate or emarginate at the
apex, usually acute at the base; flowers partly concealed by the sheathing petioles,
axillary; sepals ovate-lanceolate or lanceolate, 4-5 mm. long, pinkish or purple
within; capsule 4-5 mm. long, aristate; seeds rough, black, 2 mm. in diameter.

By some authors the plant is described as perennial, but usually
if not always it is an annual and often a short-lived one. In Guate-
mala, except on salt flats, it soon withers after the end of the rainy
season.

PORTULACACEAE. Purslane Family

Reference: Per Axel Rydberg, Portulacaceae, N. Amer. Fl. 21:
279-336. 1932.

Annual or perennial herbs, rarely shrubs, usually very succulent, glabrous, or
rarely pilose at the nodes; leaves opposite, alternate, or in basal rosettes, entire;
stipules scarious, lacerate or modified into hairs, sometimes none; flowers small,
solitary, racemose, paniculate, or cymose, terminal or axillary, perfect, regular or
nearly so; -sepals generally 2, persistent or deciduous, scarious or herbaceous;
petals mostly 4-5, sometimes slightly united at the base, often fugacious or marces-
cent; stamens inserted with the petals, sometimes adnate at the base, usually as
many as the petals, sometimes more or fewer; filaments filiform, the anthers
2-celled, longitudinally dehiscent; ovary 1-celled, superior or partly or wholly
inferior, the styles 2-7, more or less united; ovules 2-many, the placenta central
or basal; fruit capsular, loculicidally dehiscent or circumscissile, the valves as
many as the styles; seeds 3-many, or by abortion 1-2, usually rounded-reniform,
compressed, lenticular, sometimes strop hiolate, the testa often crustaceous;
embryo generally hippocrepiform, enclosing the farinaceous endosperm.

Perhaps 15 genera, mostly in America but some of them repre-
sented in the Old World. The species are most numerous, at least
in most of the groups, in temperate and arctic North America. Only
the following genera are represented in Central America. No two
authors seem to be in accord as to the genera that are to be recognized
in North America. Rydberg recognizes 17 in North America, but
most authors unite some of these, although they are not in agree-
ment as to how they are to be combined.

208 FIELDIANA: BOTANY, VOLUME 24

Ovary partly or wholly inferior; leaves flat or terete; capsule circumscissile at or

above the middle Portulaca.

Ovary superior; leaves flat; capsule valvate, or rarely circumscissile at the base.
Sepals deciduous; stems with numerous leaves; flowers in racemes, panicles, or

cymes Talinum.

Sepals persistent; stems naked or leafy, or the plants sometimes acaulescent;

flowers solitary in the leaf axils or in terminal umbels.
Leaves all basal, the pedicels arising among the leaves; plants with a thick

fleshy perennial root Oreobroma.

Leaves partly cauline, or the plants with elongate naked stems and a terminal

umbel or short raceme of flowers.
Stems very leafy, the leaves alternate; flowers solitary in the leaf axils.

Calandrinia.

Stems naked, or with several pairs of opposite leaves; flowers solitary in the
leaf axils or in terminal umbels . . . . Montia.

CALANDRINIA L.

Glabrous annuals with usually elongate stems; leaves numerous, alternate,
fleshy; flowers small, white, pink, or pale blue, pedicellate, axillary; sepals 2,
herbaceous, usually persistent; petals generally 5, ephemeral; stamens 5-12, the
filaments free or united at the base in a ring, or adherent at the base to the petals;
ovary superior, many-ovulate, the styles 3, united below; capsule globose or ovoid,
membranous or chartaceous, 3-valvate; seeds lenticular, rounded-reniform, con-
centrically lineate, sometimes muricate, strophiolate or naked at the hilum;
embryo hippocrepiform.

About 100 species, in America and Australia, mostly in South
America. Only the following is known from Central America.

Calandrinia micrantha Schlecht. Ind. Sem. Hort. Hal. 1838;
Linnaea 13: Litt.-Ber. 97. 1839. Berros; Barba de San Nicolas;
Excacahue (Quezaltenango).

Open banks or fields, often on limestone or in sand, sometimes
in Alnus forest, often an abundant weed in cultivated fields, 1,800-
3,700 meters; Chimaltenango; Totonicapan; Huehuetenango; Que-
zaltenango; San Marcos. Mexico.

A succulent annual, much branched from the base, the stems prostrate,
densely leafy, 7-30 cm. long; leaves lanceolate or linear-oblanceolate, the lowest
3-5 cm. long, petiolate, acute, attenuate to the base, the upper leaves shorter,
sessile, obscurely ciliate; pedicels 2-5 mm. long, shorter than the calyx; sepals
ovate, acute, 6 mm. long, costate, ciliate on the margins and costa; petals pale
blue, equaling the calyx; stamens 3-6; capsule oblong, almost equaling the calyx;
seeds black, lustrous, 1.5 mm. long.

A rather common plant in the mountains of the Occidente. It
is cooked and eaten like spinach, the whole plant being used, and
it is said to be one of the best of the wild pot herbs. Compact

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 209

bunches of the plants may be found in the markets of Totonicapan,
Quezaltenango, and elsewhere. This species is closely related to
C. caulescens HBK. of the South American Andes and it is somewhat
questionable whether it is really distinct from that species.

MONTIA L.

Annual or perennial, more or less succulent herbs, glabrous, with few or
numerous basal leaves; cauline leaves 2 and opposite, at the apex of the stem,
often completely united by their bases, or the cauline leaves 2 or more pairs and
often distinct; flowers small, white or pink, axillary or in terminal umbels or short
racemes; sepals 2, persistent, often unequal; petals 5, emarginate; stamens 5;
ovary subglobose, 3-ovulate; styles 3; capsule subglobose, 3-valvate from the
apex, the valves elastically involute at dehiscence; seeds 1-3, lenticular, usually
smooth and shining, with a minute strophiole.

Species perhaps 50, mostly American but a few in the Old World.
Only the following are known from Central America.

Flowers axillary; cauline leaves several pairs, distinct, oblanceolate . . .M. calcicola.

Flowers terminal, in umbelliform racemes; cauline leaves 1 pair, united to form

a rounded perfoliate disk M. mexicana.

Montia calcicola Standl. & Steyerm. Field Mus. Bot. 23: 48.
1944.

On limestone cliffs or rocks, in or near Juniperus forest, 3,700-
3,800 meters; endemic; Huehuetenango (Sierra de los Cuchumatanes;
type from Cerro Che"mal, Steyermark 50308); San Marcos(?).

Perennial, with very slender, elongate, sparsely leafy stolons, glabrous through-
out, the stems mostly 10 cm. long or less, prostrate or procumbent, simple; leaves
cauline, 5 or fewer pairs, opposite, mostly 2-2.5 cm. long (including the petiole),
oblanceolate, 3-5 mm. wide, obtuse or subacute, attenuate at the base into a
marginate petiole; flowers 1-3 in the leaf axils, the pedicels 8 mm. long or shorter,
recurved in age; sepals pale green, rounded-obovate, 1.5 mm. long or slightly
larger, rounded at the apex, shorter than the capsule; petals pale pink, somewhat
longer than the sepals; capsule subglobose, 2 mm. long, 3-valvate; seeds 2-3,
reddish black, 1 mm. in diameter, very minutely and closely reticulate.

Montia mexicana (Rydb.) Standl. & Steyerm. Field Mus. Bot.
23: 49. 1944. Limnia mexicana Rydb. N. Amer. Fl. 21: 309. 1932.

Alpine meadows or on limestone rocks in Juniperus forest,
2,600-3,800 meters; Huehuetenango. Southern Mexico.

Plants annual, variable in size and habit, 4-20 cm. high; basal leaves few or
numerous, on petioles 3-15 cm. long, very thin when dried but succulent when
fresh, the blades rhombic or broadly deltoid, often broader than long, 1-5 cm.

210 FIELDIANA: BOTANY, VOLUME 24

wide, abruptly contracted at the apex into a distinct triangular cusp, abruptly
contracted and decurrent at the base; cauline leaves 1 pair, connate to form an
orbicular perfoliate disk or cup 1.5-5 cm. wide; inflorescence subsessile, several-
flowered, umbelliform, the pedicels 1 cm. long or shorter; sepals rounded-obovate,
2 mm. long; petals white, spatulate, 3 mm. long; seeds black, smooth, lustrous,
1.5 mm. long.

This has been reported from Guatemala as Claytonia perfoliata
Bonn (Montia perfoliata Howell), and it is closely related to that
common plant of the Pacific coast of the United States, whose
succulent leaves have been much eaten in the past in salads. The
several Guatemalan collections are highly variable in size and shape
of their leaves but not more so than those of the Californian plant,
to which it is suspected M. mexicana ultimately will have to be
reduced.

OREOBROMA Howell

Perennial herbs with a fleshy taproot and a short cespitose caudex; basal
leaves numerous, densely clustered at the end of the caudex; stems or scapes
geniculate at the base, the flowers racemose, cymose, or paniculate or (as in the
Central American species) reduced to a single flower; sepals 2, persistent, some-
times dentate; petals 5-10, white or pink; stamens 5-20, the filaments filiform;
ovary ovoid, the ovules numerous, the placenta central; styles 3-7, united at the
base; capsule ovoid, circumscissile near the base, then splitting upward, several-
seeded; seeds ovate or rounded, smooth and lustrous, estrophiolate.

Species about 20, all American and chiefly in the western United
States. Only one is known in Central America.

Oreobroma megarhizum (Hemsl.) Standl. & Steyerm. Field
Mus. Bot. 23: 49. 1944. Calandrinia megarhiza Hemsl. Diag. PI.
Mex. 23. 1879. Claytonia megarhiza Kuntze, Rev. Gen. 57. 1891.
0. mexicanum Rydb. N. Amer. Fl. 21: 326. 1932.

Type from Volcan de Fuego, Sacatepe"quez, 3,300-3,600 meters,
Salvin; also in Huehuetenango (Sierra de los Cuchumatanes, 3,700
meters). High mountains of central Mexico.

Plants perennial, with a thick fleshy taproot as much as 10 cm. long and 1.5
cm. thick; leaves numerous, all basal, linear, fleshy, usually flat on the ground,
2-7 cm. long, 2-3 mm. wide, somewhat dilated and nerved at the base; flowers
numerous but solitary on basal peduncles arising among the leaves, the peduncles
5-20 mm. long; sepals elliptic-lanceolate, 6-7 mm. long; petals 5-6, spatulate,
white, 1 cm. long; capsule ellipsoid, 7 mm. long, the pericarp very thin, circum-
scissile at the base; seeds numerous, black, 1.5 mm. long.

We have seen no material from Volcan de Fuego, but have little
or no doubt that the Mexican and Guatemalan plants are conspecific.
The plant is a truly alpine species.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 211

PORTULACA L.

Reference: Karl von Poellnitz, Versuch einer Monographic der
Gattung Portulaca L., Repert. Sp. Nov. 37: 240-320. 1934.

Succulent, annual or perennial herbs, often prostrate, glabrous or pubescent;
leaves alternate or opposite, flat or terete, often verticillate about the flowers;
stipules scarious or none, often reduced to tufts of hairs; flowers perfect, solitary
or crowded at the ends of the stems, small or large; sepals 2; petals 4-6, usually 5;
stamens 8-many, inserted at the base of the petals; ovary partly or wholly inferior,
the styles 3-9; ovules numerous; capsule 1-celled, membranous, circumscissile,
many-seeded; seeds reniform or cochleate, the testa smooth or minutely tubercu-
late.

About 100 species, mostly in the Old World but more than 20
are found in North America. Only the following are known from
Central America.

Plants glabrous or nearly so, not white-pilose about the flowers and in the leaf
axils; leaves flat; petals yellow; sepals carinate P. oleracea.

Plants densely white-pilose with long soft hairs about the flowers and in the leaf
axils; leaves subterete or flat; petals yellow or rose-purple; sepals not carinate.

Petals yellow; leaves flat P. Conzattii.

Petals rose-purple; leaves terete or nearly so P. pilosa.

Portulaca Gonzattii P. Wilson, Torreya 28: 28. 1928.

Collected in Guatemala but once, in depressions on top of boulder
along a stream in a quebrada, about 1,350 meters; Jalapa (near
Jalapa, Standley 77421). Southern Mexico, the type from Oaxaca.

Plants erect, slender or stout, probably perennial, 30 cm. tall or often much
lower, sparsely branched, the stems often very stout and fleshy, bearing dense
tufts of very long, white hairs in the leaf axils; leaves alternate, flat, lanceolate to
obovate or oblanceolate, 1.5-2.5 cm. long, 2.5-4.5 mm. wide, obtuse or subacute;
flowers terminal, in fascicles of 2-3, surrounded by very long, white hairs and by
an involucre of 8 or more large leaves; sepals deltoid-orbicular, 5 mm. long; petals
yellow, obovate or elliptic-obovate, 7-8 mm. long; stamens about 20; lobes of the
style 4-5; capsule subglobose, 4 mm. in diameter, circumscissile at the middle;
seeds black, 0.8 mm. in diameter, obtusely tuberculate.

Portulaca oleracea L. Sp. PI. 445. 1753. Verdolaga; Paxlac
(Quiche") ; Graviol (Quecchi).

Moist fields or cultivated or waste ground, often along roadsides,
on open banks, or in city streets, 2,400 meters or less; Alta Verapaz;
Izabal; Zacapa; Chiquimula; Jalapa; Santa Rosa; Escuintla; Guate-
mala; Sacatepe"quez ; Chimaltenango; Suchitepe"quez ; Retalhuleu;
Quezaltenango; San Marcos; Huehuetenango; probably in all the
departments. Temperate North America; Mexico; British Hon-

212 FIELDIANA: BOTANY, VOLUME 24

duras to Salvador and Panama; West Indies; South America;
temperate and tropical regions of the Old World.

A glabrous fleshy annual, usually much branched from the base, the stems
prostrate and forming mats, mostly 20-40 cm. long, sometimes ascending, often
reddish; leaves alternate, cuneate-obovate or spatulate, 1-3 cm. long, rounded or
almost truncate at the apex, attenuate to the sessile base; flowers sessile, clustered
or solitary at the ends of the stems, the hairs surrounding them very inconspicuous
or wanting; sepals broadly ovate or orbicular, 3-4.5 mm. long, subacute, carinate;
petals yellow, 3-4.5 mm. long; stamens 6-10; style 4-6-lobate; capsule 5-9 mm.
high, circumscissile at about the middle; seeds black, almost 1 mm. in diameter,
granulate.

The Maya name of Yucatan and probably also of Guatemala is
"xucul." This is one of the most widely distributed weedy plants,
being found almost all over the earth. Probably it was native
originally in one or the other of the hemispheres, and has been
introduced into the other, but if so, it is not known which is the
original home. "Purslane," "pusley," or "pursley," as it is called
in English, is found commonly in most parts of Guatemala, at least
where there are settlements or cultivated ground. The plants
produce great numbers of seeds, and their stems show great vitality.
When pulled from the soil and placed upon some place where they
can not take root, they require weeks for withering. The plant is
of considerable economic importance in all Central America since
the young stems and leaves are much eaten as a pot herb, like
spinach, for which they afford an excellent substitute, and they are
also good when used raw in salads. They are so used occasionally
in some parts of the United States but much less commonly than in
Guatemala, where they are one of the common verduras of the
markets. The plants make excellent food for pigs and other stock.

Portulaca pilosa L. Sp. PI. 445. 1753. Colchon de nino; Anisillo
(Zacapa).

Moist or dry, often rocky plains or hillsides, often in sand, 2,500
meters or less, most frequent at low elevations; Zacapa; Chiquimula;
Jalapa; Jutiapa; Escuintla; Retalhuleu; Solola; Quezaltenango;
San Marcos. Southern United States; Mexico; British Honduras
to Salvador and Panama; West Indies; South America.

Plants very succulent, annual or sometimes persisting for more than a year,
prostrate to erect, usually branched, the stems mostly 15 cm. long or less, densely
white-pilose in the leaf axils; leaves alternate, terete, 5-16 mm. long, sessile or
nearly so; flowers clustered at the ends of the branches, surrounded by long, whitish
or brownish hairs and an involucre of 6-10 leaves; sepals not carinate, triangular-
ovate or ovate, 2-3 mm. long; petals rose-purple, obovate or broadly obovate,

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 213

3-5.5 mm. long, sometimes retuse; stamens 15-32, the filaments crimson; style
4-6-lobate; capsule subglobose, 3-4 mm. in diameter, circumscissile at about the
middle; seeds black, 0.5 mm. broad, minutely tuberculate.

Sometimes called "hiedra" in Honduras; "arroz-xiu" (Yucatan,
Spanish and Maya); "tsayoch," "tsotsiltsaioch" (Yucatan, Maya).
The plant often is grown for ornament in Guatemalan gardens and
sometimes is employed to make formal designs in flower beds in
gardens or parks.

Portulaca grandiflora Hook., native of Argentina, probably is in
cultivation for ornament in Guatemala, although we have no record
of its occurrence there. It has large flowers, the petals 1.5-2.5 cm.
long, and pink, red, yellow, orange, or white. The flowers are open
in the early morning, but the petals, as in some or all other species,
collapse about or before noon.

TALINUM Adanson

Herbs or low shrubs, annual or perennial, the stems short or elongate; leaves
carnose, alternate or subopposite, flat or terete, entire; flowers small or rather
large, in cymes, these on long or short peduncles, often paniculate, or the flowers
sometimes solitary in the leaf axils; sepals 2, deciduous; petals 5 or more, soon
withering; stamens few or numerous, the filaments filiform; ovary superior, the 3
styles more or less united, the ovules numerous; capsule 1-celled, 3-valvate;
seeds compressed, rounded-reniform; embryo incompletely annular; endosperm
farinaceous.

About 35 species, mostly in temperate and tropical North
America, a few in Africa and Asia. Only the following are found in
Central America.

Plants with annual stems; leaves mostly 2-4 cm. wide; flowers in elongate panicles;

sepals 3-4 mm. long T. paniculatum.

Plants with perennial stems, somewhat suffrutescent below; leaves mostly less

than 2 cm. wide; flowers few, racemose or cymose; sepals 5-6 mm. long.

T. triangulare.

Talinum paniculatum (Jacq.) Gaertn. Fruct. & Sem. 2: 219.
pi. 128, f. 13. 1791. Portulaca paniculata Jacq. Enum. PI. Carib. 22.
1760. P. patens L. Mant. PL 242. 1771. T. patens Willd. Sp. PL 2:
863. 1800. Verdolaga; Orejilla.

Moist or wet fields or thickets, often in waste ground, sometimes
in cultivated fields, 2,400 meters or less, mostly at low elevations;
Pete*n; Baja Verapaz; Zacapa; Chiquimula; Santa Rosa; Escuintla;
Guatemala; Quiche"; Huehuetenango. Southern United States;
Mexico; southward to Panama; West Indies; South America.

214 FIELDIANA: BOTANY, VOLUME 24

Plants erect from somewhat fleshy or tuberous roots, the stout stems simple
or branched, a meter high or less, the plants glabrous throughout; leaves fleshy,
elliptic or obovate, 3-10 cm. long and 1.5-4.5 cm. wide or sometimes larger, obtuse
or acute, attenuate to the base and sessile or nearly so; inflorescence a rather open
but narrow, many-flowered, terminal panicle 10-40 cm. long, the flowers in open
cymes, yellow; pedicels slender, 1-2 cm. long, terete; sepals oval or orbicular, 3-4
mm. long; petals oval or orbicular, 3.5-5 mm. long; stamens 15-20; capsule sub-
globose, 3-4.5 mm. in diameter; seeds black, 1 mm. in diameter, striolate and some-
times minutely tuberculate, lustrous.

Called "lechuguilla" in Salvador; "saioch," "dzum-yail" (Yuca-
tan, Maya). Undoubtedly the leaves of this species could be used
like those of T. triangular e and make a good substitute for spinach,
but we have no information that they are so employed in Central
America. The growing plants are found only during the rainy
season, the stems withering when the rains cease.

Talinum triangulate (Jacq.) Willd. Sp. PL 2: 862. 1800.
Portulaca triangularis Jacq. Enum. PI. Carib. 22. 1760.

Moist or rather dry, often rocky thickets or low forest, 650
meters or less; Zacapa; Chiquimula; Suchitepe"quez. Southern
Florida; Mexico; Honduras; West Indies; South America.

Plants perennial from a stout, often woody root, usually 50 cm. high or often
much lower, much branched, the stems very thick and fleshy, persisting for several
years; leaves usually deciduous during the dry season, oblanceolate or obovate,
mostly 2-5 cm. long and 1.5 cm. wide or less, usually rounded at the apex, attenu-
ate to the base, more or less petiolate, thick and succulent; inflorescence a few-
many-flowered raceme or a small cyme, the pedicels 7-11 mm. long, 3-angulate;
sepals lance-ovate or broadly ovate, 5-7 mm. long, cuspidate, somewhat persistent;
petals white, broadly elliptic or oval, sometimes pink or purple, 7-10 mm. long;
stamens about 30; capsule subglobose, 4.5-6 mm. in diameter; seeds black, lustrous,
almost 1 mm. in diameter, minutely striolate.

Sometimes known as Philippine spinach, and called "espinaca"
in Honduras. The fleshy leaves make an excellent substitute for
spinach and the plant has been introduced into many remote regions
as a garden plant for this reason. It is said to be much grown in the
Philippines and the East Indies, and some years ago was introduced
from the Philippines into the Atlantic coast of Honduras by persons
who did not know that it was actually native in the latter area.
The stems of this species do not die to the ground during the dry
season as do those of T. paniculatum.

BASELLACEAE

Reference: Percy Wilson, Basellaceae, N. Amer. Fl. 21: 337-339.
1932.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 215

Herbaceous vines, usually with tuberous roots, glabrous, succulent; leaves
alternate, without stipules, entire, often cordate; flowers small, perfect, in simple
or branched racemes or spikes, regular, a bract present at the base of the pedi-
cel and 2 bractlets at its apex; sepals 2, sometimes winged in fruit; petals 5;
stamens 5, inserted on a hypogynous disk adnate to the base of the corolla, opposite
the petals; filaments terete or complanate, sometimes reflexed in bud; ovary
superior, 1-celled, 1-ovulate; styles 1-3, the stigmas entire or cleft; fruit included
in the perianth, utricular; seed erect, with endosperm.

Five small genera, four in tropical America, one in Asia. Only
the following are known in North America. Ullucus tuberosus
Lozano is an important food plant of the South American Andes,
cultivated on a large scale for its potato-like tubers, which are
cooked and eaten. Basella rubra L., native of tropical Asia, some-
times is cultivated for its succulent leaves, which are cooked and
eaten like spinach.

Sepals broadly winged Anredera.

Sepals not winged , Boussingaultia.

ANREDERA Jussieu

Roots tuberous; leaves petiolate, fleshy; flowers small, white, in dense peduncu-
late curving racemes; bracts lance-subulate, deciduous, the bractlets triangular,
persistent; sepals navicular, enclosing the petals, broadly winged dorsally; petals
hyaline, subequal, oblong, obtuse, 1-nerved; filaments subulate, in bud reflexed
below the apex, the anthers sagittate, included; styles 3, connate at the base, the
stigmas dilated, 2-3-lobate; utricle included in the perianth, stipitate, ovoid-
globose, the pericarp fleshy, adherent to the seed; seed erect, compressed, the
testa coriaceous, brown; embryo almost annular, the endosperm scant, farinaceous.

The genus consists of a single species.

Anredera vesicaria (Lam.) Gaertn. f. in Gaertn. Fruct. 3: 176.
1807. Polygonum scandens L. Sp. PI. 364. 1753, in part. Basella
vesicaria Lam. Encycl. 1: 382. 1785. A. scandens Moq. in DC.
Prodr. 13, pt. 2: 230. 1849. Hiedra de monte.

Moist thickets or hedges, 400-1,300 meters; Chiquimula;
Jutiapa; Huehuetenango. Texas and Mexico; Panama; Cuba and
Jamaica; western South America.

A small or large vine, much branched, climbing over shrubs; leaves broadly
ovate to oblong-ovate, 3-6.5 cm. long, 2-4 cm. wide, acute or acuminate, abruptly
narrowed at the base or truncate; racemes very dense, 2-12 cm. long, 1 cm. broad
in fruit, the flowers short-pedicellate, whitish, somewhat fragrant; wings of the
sepals in fruit 4-5 mm. long, membranous; petals 2 mm. long.

Called "suelda con suelda" in Salvador. An infusion of the
succulent leaves is said to be used in Huehuetenango for shampooing
the hair.

216 FIELDIANA: BOTANY, VOLUME 24

BOUSSINGAULTIA HBK.

Slender vines with much branched stems; leaves succulent, on long or short
petioles, broad; flowers in axillary or terminal racemes or panicles; sepals 2, some-
what shorter than the petals; stamens inserted at the base of the petals, the fila-
ments subulate or lanceolate, recurved in bud, the anthers versatile; ovary ovoid,
with 3 stigmas, or the stigma simple and 3-lobate; ovule subsessile; utricle included
in the persistent perianth; seed erect, the testa crustaceous; embryo semiannular,
the cotyledons plano-convex, the radicle thick.

About 10 species, in tropical America. Only the following are
known in North America.

Styles united throughout, with a 3-lobate stigma. Flowers less than 3 mm. broad,

dark purple when dried B. ramosa.

Styles connate only at or near the base, the stigmas 3.

Flowers 5-6 mm. broad, dark purple or blackish when dried; stigmas entire.

B. baselloides.
Flowers 3-3.5 mm. broad, white when dried; stigmas usually 2-cleft.

B. leptostachys.

Boussingaultia baselloides HBK. Nov. Gen. & Sp. 7: 196.
1825. Hiedra.

Often planted for ornament, and sometimes naturalized in hedges
and thickets, 1,300-2,300 meters; Alta Verapaz; Jalapa; Quezal-
tenango. Mexico; South America.

Stems rather stout and thick; leaves broadly ovate to orbicular-ovate, 3-10
cm. long, acute to short-acuminate, very succulent, deeply cordate to abruptly
cuneate at the base; racemes slender or stout, simple or compound, 5-20 cm. long,
the pedicels 2 mm. long; flowers 5-6 mm. wide, white at first, turning dark purple;
bractlets connate and persistent; sepals suborbicular, 2-2.5 mm. long, the petals
elliptic or oval; filaments lanceolate; stigmas stout, entire; fruit globose, 1 mm.
in diameter, brown.

The vine grows plentifully in the hedge at the cathedral in
Quezaltenango. Called "Madeira vine" in the United States, where
it is often grown for ornament. The plant is said to be a native of
Ecuador, and it probably is not native anywhere in North America.
Produced along the stems are many fleshy aerial tubers somewhat
suggestive of tiny potatoes, by which the plant may be propagated
readily. The flowers are fragrant.

Boussingaultia leptostachys Moq. in DC. Prodr. 13, pt. 2:
229. 1849.

Moist hedges or thickets, 1,200 meters or less; Zacapa; Chiqui-
mula; Santa Rosa. Southern Florida; Mexico; British Honduras;
West Indies; South America.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 217

A small or large vine, sometimes covering rather large trees; leaves petiolate,
ovate to ovate-elliptic or rounded-ovate, 2-8 cm. long, acute or acuminate, gradu-
ally or abruptly narrowed at the base; racemes slender and lax, 6-20 cm. long, the
pedicels 1 mm. long, the flowers white; bractlets free, often deciduous; sepals 1.3-
1.6 mm. long; petals 2 mm. long; filaments subulate; styles connate near the base,
the stigmas slender, mostly 2-cleft.

The name "xayillol" is reported from Yucatan.

Boussingaultia rarnosa (Moq.) Hemsl. Biol. Centr. Amer.
Bot. 3: 27. 1882. Tandonia ramosa Moq. in DC. Prodr. 13, pt. 2:
227. 1849. Dioscorea calyculata Bonn. Smith, Bot. Gaz. 20: 295.
1895 (type from Guachipelin, Guatemala, Heyde & Lux 6260).
Llovizna.

Moist thickets, 750-2,000 meters; type collected somewhere in
Guatemala by Skinner; Guatemala; Sacatepe"quez; Quiche"; Hue-
huetenango. Southern Mexico; Costa Rica.

Plants sometimes climbing over tall trees, the stems then pendent from the
high branches; leaves petiolate, ovate to rounded-ovate, 2-7 cm. long, acute or
acuminate, subcordate or truncate at the base; racemes slender and lax, 4-15 cm.
long, the flowers white, turning dark purple, the pedicels 1-2 mm. long; bractlets
ovate to triangular-ovate, persistent; sepals oval, 1.4 mm. long; petals 2 mm. long;
filaments subulate; styles united throughout, the stigma 3-lobate.

CARYOPHYLLACEAE. Carnation Family

Mostly annual or perennial herbs, the stems often articulate at the nodes
and usually more or less thickened; leaves opposite, entire, mostly 1-3-nerved,
often connate at the base by a transverse line, with or without stipules, the
stipules, when present, small and scarious; flowers small or large, white or colored,
perfect or rarely by abortion unisexual; inflorescences centrifugal, cymose and
many-flowered or simple or dichotomous; flowers regular, the sepals 4-5, persistent,
free or connate, imbricate; petals as many as the sepals, inserted on a hypogynous
annulus or sometimes short-perigynous, entire, 2-fid, or lacerate, imbricate and
usually contorted, sometimes minute or none; stamens 8-10 or fewer, inserted
with the petals; filaments filiform, the anthers 2-celled, the cells parallel, longitu-
dinally dehiscent; torus usually small; ovary free, 1-celled or rarely partially
2-5-celled; styles 2-5, stigmatose on the inner side, free or connate below; ovules
2-many, the funicles arising from the base of the ovary or affixed to a central
column, amphitropous, ascending; fruit capsular, membranaceous or crustaceous,
opening by as many or twice as many valves or teeth as there are styles; seeds
numerous or few, the testa membranaceous or crustaceous, various in shape, the
hilum marginal; endosperm farinaceous or rarely carnose; embryo more or less
curved, the radicle terete; seeds smooth or often granulate or echinate, rarely
winged.

Genera about 80, chiefly in temperate and cold regions, in the
tropics found mostly in the mountains. One other genus, Poly-
carpaea, is represented in Central America (Panama).

218 FIELDIANA: BOTANY, VOLUME 24

Calyx of united petals, dentate or lobate; petals unguiculate. Stipules none.
Calyx multistriate, subtended at the base by bracts. Cultivated plants.

Dianthus.
Calyx 5-10-nerved, not bracteate at the base.

Calyx 5-nerved; styles 2. Plants glabrous; flowers small, white . . . Gypsophila.
Calyx with 10 or more nerves; styles 3-4. Plants glabrous or pubescent;

flowers often large, variously colored Silene.

Calyx of distinct sepals; petals not unguiculate; stipules sometimes present.
Styles united below, with usually 3 branches above. Stamens 5; petals 2-parted;

small scarious stipules present but sometimes deciduous Drymaria.

Styles free, with 2-5 branches.

Stipules present, scarious, mostly small and inconspicuous; leaves linear,

apparently in whorls of very numerous leaves Spergula.

Stipules none; leaves not verticillate.

Capsule cylindric, usually somewhat curved, opening by usually 10 minute

teeth. Petals 2-cleft Cerastium.

Capsule ovoid, not cylindric, opening by 5 or fewer valves.

Petals 2-cleft or 2-parted Stellaria.

Petals entire or shallowly emarginate, sometimes none.

Styles as many as the sepals; plants small, annual, the stems mostly

2-5 cm. long Sagina.

Styles fewer than the sepals; plants usually much larger and with
elongate stems, or the stems sometimes short and cespitose but
the plants then perennial Arenaria.

ARENARIA L.

Reference: F. N. Williams, A revision of the genus Arenaria,
Journ. Linn. Soc. Bot. 33: 326-437. 1898.

Annual, biennial, or perennial herbs, rarely suffrutescent, often cespitose;
leaves subulate and stiff or broad but small and membranaceous; inflorescence
usually dichasiiform, the flowers terminal and cymose-paniculate, thyrsoid,
capitate, or solitary, sometimes axillary and solitary, the petals generally white;
sepals 5, connate at the very base; petals 5, entire, rounded to obtuse, retuse, or
emarginate at the apex, rarely erose or laciniate, sometimes none; stamens 10,
rarely 5; disk perigynous, bearing the stamens, sometimes annular, sometimes
5- or 10-lobate, often glanduliferous; ovary 1-celled, the styles 3 or 2, distinct; cap-
sule globose, ovoid, short-oblong, or rarely cylindric-conic, sometimes depressed,
dehiscent by twice as many teeth as the number of the styles, usually split finally
into 3-2 bidentate valves; seeds estrophiolate, naked, reniform-globose or laterally
compressed, tuberculate, scabrous, or smooth.

Species about 170, widely distributed in both hemispheres,
mostly in temperate or cold regions, in the tropics almost wholly
confined to mountain regions. Only the following species have been
found in Central America.

Plants cespitose, forming very dense, rounded tufts or mats; leaves densely imbri-
cate, 4 mm. long or less, rounded at the apex A. bryoides.

Plants with elongate branched stems; leaves not imbricate, mostly much more
than 5 mm. long, sometimes acute or attenuate.

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 219

Pedicels glabrous A. paludicola.

Pedicels puberulent or short-pilose.

Leaves linear or subulate; sepals glabrous, sometimes ciliate.

Leaves aristate-acuminate, with a very thick and prominent costa, stiff
and rather rigid; costa of the sepals thick and conspicuous.

A. lycopodioides.

Leaves subacute, not aristate, the costa inconspicuous, the blades herba-
ceous, soft; costa of the sepals inconspicuous A. altorum.

Leaves lanceolate to ovate; sepals pubescent, at least on the costa.

Petals equaling or usually shorter than the sepals, often none; sepals 2-3

mm. long.
Petals none; leaves abruptly contracted at the base and conspicuously

petiolate A. reptans.

Petals present; leaves sessile or gradually narrowed into a short petiole.

A. lanuginosa.
Petals conspicuously longer than the sepals; sepals 4-5 mm. long.

Leaves lanceolate or elliptic-lanceolate, puberulent, short-ciliate; stems
short-pubescent with subreflexed hairs; leaf margins not thickened;

pedicels puberulent A. guatemalensis.

Leaves ovate, pilose with rather long, spreading hairs, long-ciliate; stems
pilose with soft spreading hairs; leaf margins conspicuously thick-
ened; pedicels pilose with rather long, spreading hairs.

A. megalantha.

Arenaria altorum Standl. & Steyerm. Field Mus. Bot. 23: 49.
1944. Clarincillo.

Dry, open, often rocky mountain slopes, 1,500-2,900 meters;
endemic; Jalapa (type collected near Minas de Croma, Potrero
Carrillo, 13 miles northeast of Jalapa, Steyermark 33091; flowering
in December); Huehuetenango (Sierra de los Cuchumatanes).

Perennial, ascending, the roots thick and lignescent, the stems several, slender,
6-14 cm. long, simple or sparsely branched above, minutely puberulent; leaves
sparse, linear, spreading, slightly fleshy, sessile, 10-18 mm. long, scarcely 1 mm.
wide, glabrous, ciliate near the base; flowers axillary or subpaniculate, often also
terminal, few or numerous, the pedicels straight, 8 mm. long or less, very minutely
puberulent, erect or suberect; sepals 3-3.5 mm. long, glabrous or microscopically
puberulent on the keel, acute or subulate-acuminate, the apex subrecurved, con-
spicuously carinate, green along and near the keel, the margins scarious, white;
petals entire, slightly longer than the sepals; styles 3; capsule 4 mm. long, lustrous,
shortly 3-valvate, the valves emarginate.

A relative of the Mexican A. Bourgaei Hemsl. which might well
occur in western Guatemala but apparently has not been collected
there thus far.

Arenaria bryoides Willd. ex Schlecht. Ges. Naturf. Freund.
Berlin Mag. 7: 201. 1813. A. bryoides var. guatemalensis Hemsl.

220 FIELDIANA: BOTANY, VOLUME 24

Biol. Centr. Amer. Bot. 1: 70. 1879 (type from summit of Volcan de
Fuego, Salvin & Godman 224).

Open rocky alpine slopes or summits, sometimes in alpine
meadows, or on limestone, 3,300-4,600 meters; Sacatepe"quez
(Volcan de Fuego); Chimaltenango (Volcan de Acatenango);
Huehuetenango (Che"mal, Sierra de los Cuchumatanes) ; San Marcos
(volcanoes of Tajumulco and Tacana). Higher mountains of central
and southern Mexico.

Plants cespitose and forming very dense, cushion-like mats 10 cm. broad or
sometimes larger, 1-3 cm. high; leaves very densely crowded and imbricate,
coriaceous, oval or oblong, very obtuse, concave, carinate beneath, ciliolate or
eciliolate, glabrous, scarcely more than 3 mm. long; flowers sessile at the ends of
the branches, about 4 mm. long; sepals concave, coriaceous, ciliolate at the base
or eciliate, glabrous, obtuse; capsule 3-valvate; seeds 1-3, black, lustrous.

Var. guatemalensis differs from the type but little, except that
the leaves are mostly eciliate rather than conspicuously ciliate as
are most of the Mexican specimens. This is one of the typical high
alpine plants of Guatemalan mountains, extending to the very
summits of most of the high volcanoes.

Arenaria guatemalensis Standl. & Steyerm. Field Mus. Bot.
23: 50. 1944.

Moist, shaded, brushy or open banks, often in thickets, sometimes
in oak-pine or Juniperus forest, 1,500-3,300 meters; Zacapa; Jalapa;
Sacatepe"quez; Chimaltenango; Solola; Huehuetenango; Quezal-
tenango; San Marcos (type from Rio Vega near San Rafael and
Guatemala-Mexico boundary, Volcan de Tacana, Steyermark 36268).
Doubtless also in southern Mexico, and a variety is known from that
country; Costa Rica; Panama.

Perennial, the stems usually laxly branched, procumbent, prostrate, or often
pendent from banks, sometimes 3 meters long and sprawling or subscandent over
bushes, densely puberulent with mostly reflexed hairs, the internodes mostly longer
than the leaves; leaves sessile or very shortly petiolate, herbaceous, linear-lanceo-
late to elliptic-lanceolate, 1.5-3.5 cm. long, 2-8 mm. wide, acute, densely and
minutely puberulent above, minutely hispidulous beneath on the costa and some-
times puberulent elsewhere, 1-nerved, the margins not thickened and not con-
spicuously ciliate; flowers axillary, the slender pedicels usually much longer than
the leaves, densely and minutely puberulent; sepals about 5 mm. long, the outer
ones lanceolate or linear-lanceolate, attenuate-acuminate, minutely hispidulous,
the inner ones broader, hispidulous only on the costa, the margins scarious, white;
petals about 8 mm. long, always longer than the sepals; styles 3; capsule 5-6 mm.
long, 3-valvate, the valves deeply 2-lobate.

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 221

This has generally been confused with A. megalantha, which it
much resembles, but it is fully and constantly distinct from that
species by the characters enumerated in the key.

Arenaria lanuginosa (Michx.) Rohrb. in Mart. Fl. Bras. 14, pt.
2: 274. 1872. Spergulastrum lanuginosum Michx. Fl. Bor. Amer. 1:
275. 1803. A. alsinoides Willd. ex Schlecht. Ges. Naturf. Freund.
Berlin Mag. 7: 201. 1813.

Moist thickets, brushy or shady banks, oak-pine forest, rocky
slopes, sometimes among rocks along streams or on sandbars, often
in open fields, 700-2,800 meters; Alta Verapaz; Baja Verapaz;
Zacapa; Chiquimula; Jalapa; Santa Rosa; Guatemala; Sacatepe"-
quez; Chimaltenango; Solola; Quiche"; Huehuetenango; Quezalte-
nango. Southern United States; Mexico; Honduras to Panama;
West Indies; South America.

Perennial or sometimes annual, usually much branched from the base, the
stems 15-50 cm. long, slender, procumbent, puberulent or pubescent, rarely gla-
brate; leaves linear to oblong-lanceolate, acute or obtuse, mostly 1-2.5 cm. long,
usually puberulent, often densely so, herbaceous, 1-nerved; pedicels axillary,
slender, longer or shorter than the leaves, finely puberulent; sepals ovate or lance-
ovate, 2-3 mm. long or in fruit slightly elongate, acuminate, puberulent, scarious-
margined; petals white, oblong or oval, obtuse, equaling or somewhat shorter than
the petals; stamens slightly shorter than the calyx; capsule ovoid-oblong; seeds
dark brown, smooth, lustrous.

A common plant in many mountain regions of Central America,
often of a decidedly weedy nature.

Arenaria lycopodioides Willd. ex Schlecht. Ges. Naturf. Freund.
Berlin Mag. 7: 212. 1813. A. decussata Willd. ex Schlecht. loc. cit.

Usually on limestone cliffs or boulders, sometimes in alpine
meadows, 2,400-3,700 meters; Huehuetenango (Sierra de los
Cuchumatanes, where collected in various localities). Mountains
of southern Mexico.

Plants perennial, prostrate, often forming rather dense mats, much branched,
the stems mostly 30 cm. long or less, sometimes laxly branched, slender but rather
stiff, 2-sulcate or subangulate, minutely puberulent in 2 lines or almost wholly
glabrous, often densely leafy; leaves linear or lance-linear, 1.5 cm. long or shorter,
1-2 mm. wide, almost subcoriaceous and rigid, spreading or ascending, subulate-
acuminate, ciliate near the base, otherwise glabrous, the margins cartilaginous-
thickened, the costa stout and very prominent beneath; flowers few, axillary or
terminal, sometimes subpaniculate at the ends of the branches, the pedicels usually
short but often much longer than the leaves, slender, densely and very minutely
puberulent; sepals subcoriaceous, 4 mm. long, oblong-ovate, glabrous, minutely
ciliolate near the base, acute, the costa very stout and prominent; petals white,

222 FIELDIANA: BOTANY, VOLUME 24

about equaling the sepals or sometimes exceeding them; stamens equaling the
sepals.

A typically alpine plant of the Cuchumatanes.

Arenaria megalantha (Rohrb.) F. N. Williams, Journ. Linn.
Soc. Bot. 33: 379. 1898. A. lanuginosa var. megalantha Rohrb.
Linnaea 37: 264. 1871-72. A. alsinoides var. ovatifolia Bonn. Smith,
Bot. Gaz. 18: 198. 1893 (type from Volcan de Agua, W. C. Shannon
3635).

Moist steep slopes or on shaded banks or cliffs, sometimes in
pine forest, 2,200-3,500 meters; Sacatepe"quez (Volcan de Agua);
Quezaltenango. Mountains of southern Mexico.

Perennial, prostrate, the stems usually much branched and interlaced, slender,
densely pilose with short, rather stiff, fulvous, spreading hairs; leaves sessile or
nearly so, stiff, spreading, ovate or broadly ovate, as much as 12 mm. long and
8 mm. wide, acute or subulate-acuminate, obtuse or broadly rounded at the base,
densely hispidulous on both surfaces, long-ciliate, the margins cartilaginous-
thickened, the costa stout and prominent beneath; flowers axillary, the pedicels
usually several times as Ipng as the subtending leaves, very slender, densely
hispidulous; sepals oblong-ovate, 4.5 mm. long, subulate-acuminate, densely
hispidulous over the whole outer surface; petals white, sometimes twice as long
as the sepals; styles 3; capsule ovoid-oblong; seeds spheroid-lenticular.

Arenaria paludicola Robinson, Proc. Amer. Acad. 29: 298.
1894.

Around edge of water of small depressions in alpine meadow,
3,400-3,500 meters; Huehuetenango (vicinity of Tunima, Sierra de
los Cuchumatanes, Steyermark 48318). California; Chihuahua.

Plants glabrous, flaccid, the stems branched, sometimes creeping, rooting
at the lower nodes, remotely leafy; leaves linear, flat, 1-nerved, acute, spreading,
3.5 cm. long or less, about 2 mm. wide, slightly scabrous on the margins, often
somewhat connate at the base; pedicels axillary, solitary, as much as 5 cm. long
but usually shorter, spreading or somewhat deflexed; sepals ecostate, herbaceous,
4.5-5 mm. long, subacute, glabrous; petals obovate, about twice as long as the
sepals; capsule ovoid, 3-valvate, the valves entire.

If the Guatemalan plant is correctly determined, the range of
this species is an extraordinary one. The material is not in good
condition for study, and it may well be that a new species is repre-
sented, but there are no obvious characters by which it may be
separated from collections made in northern Mexico.

Arenaria reptans Hemsl. Diagn. PI. Mex. 22. 1879.
Moist or wet, usually dense forest, often in forest of pine, Juni-
perus, Cupressus, or Abies, sometimes on moist open banks or in wet

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 223

fields or on mossy logs, frequently on white-sand slopes, 1,500-4,600
meters, chiefly at high elevations; El Progreso; Chiquimula; Jalapa;
Sacatepe"quez; Retalhuleu; Solola; Chimaltenango; Totonicapan;
Huehuetenango; Quezaltenango; San Marcos. Central and southern
Mexico.

Plants perennial from a slender root, the stems usually branched, prostrate,
and often forming dense mats, often rooting at the nodes, more or less angulate,
hispidulous or puberulent; leaves numerous, very small, obovate-lanceolate or
ovate-lanceolate, mostly less than 5 mm. long, obtuse and cuspidate-apiculate or
often acuminate, usually abruptly contracted at the base into a rather long petiole,
sometimes attenuate to the petiole, often fasciculate, usually conspicuously long-
ciliate with white hairs, at least on the petiole, generally hispidulous or white-
pilose beneath, white-punctate; flowers axillary, the pedicels very slender, mostly
much longer than the leaves; sepals 2.5-3 mm. long, ovate-oblong or lanceolate,
obtuse or cuspidate, membranaceous-marginate, glabrous, sometimes ciliate;
petals none; capsule about equaling the sepals; seeds lenticular, rufous-black.

Var. Pringlei F. N. Williams (Journ. Linn. Soc. Bot. 33: 383.
1898; type from Sierra de San Felipe, Oaxaca) is a densely cespitose
form with short and densely leafy rather than elongate branches.
It scarcely deserves nomenclatorial recognition and at best is a mere
form.

CERASTIUM L.

Plants annual or perennial, usually pubescent, often viscid; flowers small,
white, in terminal dichotomous cymes; sepals normally 5; petals 5, emarginate or
bifid at the apex, rarely absent; stamens 10 or rarely fewer; styles as many as the
sepals and opposite them, sometimes fewer; capsule 1-celled, cylindric, often
curved, dehiscent by 10 or rarely 8 apical teeth; seeds numerous, rough, more or
less compressed, attached by their edges.

Species about 50, widely distributed, chiefly in temperate regions.
Only the following are known from Central America.

Cauline leaves ovate, elliptic, or obovate, very obtuse or rounded at the apex;

pedicels mostly shorter than the calyx C. riscosum.

Cauline leaves linear to lance-oblong, acute or acuminate.
Pedicels shorter than the calyx, the inflorescence dense and congested.

C. brachypodum.
Pedicels all or mostly much longer than the calyx, the inflorescence open.

Calyx scarcely 4 mm. long; stems white-lanate below C. vulcanicum,

Calyx 5-8 mm. long; stems villous, not lanate.

Calyx 7-8 mm. long, the petals much longer; cauline leaves lance-oblong,

mostly 5-8 mm. wide C. Juniperorum.

Calyx about 5 mm. long; cauline leaves linear or lance-linear, usually
narrower C. guatemalense.

Cerastium brachypodum (Engelm.) Robinson ex Britton,
Mem. Torrey Club 5: 150. 1894. C. nutans var. brachypodum
Engelm. ex Gray, Man. ed. 5. 94. 1867.

224 FIELDIANA: BOTANY, VOLUME 24

Alpine meadows, 3,300-3,700 meters; Huehuetenango (Sierra de
los Cuchumatanes). Central and western United States, south
through Mexico.

Annual, the plants light green, especially when dried, viscid-pubescent or
short- villous throughout; stems simple or branched, often several from each root,
mostly 10-15 cm. high, erect or ascending; lower and basal leaves spatulate or
oblanceolate, obtuse or subacute, 2.5 cm. long or less, narrowed into a short
petiole; upper cauline leaves linear or lance-linear, sessile; cymes few-several-
flowered, the fruiting pedicels nutant or deflexed, shorter than the calyx or but
slightly longer, the inflorescence congested at anthesis but in fruit more open and
the pedicels more elongate; sepals ovate-oblong, about 4.5 mm. long; petals shorter
than the sepals or slightly exceeding them; capsule about 9 mm. long, straight or
slightly curved, pale stramineous, transparent, 2-3 times as long as the calyx.

More ample specimens may show the Guatemalan plant to be
an undescribed species, since it does not appear to be referable to
any other species known from Mexico.

Cerastium guatemalense Standl. Field Mus. Bot. 17: 244.
1937.

Alpine or subalpine slopes, mostly in open pine forest, some-
times on open exposed ridges, 2,300-4,000 meters or even higher;
endemic so far as known, but to be expected in the mountains of
southern Mexico; Sacatepe"quez (type from upper slopes of Volcan
de Agua, J. R. Johnston 816); Chimaltenango; Quezaltenango; San
Marcos.

Probably perennial, or sometimes annual, the stems often several from each
root, erect or decumbent, 40 cm. long or less, densely viscid-villous with spreading
hairs, the whole plant rather pale green; leaves linear or lance-linear, sessile,
2-3.5 cm. long, 2-4 mm. wide near the base, gradually attenuate to the acute apex,
1-nerved, viscid- villosulous on both surfaces; inflorescence laxly cymose, few-many-
flowered, the flowers nutant, the slender pedicels mostly 1-3 cm. long, densely
viscid-villous; sepals 5-6 mm. long, oblong-ovate, acuminate, densely viscid-
villosulous, the margins pale and hyaline; petals white, 7 mm. long or less; capsule
slightly curved, 12-13 mm. long, with very short teeth; seeds brown, coarsely
tuberculate, 1.2 mm. in diameter.

One of the characteristic plants of the summits of the higher
volcanoes. The plants are dry during the verano, growing only during
the rainy season.

Cerastium Juniperorum Standl. & Steyerm. Field Mus. Bot.
23: 51. 1944.

Alpine meadows, 3,400-3,700 meters; endemic; Huehuetenango
(type from vicinity of Tunima, Sierra de los Cuchumatanes, Steyer-
mark 48413; known only from this locality).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 225

Probably perennial, the stems solitary or few together, erect or decumbent,
20-35 cm. long, simple, densely viscid-villosulous with short spreading hairs, the
internodes mostly much longer than the leaves; leaves sessile, spreading, herba-
ceous, oblong-lanceolate, 1.5-3.5 cm. long, 5-8 mm. wide, acute, with a callous-
thickened tip, obtuse at the base, densely pubescent on both surfaces with short
spreading hairs, 1-nerved; cymes terminal, few-flowered, the pedicels very slender,
apparently straight, as much as 3.5 cm. long, densely viscid-pubescent; sepals 7-8
mm. long, oblong-lanceolate, green, scarious-margined, viscid-villosulous; petals
white, 1 cm. long, conspicuously longer than the sepals.

Perhaps only an extreme form of C. guatemalense, but it appears
to be a quite distinct species.

Cerastium viscosum L. Sp. PI. 437. 1753.

Moist thickets, open fields or banks, dry rocky hillsides, sand-
bars along streams, or very often a weed in cultivated ground,
1,350-3,300 meters; Alta Verapaz; Baja Verapaz; Jalapa; Guate-
mala; Sacatepe'quez ; Chimaltenango; Quiche"; Huehuetenango;
Quezaltenango; San Marcos. Native of Europe but now widely
naturalized in North America, from Canada south to Mexico; Costa
Rica; West Indies; South America.

Annual, erect or decumbent, the stems sometimes spreading, often much
branched, very leafy, densely viscid-pubescent, 30 cm. long or less; leaves green,
sessile or petiolate, ovate to obovate or elliptic, 1-2.5 cm. long, 6-15 mm. wide,
obtuse or rounded at the apex, apiculate, pilose on both surfaces; inflorescence
many-flowered, the flowers short-pedicellate, congested, the cymes more open in
fruit; sepals lance-oblong, 4 mm. long, acuminate, green, pilosulous; petals usually
shorter than the sepals, 2-cleft; capsule 7-8 mm. long, almost straight, twice as
long as the sepals.

A very common weed in waste and cultivated ground in the
central and western mountains. It is thoroughly established also
in the mountain pastures of Costa Rica.

Cerastium vulcanicum Schlecht. Linnaea 12: 208. 1838.

Alpine meadows, 3,250 meters; Huehuetenango (Sierra de los
Cuchumatanes, A.F. Skutch 1219). High mountains of central and
southern Mexico.

Annual, erect or decumbent, often much branched from the base, densely
lanate almost throughout, especially on the leaves and lower part of the stems,
with matted white hairs, also viscid-pubescent, 12-24 cm. high; cauline leaves
linear-lanceolate or oblanceolate, mostly 2 cm. long or less, acute, sessile; stems
cymosely branched above, usually many-flowered, the pedicels mostly 6-10 mm.
long, the flowers nutant; sepals about 4 mm. long, ovate-oblong, acute, scarious-
margined, viscid-villosulous; petals white, deeply bifid, scarcely longer than the

226 FIELDIANA: BOTANY, VOLUME 24

sepals; capsule 8 mm. long, twice as long as the calyx, rather broad, hyaline, pale-
stramineous.

DIANTHUS L.

Rather stiff, perennial herbs, sometimes biennial or annual, the leaves narrow;
flowers small or large, often very showy, terminal, solitary or cymose-paniculate,
usually colored; calyx 5-dentate, multistriate, tubular, with several bracts at the
base; petals 5, unguiculate, dentate or crenate; stamens 10; styles 2; ovary 1-celled,
stipitate; capsule cylindric or oblong, stipitate, dehiscent at the apex by 4-5
short teeth; seeds numerous, compressed, attached laterally; embryo straight,
excentric.

Species 200 or more, all except one in temperate and cool regions
of the Old World. Several have been naturalized in America, and
some are widely cultivated for ornament. The following species are
common in cultivation in Guatemala and some other ones may be
planted occasionally.

Flowers densely crowded in flat-topped clusters D. barbatus.

Flowers solitary or in loose 2-3-flowered inflorescences.

Bracts at the base of the calyx short and broad, closely appressed; leaves

glaucous; flowers double D. Caryophyllus.

Bracts at the base of the calyx linear, spreading or recurved; flowers rarely
double D. chinensis.

Dianthus barbatus L. Sp. PI. 409. 1753. Clavel imperial.
Sweet William.

Native of Europe, but grown widely as an ornamental plant;
planted commonly in Guatemalan gardens.

Perennial or biennial, erect, glabrous, the stems rather stout, 30-60 cm. high,
usually branched above; leaves lanceolate or ovate-lanceolate, green, 3.5-7 cm.
long, acute, the basal leaves oblong or obovate; bracts at the base of the calyx
linear-filiform, about equaling the calyx, this deeply dentate; flowers small, very
variable in coloring, crowded in dense flat-topped clusters.

Sweet William is a popular garden flower of the mountain regions
of Guatemala and is often grown for sale in the markets.

Dianthus Caryophyllus L. Sp. PI. 410. 1753. Clavel. Carna-
tion.

Usually stated to be native in the Mediterranean region, but
cultivated in most civilized regions of the earth for its beautiful,
often sweet-scented flowers; a common garden plant of the moun-
tains of Guatemala. The carnation is found in most gardens at
middle and high elevations, and thrives exceptionally well when

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 227

planted in the ground. Vast quantities of the flowers are offered
for sale in most of the upland markets, particularly that of Guate-
mala. They are one of the best flowers for making funeral coronas
and also are used for decorating houses, churches, and roadside
shrines. There are many color varieties. In central Guatemala
the chief region in production of this and other cut flowers is San
Juan Sacatepe"quez, not far from Guatemala City, where there are
large areas, consisting of many small properties, devoted to flower
growing, the ground usually formed into small rectangular elevated
beds and carefully watered by hand during the long dry season
(verano). A newspaper account was noted in which it was stated
that more than a million carnation plants were growing about San
Juan. Large loads of carnations and other cut flowers are carried
on men's backs for sale from San Juan to Guatemala City, about
25 km. distant, to Antigua, 60 km. away, and even to more remote
markets. Packed tightly, covered with moist cloths, and usually
transported before daylight, ' the flowers retain their freshness
perfectly in these distant markets.

Dianthus chinensis L. Sp. PI. 411. 1753. Clavellina.

Probably native in China and Japan, but widely grown for
ornament in other regions; a common garden flower of Guatemala,
from the lowlands to the highlands, and also in other parts of Central
America.

Plants erect, cespitose, glabrous, sometimes repent at the base; leaves linear
or lance-linear, 3-5-nerved; flowers rather large, solitary or in lax clusters, mostly
pink or lilac or in part dark red, rarely if ever double, the petals dentate or laciniate
on the margins.

DRYMARIA Willdenow

Mostly small, annual or perennial, very slender herbs, diffuse or erect, dicho-
tomously branched, glabrous or pubescent; leaves small, broad or narrow, the
stipules small, often fugacious; flowers small, pedicellate, solitary in the forks of
the branches or in terminal or axillary cymes; sepals 5, herbaceous or scarious-
margined; petals 5 and 2-6-cleft; stamens 5 or by abortion fewer, subperigynous;
ovary 1-celled, many-ovulate, the style 3-fid; capsule 3-valvate; seeds reniform-
globose or laterally compressed, the hilum lateral; embryo peripheral.

Species 30 or more, in tropical America, a few reaching the south-
western United States. Only the following are known in Central
America, most of the species being Mexican.

Plants dwarf, erect, mostly 3 cm. high or less D. minuscula.

Plants with elongate stems mostly 10 cm. long or more, often decumbent or
prostrate.

228 FIELDIANA: BOTANY, VOLUME 24

Flowers sessile or nearly so, in dense head-like cymes.

Stems glabrous D. leptoclados.

Stems densely glandular-puberulent D. ramosissima.

Flowers slender-pedicellate, mostly in lax open cymes.

Pedicels glabrous; sepals very obtuse, broadly ovate, less than 3 mm. long.

D. palustris.
Pedicels variously pubescent, villosulous, glandular-pubescent, or minutely

farinose-puberulent; sepals often acute or acuminate.
Upper leaves ovate or broadly ovate, acute or obtuse, mucronate, distinctly
longer than broad.

Sepals 3.5 mm. long; leaves laxly reticulate- veined D. hyperidfolia.

Sepals 5.5 mm. long; leaves not reticulate-veined D. laxiflora.

Upper leaves suborbicular, broadly rounded at the apex, not or scarcely

mucronate, as broad as long or often broader.

Young pedicels appearing thickened, covered with a dense minute
whitish tomentum-like puberulence, this becoming more sparse in

age D. cordata.

Young pedicels filiform, not minutely tomentulose, villosulous, sparsely

puberulent, or rarely glabrous.
Stems glandular-puberulent throughout, or at least above.

D. glandulosa.
Stems glabrous or villosulous.

Leaves usually densely or sparsely villous beneath; stems sparsely

villous D. villosa.

Leaves usually glabrous; stems glabrous '. . .D. gracilis.

Drymaria cordata (L.) Willd. ex Roem. & Schult. Syst. Veg. 5:
406. 1819. Holosteum cordatum L. Amoen. Acad. 3: 21. 1756.

Moist thickets, shaded banks, or forest, often a weed in waste
or cultivated ground, especially in cafetales, 900 meters or less;
Pete"n; Izabal; Alta Verapaz; Zacapa; Chiquimula; Jutiapa; Santa
Rosa; Suchitepe"quez; Retalhuleu; Quezaltenango; San Marcos.
Mexico; Honduras to Costa Rica; West Indies; South America.

Plants very slender, annual, erect or procumbent, usually much branched,
the stems 10-30 cm. long or more, glabrous or nearly so; leaves short-petiolate,
ovate-rounded or orbicular, 5-25 mm. long, pale green, rounded at the apex,
rounded or shallowly cordate at the base; flowers greenish white, in lax terminal
cymes, the pedicels, at least at first, covered with a dense whitish glandular
tomentum-like pubescence and appearing thickened; sepals lanceolate or ovate,
3-4 mm. long, usually glabrous, acute, scarious-margined; petals 2-parted, gener-
ally shorter than the sepals; capsule ovoid, slightly exceeding the persistent sepals.

Called "palitaria" or "pelitaria" in Honduras; "petatillo,"
"comida de canario," "trencilla," "comapa," "comapona" (Salva-
dor). One of the common weedy plants of Central American low-
lands, mostly in moist shaded places. It is easily recognized, for
it has whitish pedicels, which are somewhat thickened rather than

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 229

almost capillary as in other species, and have the appearance of
having been attacked by a mildew.

Drymaria glandulosa Bartling in Presl, Rel. Haenk. 2: '9.
1835-36.

Moist forest or thickets, 2,700 meters or less; Izabal; Jalapa;
Escuintla; Sacatepe"quez ; San Marcos; reported from Quiche".
Central and southern Mexico.

Similar in habit and appearance to D. cordata but finely glandular-pubescent
on the stems and pedicels; leaves on short slender petioles, small, ovate-rounded
or broadly orbicular, rounded at the apex, rounded or subcordate at the base;
flowers greenish white, in lax terminal cymes, slender-pedicellate, very numerous;
sepals lance-oblong, 4-5 mm. long, acute, glandular-puberulent or glabrous; petals
shorter than the sepals, short-bifid; stamens 5; capsule much shorter than the
calyx, 5-6-seeded; seeds subreniform, fuscous, granulate-tuberculate in lines.

Drymaria gracilis Cham. & Schlecht. Linnaea 5: 232. 1830.
D. multiflora Brandeg. Zoe 5: ^32. 1906.

Moist or wet forest or thickets, 1,500-2,700 meters; Chiquimula;
Sacatepe"quez; Chimaltenango; Quiche"; Quezaltenango. Southern
Mexico.

A slender annual, in general appearance like D. cordata, ascending, procum-
bent, or sometimes subscandent, usually much branched, the stems glabrous;
leaves on slender, often much elongate petioles, thin, pale green, orbicular or
ovate-orbicular, rounded at base and apex, glabrous; flowers greenish, in lax
cymes, the pedicels capillary, glabrous or sometimes puberulent or glandular-
puberulent; sepals ovate, green, scarious-margined, 3-4 mm. long, obtuse or
acute; petals scarcely equaling the sepals; seeds few, much larger than in D.
cordata, dark reddish brown, coarsely granulate-tuberculate.

Drymaria hypericifolia Briquet, Ann. Cons. Jard. Geneve 13
& 14: 369. 1911.

Moist or wet, usually dense forest or mountain thickets, often in
oak, pine, or Alnus forest, sometimes on white-sand slopes, 1,500-
3,400 meters; Zacapa; Jalapa; Guatemala; Sacatepe"quez; Chimal-
tenango; Quezaltenango; San Marcos. Southern Mexico.

Plants probably perennial, very slender, erect or often reclining on other
plants, frequently pendent from banks, sparsely or much branched, the stems
often a meter long, green, terete, glabrous or sparsely puberulent; leaves on short
slender petioles, broadly ovate or rounded-ovate, 1-2.5 cm. long, glabrous, obtuse
or subobtuse, rounded at the base and abruptly contracted, thin but rather stiff,
deep green above, paler beneath, conspicuously and laxly reticulate-veined, tripli-
nerved; cymes few-flowered, very lax, glabrous or minutely pilosulous, the flowers

230 FIELDIANA: BOTANY, VOLUME 24

on capillary pedicels; sepals in an thesis 3.5 mm. long, somewhat elongate in fruit,
oblong-lanceolate, subobtuse, minutely pilosulous or almost glabrous, green,
white-marginate; petals white, 2-fid, somewhat exceeding the sepals; stamens 5,
equaling the petals.

Drymaria laxiflora Benth. PI. Hartweg. 73. 1841.

Type collected on rocks near Zunil, Quezaltenango, Hartweg 523.
Reported from southern Mexico.

Plants glabrous, much branched, diffuse; leaves on rather long, slender petioles,
broadly ovate, acute, mucronate, 4-8 mm. long and almost as broad, rounded at
the base and abruptly short-decurrent; stipules several, setaceous, almost equaling
the petioles; cymes lax, few-flowered, the bracts lanceolate, scarious-margined;
pedicels 4-6 mm. long, filiform, glabrous, or with a few minute viscid hairs; sepals
5.5 mm. long, narrowly lanceolate, scarious-margined; petals deeply 2-fid, scarcely
longer than the sepals; stamens usually 5; valves of the capsule generally 3, some-
times 4; seeds about 20, muriculate.

Rather strangely, this species is not represented in recent Guate-
malan collections although we have collected many plants in the
general region of the type locality. We have seen type material of
the species and find the sepals in two available specimens much
longer than described by Bentham (he states they are 2 lines long,
i.e. 4 mm.).

Drymaria leptoclados Hemsl. Diagn. PL Mex. 2. 1878.

Known only from the type, Bernoulli 240, from "Camino del
Sapote"; there are at least 23 settlements in Guatemala that bear
the name Zapote, and we do not know which is the one where the
type was collected.

Plants annual, erect, 7-15 cm. high, glabrous throughout, the branches terete,
almost filiform; leaves on very short petioles, membranaceous, broadly ovate-
rounded, acute or mucronulate, 5-7-nerved, 6-10 mm. wide, the stipules setiform;
flowers small, in dense terminal cymes, almost sessile; sepals paleaceous, oblong-
lanceolate, mucronulate, 4 mm. long or less, the costa prominent, the 2 lateral
nerves inconspicuous; petals very narrow, shorter than the sepals, deeply 2-parted;
capsule oblong, about as long as the sepals, 3-valvate, few-seeded; seeds minute,
hippocrepiform, punctulate.

We have seen no material of this species. If correctly described
as glabrous, and there is no reason to doubt that it is, it must be a
rare species.

Drymaria minuscula Standl. & Steyerm. Field Mus. Bot. 23:
52. 1944.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 231

On rocky limestone outcrops under Juniperus Standleyi, 3,700
meters; Hue'huetenango (type from Che'mal, summit of Sierra de
los Cuchumatanes, Steyermark 50243). State of Mexico, Mexico.

An erect annual only 1-3 cm. high, glabrous, densely branched from the base,
the stems slender, terete, pale; basal leaves rosulate, oblanceolate-spatulate, 8 mm.
long or shorter, obtuse, attenuate to the base; cauline leaves linear-oblanceolate,
of about the same length, obtuse, sessile, attenuate to the base, the upper leaves
minute and bract-like; inflorescence repeatedly dichotomous, dense, many-
flowered, the flowers small, on very short pedicels; sepals 1.5-2 mm. long, obtuse,
erect, slightly excurved at the apex, obscurely carinate; petals shorter than the
sepals, white; stamens 5, much shorter than the sepals; style short, with 3 short
branches.

Differing from all other local species in its greatly reduced size.

Drymaria palustris Cham. & Schlecht. Linnaea 5: 232. 1830.

Wet banks or marshes, most often at the edges of streams, some-
times on rocks at the edge of water or in pine forest, 650-3,400
meters, most common at higher elevations; Jalapa; Guatemala;
Sacatepe"quez; Chimaltenango; Retalhuleu; Totonicapan; Quezalte-
nango; San Marcos; Huehuetenango. Southern Mexico.

Plant probably perennial, in general appearance similar to D. cordata but
smaller in most of its parts, glabrous throughout or nearly so, the stems very
slender, usually branched, prostrate or ascending; leaves bright green, on short
filiform petioles, orbicular, ovate-orbicular, or reniform-orbicular, 4-8 mm. wide,
rounded at the apex, rounded or subcordate at the base, sometimes sparsely
villous beneath; flowers greenish white, solitary or in few-flowered cymes, on
filiform pedicels; sepals ovate, obtuse, 2 mm. long, green, white-margined; petals
equaling or shorter than the sepals; seeds slightly larger than in D. cordata, brown,
minutely tuberculate.

We have seen a photograph of the type, formerly in the Berlin
herbarium. The photograph is not a very good one nor was the
specimen an ample one, but both it and the original description seem
to agree well with the numerous Guatemalan specimens. The
specific name is a most appropriate one since the plant usually is
found in wet soil, most often in places where the leaves are always
wet with dew or the spray of running water.

Drymaria ramosissima Schlecht. Linnaea 12: 206. 1838.

Open banks or fields, often a weed in cornfields or cafetales, some-
times in oak forest, frequent on sandbars along streams, 1,350-2,700
meters; Jalapa; Guatemala; Sacatepe"quez ; Chimaltenango; Quiche";
Huehuetenango ; Quezaltenango ; San Marcos. Central and southern
Mexico.

232 FIELDIANA: BOTANY, VOLUME 24

Plants annual, erect or ascending, usually much branched from the base or
throughout, the stems sometimes 40 cm. long but usually shorter, rather densely
glandular-puberulent; leaves short-petiolate, ovate-orbicular or orbicular-reni-
form, mostly 6-15 mm. wide, usually abruptly acute or apiculate, rounded or
somewhat cordate at the base, glabrous or puberulent; cymes dense and con-
gested, terminal, numerous, the flowers numerous, sessile' or short-pedicellate;
sepals lance-oblong, rather rigid, acute or subulate-acuminate, glandular-puberu-
lent, green, white-margined, carinate, the 2 lateral nerves obvious or obscure;
petals white, shorter than the sepals, 2-fid to below the middle; stamens 5, shorter
than the petals; capsule half as long as the sepals, 3-valvate; seeds usually 2-3,
small, brown, orbicular, tuberculate-papillate.

A very common, weedy plant in the central mountains, especially
in old cornfields.

Drymaria villosa Cham. & Schlecht. Linnaea 5: 232. 1830.
D. idiopoda Blake, Contr. U. S. Nat. Herb. 24: 4. 1922 (type from
Dept. Copan, Honduras). Poleo; Llovizna blanca; Milldn.

Moist or wet fields, sometimes in marshes, often in thickets or
on sandbars along streams, frequently a weed in cultivated fields,
300-2,500 meters; Alta Verapaz; Baja Verapaz; El Progreso; Zacapa;
Chiquimula; Santa Rosa; Guatemala; Sacatepe"quez; Chimaltenango;
Quich^ ; Huehuetenango; Quezaltenango. Mexico; Honduras to
Salvador and Panama.

Plants very slender, erect or decumbent, annual, usually much branched, the
stems 10-30 cm. long, sparsely or densely short- villous; leaves on slender, often
long petioles, ovate-orbicular to reniform-orbicular, 1-2 cm. wide, rounded or
very obtuse at the apex, sometimes subacute, rounded or subcordate at the base,
usually villous beneath and often also on the upper surface; cymes very lax,
generally few-flowered, numerous, the pedicels almost filiform, short or elongate,
viscid-villous; sepals acute or obtuse, 3 mm. long, usually villosulous, green with
whitish margins; petals 2-parted, shorter than the sepals; seeds small, tuberculate.

GYPSOPHILA L.

Plants annual or perennial, branched, erect or spreading, mostly glabrous and
glaucous, the leaves narrow; flowers small, numerous, axillary or paniculate; calyx
turbinate or campanulate, 5-nerved, 5-dentate, naked at the base; petals 5, entire
or emarginate, narrow-unguiculate; stamens 10; styles 2; capsule 4-valvate to the
middle or less deeply; seeds reniform, attached laterally; embryo coiled.

About 60 species, in Europe, Asia, and northern Africa. One
or two species often are grown for ornament.

Gypsophila elegans Bieb. Fl. Taur. Cauc. 1 : 319. 1808. Gipso-
fila; Sofilia; Llovizna.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 233

Native of the Caucasus region but often grown for ornament in
other parts of the earth; much planted in gardens of Guatemala,
especially for market; found thoroughly naturalized in an opening
in oak forest near San Juan Sacatepe*quez, Guatemala, 1,800 meters.

Annual, erect or ascending, 30-60 cm. high, glabrous, repeatedly dichotomous,
somewhat glaucous; leaves sessile, the cauline ones linear or linear-lanceolate, the
lowest oblong or narrowly spatulate; flowers very numerous, small, white, forming
large cymes or panicles; sepals about 3 mm. long, the petals twice as long or more,
truncate, almost recurved.

Much grown in the mountains of Guatemala, especially for use
as a filler in funeral wreaths or other formal designs. As they appear
in the markets, the flowers, as happens so frequently with other
white blossoms, often are colored red, pink, blue, yellow, etc. with
dyes purchased in the shops for the purpose.

Lychnis coronaria (L.) Desr. is planted rarely for ornament in
gardens of Guatemala. It is a native of Europe, a tall coarse plant,
densely white-tomentose throughout, the few large flowers 2.5-3 cm.
broad and dull rose-colored.

SAGINA L.

Dwarf, annual or perennial herbs, often tufted or matted; leaves small, linear
or subulate, few, the flowers minute, pedicellate, whitish; sepals 4-5; petals 4-5,
entire or emarginate, sometimes none; stamens as many as the sepals or twice as
many; ovary 1-celled, many-ovulate; styles as many as the sepals and alternate
with them ; capsule 4-5-valvate, finally dehiscent to the base, the valves opposite
the sepals.

Species about 10, in the northern hemisphere. No others are
known from Central America.

Sagina procumbens L. Sp. PI. 128. 1753.

Dry or moist, shaded banks in forest or often in open fields, waste
ground about dwellings, on sandbars along streams, very often a
weed between cobblestones in streets, sometimes on limestone in
forest of Juniperus Standleyi, 1,400-4,000 meters; Alta Verapaz;
Guatemala; Sacatepe"quez; Chimaltenango; Quiche*; Huehuetenango;
Quezaltenango; San Marcos. Widely distributed in North America,
south through Mexico (where uncommon); South America; Eurasia.

Plants mostly annual, usually much branched, decumbent or spreading,
glabrous or nearly so, the slender stems 2-6 cm. long; leaves linear-subulate, mostly
6 mm. long or less, connate at the base; flowers greenish white, numerous, about
2 mm. broad; pedicels capillary, longer than the leaves, the flowers sometimes

234 FIELDIANA: BOTANY, VOLUME 24

nutant; sepals generally 4, ovate-oblong, obtuse; petals shorter than the sepals or
absent; stamens 4; capsule about equaling the calyx, the seeds dark brown.

This is a very common weed between cobblestones in Quezalte-
nango, Chichicastenango, and other cities of Guatemala, a small
and inconspicuous plant. There is little doubt that it is native in
the mountains of Guatemala, rather than imported from Europe
as it is believed to be in some regions of America.

Saponaria officinalis L., bouncing Bet, a native of Europe, was
observed in patios at Antigua and Jalapa, but it is a rare plant in
Central America. It is a rather coarse, glabrous perennial with
large pink flowers that has become naturalized in many parts of the
United States.

SILENE L.

Annual or perennial herbs, often with viscid pubescence; flowers small or large
and showy, mostly pink, red, or white, solitary or cymose; calyx somewhat inflated,
tubular to campanulate, 5-dentate or 5-cleft, 10-many-nerved, not bracteate at
the base; petals 5, unguiculate, usually with a scale near the base of the blade;
stamens 10; styles 3, rarely 4-5; ovary 1-celled or incompletely 2-4-celled; capsule
dehiscent by 6 or rarely 3 apical teeth; seeds mostly echinate or tuberculate.

About 250 species, in both hemispheres, mostly in temperate
regions. None are native in Central America but several are indige-
nous in Mexico, and one or two of them might be expected in the
mountains of western Guatemala.

Plants glabrous, cultivated; flowers deep pink or rarely white S. Armeria.

Plants abundantly and coarsely pubescent, naturalized as a weed; flowers white
or pale purplish S. gallica.

Silene Armeria L. Sp. PI. ed. 2. 601. 1762. Llovizna; Espanola.

Native of Europe but often grown for ornament in other parts
of the earth; planted frequently in Guatemalan gardens, at almost
all elevations.

Annual, erect, branched, glabrous and glaucous, sometimes minutely puberu-
lent, 60 cm. high or less, the stems glutinous below the nodes; basal leaves oblance-
olate, 5-7 cm. long, obtuse; cauline leaves ovate or ovate-lanceolate, 2.5-7 cm.
long, acute or obtuse; flowers numerous, in a dense terminal compound cyme;
flowers deep pink or rarely white, 12-18 mm. broad; calyx clavate, 1-1.5 cm. long,
slightly dilated by the ripe capsule; petals emarginate, each bearing a narrow
scale.

Silene gallica L. Sp. PI. 417. 1753. S. anglica L. op. cit. 416.
Hierba de recluta (fide Aguilar).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 235

Most common as a weed in cornfields or other cultivated ground,
sometimes on sandbars or in moist open fields or thickets, common
in many localities, 1,300-2,700 meters; Jalapa; Guatemala; Sacate-
pe"quez; Chimaltenango; Quiche"; Quezaltenango; San Marcos.
Native of Europe, but widely naturalized in temperate and sub-
tropical America, in Central America in the mountains; Costa
Rica; South America.

Annual, villous-hirsute throughout with whitish hairs, viscid above, the stems
usually several, branched, erect or spreading, 15-50 cm. high; leaves spatulate or
oblanceolate, mostly 2-5 cm. long, obtuse or rounded at the apex, often apiculate,
narrowed to the broadly winged petiole, the uppermost leaves often narrower and
acute; flowers in terminal simple secund spike-like racemes, subsessile or the lower
flowers distant and conspicuously pedicellate; calyx cylindric or oblong-tubular in
anthesis, much enlarged and ovoid in age, 10-nerved, 8-10 mm. long, villous,
contracted at the apex in fruit, the teeth lanceolate, spreading; petals dentate,
entire, or 2-cleft, white or dull purplish, slightly longer than the calyx.

This is often an abundant weed in old cornfields in the central
mountains.

Silene pendula L., native of the Mediterranean region, was found
in cultivation in a garden at San Sebastian, San Marcos, but it is
uncommon in Guatemala. It is a rather coarse plant, abundantly
pubescent, with spatulate leaves, the rather large, rose-colored
flowers axillary and pedicellate, the calyx somewhat inflated and
conspicuously green-costate.

SPERGULA L. Spurry

Annuals, erect or spreading, usually much branched, viscid-pubescent; leaves
subulate, fasciculate in the leaf axils and appearing verticillate, stipulate; flowers
very small, whitish, in lax terminal cymes; sepals and petals each 5; stamens 10
or 5; styles 5, alternate with the sepals; capsule 5-valvate, the valves opposite the
sepals; seeds compressed, with acute or winged margins.

Species about 6, natives of the Old World. One has become
naturalized rather sparingly in North America.

Spergula arvensis L. Sp. PI. 440. 1753.

Sandy, moist or dry fields, especially in cultivated ground,
sometimes on sandbars along streams, 1,500-2,600 meters; Quezal-
tenango. Native of Europe; naturalized in some parts of Canada
and United States; apparently unknown in Mexico or elsewhere in
Central America.

Plants much branched, suberect or spreading, rather sparsely pubescent, the
stems 50 cm. long or usually much shorter; leaves linear or subulate, 2-5 cm. long,

236 FIELDIANA: BOTANY, VOLUME 24

appearing to be inserted in verticels of very numerous leaves, the stipules small,
scarious, connate; flowers numerous, 4-6 mm. broad, in lax cymes, often subumbel-
late, the pedicels long and very slender, divaricate; sepals ovate, obtuse, 3-4 mm.
long, viscid-pubescent; petals slightly shorter than the sepals; stamens 10 or 5 on
flowers of the same plant; capsule ovoid, slightly longer than the calyx; seeds
black, minutely white-papillose.

Rather widely distributed in the vicinity of Quezaltenango but
not common, at least during the dry months.

STELLARIA L. Chickweed

Mostly annual herbs, generally diffusely branched and spreading, the leaves
broad or narrow (broad in Central American species) ; flowers small, white, cymose;
sepals 5, rarely 4; petals as many as the sepals, usually deeply 2-cleft or 2-parted,
rarely none; stamens 10 or fewer, hypogynous; ovary 1-celled, the ovules several or
many; styles mostly 3, rarely 4-5, usually opposite the sepals; capsule globose,
ovoid, or oblong, dehiscent by twice as many valves as there are styles; seeds
smooth or roughened, globose or compressed.

About 75 species, widely distributed, most numerous in temperate
or cold regions, in the tropics confined to mountain regions. One
other species is known from the mountains of southern Central
America (Costa Rica).

Leaves rounded to acute at the apex, often short-cuspidate, rounded to acute at
the base, never cordate.

Petals shorter than the sepals; flowers mostly in compact cymes S. media.

Petals longer than the sepals; flowers solitary in the leaf axils S. ovata.

Leaves mostly acuminate or long-acuminate, often merely acute, mostly truncate

or cordate at the base, at least the lower leaves conspicuously cordate.
Sepals scarcely 2 mm. long; cymes very lax and open, the bracts inconspicuous.

S. irazuensis.
Sepals 3-6 mm. long; cymes usually few-flowered and conspicuously leafy or

bracteate.

Sepals 3-4 mm. long; petals usually little exceeding the sepals . . . S. prostrata.
Sepals 5-6 mm. long; petals usually twice as long as the sepals. S. cuspidata.

Stellaria cuspidata Willd. ex Schlecht. Ges. Naturf. Freund.
Berlin Mag. 7: 196. 1816. S. limitanea Standl. Field Mus. Bot. 22:
74. 1940 (type from Volcan de Tacana, Chiapas, E. Matuda 2775).

Moist thickets or forest or on moist shaded banks, sometimes in
dense pine forest, frequent on white-sand slopes, 1,400-4,000 meters;
Sacatepe*quez (Volcan de Agua); Chimaltenango (above Las Cal-
deras); Quezaltenango; San Marcos. Southern Mexico; Costa
Rica; Panama; western South America.

Plants prostrate or procumbent, sometimes pendent from banks or subscan-
dent on bushes, probably perennial, usually much branched, the stems very

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 237

brittle, glabrous or densely villous, the pubescence often viscid, frequently 50 cm.
long or more; leaves long-petiolate, ovate or deltoid-ovate, mostly 1-3 cm. long,
acute or acuminate, truncate to deeply cordate at the base, sometimes glabrous
but often sparsely or densely villous, especially beneath, the petioles always
villous; flowers mostly solitary in the leaf axils but sometimes cymose, the petioles
usually long and slender, often several times as long as the subtending leaves;
sepals 5-6 mm. long, ovate or lance-ovate, viscid-villosulous or glabrous except
at the base, acute; petals white, generally twice as long as the sepals and
sometimes 12 mm. long; capsule about equaling the sepals; seeds dark brown,
tuberculate.

This and the plant here called S. prostrata have been treated by
the senior author in various publications as S. nemorum L., a species
of Europe. That apparently has been introduced as a weed in
some localities of the South American Andes, but the native Ameri-
can plants are now believed to be specifically distinct from the
European species. There is some question whether S. cuspidata
and S. prostrata are distinct species but their characters are fairly
well marked, and more ample collections may strengthen the dif-
ferential characters. When described, S. limitanea was thought to
be distinct, but it is now believed that it is only an exceptionally
densely pubescent form of S. cuspidata. The pubescence in this
group is so variable in quality and density that it probably is of little
value for separating species.

Stellaria irazuensis Bonn. Smith, Bot. Gaz. 23: 236. 1897
(type from Volcan de Irazu, Costa Rica).

Dense moist forest, sometimes in Juniperus, Cupressus, or Abies
forest, found also as a weed in a wheat field, 1,500-3,500 meters;
Chiquimula; Jalapa; Chimaltenango; Solola; Huehuetenango; Que-
zaltenango (Volcan de Santo Tomas). Costa Rica.

Plants prostrate or spreading, as much as a meter long but usually smaller,
usually much branched, brittle, the stems glabrous; leaves slightly fleshy, pale
green, long-petiolate, deltoid-ovate or broadly ovate, mostly 1-2 cm. long, acute
to long-acuminate, mostly truncate or cordate at the base, glabrous, the petioles
sparsely villous; peduncles terminal, the inflorescence laxly much branched, mostly
10-20 cm. long, many-flowered, the branches glabrous or sparsely villosulous, the
leaves bract-like, small and inconspicuous, the flowers slender-pedicellate, 4-
parted; sepals scarcely 2 mm. long, oblong-elliptic, obtuse, glabrous or nearly so;
petals 2-parted almost to the base, mostly shorter than the sepals; stamens 4;
capsule 4-valvate; seeds reniform, red, puncticulate.

Stellaria media (L.) Villar, Hist. PL Dauph. 3: 615. 1789.
Alsine media L. Sp. PI. 272. 1753. Pelitaria.

238 FIELDIANA: BOTANY, VOLUME 24

Moist fields or banks, often in moist thickets, frequently a weed
in cultivated or waste ground, sometimes on rock walls, 1,500-3,300
meters; Guatemala; Sacatepe*quez ; Chimaltenango; Huehuetenango;
Quezaltenango; San Marcos. Native of Europe and Asia, but widely
naturalized in North and South America; unknown elsewhere in
Central America.

Plants annual, weak and somewhat flaccid, usually much branched, ascending
to prostrate, the stems mostly 30 cm. long or less, glabrous except for a line of
hairs along the stems; leaves ovate or oval, mostly 1-2 cm. long, obtuse or acute,
rounded to acute at the base, usually glabrous, the petioles often villosulous, the
upper leaves mostly sessile, the lower ones on rather long petioles; flowers in
generally rather dense, leafy cymes, the pedicels puberulent, little longer than the
calyx; sepals lance-oblong, 3-3.5 mm. long, pubescent, acute; petals 2-parted,
shorter than the sepals; stamens 2-10; capsule ovoid, slightly longer than the
calyx; seeds roughened and sometimes cristate.

An abundant weed in some localities of Guatemala.

Stellaria ovata Willd. ex Schlecht. Ges. Naturf. Freund. Berlin
Mag. 7: 196. 1816. Tripa de potto; Culantro de monte; Cuartillera
(fide Aguilar).

Moist or wet thickets or open forest, often on open or shaded
banks or in pastures, frequently on banks along streams or lakes,
sometimes in pine-oak forest, 300-2,500 meters; Alta Verapaz;
Zacapa; Chiquimula; Santa Rosa; Escuintla; Guatemala; Sacate-
pe"quez; Chimaltenango; Suchitepe'quez ; Retalhuleu; Huehuete-
nango; Quezaltenango; San Marcos. Mexico; Costa Rica; Panama;
northern and western South America.

Plants perennial, mostly prostrate and rooting at the nodes, often much
branched, the stems often 60 cm. long, usually glabrous; leaves on rather short
petioles, broadly ovate to suborbicular, 1-4.5 cm. long, rounded or obtuse at the
apex and apiculate, rounded at the base, rather thick and firm, pale green, glabrous,
sometimes ciliate; flowers axillary, solitary, the pedicels long and slender, sparsely
villous or glabrous; sepals 5, generally villous at the base, 3-5 mm. long, green,
obtuse; petals somewhat longer than the sepals; seeds brown, tuberculate.

This has been reported from Guatemala as S. prostrata Baldw.

Stellaria prostrata Ell. Bot. S. C. & Ga. 1: 518. 1821.

Moist or wet thickets or forest, sometimes in oak, pine, or
Cupressus forest, occasionally on sandbars along streams or in
crevices of rocks, 1,200-4,000 meters; Alta Verapaz; Baja Verapaz;
Chiquimula; Escuintla; Guatemala; Sacatepe"quez ; Chimaltenango;
Huehuetenango; Quezaltenango. Southern United States; Mexico;
•Costa Rica; western South America.

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 239

Plants annual or perennial, usually much branched, generally prostrate or
procumbent, the stems 60 cm. long or less, very brittle, glabrous or villous in lines;
leaves pale green, on long slender petioles, deltoid-ovate or broadly ovate, 1.5-3
cm. long, acute to long-acuminate, generally truncate or cordate at the base,
glabrous or often sparsely villous beneath ; flowers numerous, in small leafy cymes,
the pedicels filiform, as much as 3 cm. long; sepals ovate, obtuse or subacute,
villosulous or almost glabrous, 3-4 mm. long, green; petals sometimes twice as
long as the sepals but in Central American plants usually little if at all exceeding
them; capsule ovoid, slightly longer than the calyx; seeds minutely tuberculate.

NYMPHAEACEAE. Waterlily Family

Aquatic herbs with submerged rhizomes, the flowers usually produced on
naked scapes, the stems rarely leafy; leaves usually floating on the surface of the
water, rarely emersed, often peltate, involute in bud, the submersed leaves some-
times dissected; flowers small or large and showy, floating or emersed; sepals 3-5;
petals 3-many; stamens 6-many, free and hypogynous or sometimes perigynous or
epigynous; anthers erect, the cells dehiscent by introrse or extrorse, longitudinal
slits; carpels of the ovary 3-many, free or more or less immersed in the torus and
concrete with it; stigmas distinct or adnate to the apex of the ovary and radiating;
ovules solitary and pendulous from the apex of the cell, or numerous and attached
to the walls of the cell; mature carpels indehiscent, distinct or united to form a
fleshy or pulpy fruit; seeds surrounded by an aril or by pulp, or naked, with or
without endosperm.

Eight genera, in tropical and temperate regions. The only other
genus known from Central America is the American lotus, Nelumbo
pentapetala (Walt.) Fernald, which has been collected in Lake Yojoa,
Honduras. It is a handsome plant with large rounded peltate
leaves and showy, pale yellow flowers, both flowers and leaves
usually held well above the surface of the water.

Leaves dissected into linear lobes. Flowers very small, with 3 sepals and 3 petals;

carpels of the fruit usually 3, free Cabomba.

Leaves entire or undulate-dentate.

Sepals 4; petals numerous; leaves not peltate, with a deep basal sinus; carpels

coalescent and forming a berry-like fruit Nymphaea.

Sepals 3; petals 3; leaves peltate, without a basal sinus; carpels free. . .Brasenia.

BRASENIA Schreber

Plants with slender elongate leafy stems, covered with a gelatinous substance
like most other parts of the plant; leaves alternate, oval, entire, long-petiolate,
peltate centrally, floating, palmately nerved; flowers small, axillary, purple; sepals
3; petals 3, linear; stamens 12-18, the filaments filiform; carpels 4-18, free; ovules
2-3 in each cell, pendulous from the dorsal suture; mature carpels indehiscent,
coriaceous, 1-2-seeded.

A single species is known.

240 FIELDIANA: BOTANY, VOLUME 24

Brasenia Schreberi Gmel. Syst. Veg. 1: 853. 1796. Hydro-
peltis purpurea Michx. Fl. Bor. Amer. 1: 324. pi. 29. 1803. B. pur-
purea Gasp, in Engl. & Prantl, Pflanzenfam. III. Abth. 2: 6. 1890.

Known in Guatemala only from Laguna de Carrizal, Santa Rosa,
at 1,500 meters, Heyde & Lux 3062. Canada and United States;
Mexico; British Honduras; Cuba.

Plants with slender rootstocks; leaf blades floating, 5-10 cm. long, 3.5-5 cm.
wide, rather thick; flowers 10-12 mm. broad, long-pedunculate; carpels of the
fruit oblong, 6-8 mm. long.

CABOMBA Aublet

Plants mostly submerged, the stems slender, very leafy; leaves of 2 kinds, the
submerged ones opposite or verticillate, palmately dissected into numerous capil-
lary segments; floating leaves, when present, few, alternate, and centrally peltate,
usually absent; flowers small, white or purple; sepals 3; petals 3; stamens 3-6,
the filaments slender, the anthers short, extrorse; carpels 2-4, free, the stigmas
small, terminal; ovules generally 3, pendulous; fruiting carpels coriaceous, inde-
hiscent, 2-3-seeded.

Species about 4, in tropical or subtropical America. Only one is
known definitely from Central America.

Cabomba piauhyensis Gardner in Hook. Icon. PI. 7: pi. 641.
1844. Uchul (Peten, Maya).

In quiet fresh-water lakes or ponds, 500 meters or less; Pete"n;
Alta Verapaz; Izabal. Southern Mexico; British Honduras; Hon-
duras; West Indies; South America.

Plants very slender, the stems often 50 cm. long or more, simple or branched,
densely leafy; submerged leaves opposite or sometimes verticillate, mostly long-
petiolate, 2-4 cm. wide, divided into numerous soft linear segments; floating
leaves (often absent) linear or broader, peltate; flowers purple or white, long-
pedunculate in the upper leaf axils; sepals oblong, 6 mm. long; petals oblong or
elliptic-oblong, about equaling the sepals; stamens 3; carpels short-lanceolate,
somewhat echinate.

The plant has been reported from British Honduras as C. aquatica
Aubl., but apparently incorrectly so. The numerous specimens of
the latter species now available all have conspicuous oval floating
leaves. No such leaves are found on any of the Central American
and Mexican specimens we have examined.

NYMPHAEA L. Waterlily

Reference: Henry S. Conard, The waterlilies, a monograph of the
genus Nymphaea, Carnegie Inst. Wash. Publ. 4. 1905.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 241

Mostly large and rather coarse plants from thick short rootstocks, the leaves
and flowers floating, the flowers generally large and showy; leaves long-petiolate,
cleft basally almost to the center, entire or undulate-dentate; sepals 4; petals
(passing gradually into stamens) and stamens numerous, in many series on the
receptacle; filaments petaloid, the outer ones broad and with small anthers, the
inner ones narrow, with longer anthers; carpels immersed in the fleshy receptacle,
united with it to form a many-celled semi-inferior ovary; ovules numerous, pendu-
lous from the cell walls; fruit baccate, spongious, ripening under water and ruptur-
ing or breaking irregularly; seeds immersed in pulp, with a sac-like aril open at the
apex; endosperm scant.

Species 30 or more, widely dispersed in tropical and temperate
regions of both hemispheres. No other species are known from
Central America.

Leaves thin, entire, green beneath N. blanda.

Leaves thick, dentate or undulate, usually purple-red beneath.

Leaves coarsely and deeply dentate; flowers open in daylight AT. ampla.

Leaves merely undulate; flowers open only at night N. Rudgeana.

Nymphaea ampla (Salisb.) DC. Syst. Veg. 2: 54. 1821. Cas-
talia ampla Salisb. Parad. Lond. 1: pi. 14- 1805. N. ampla var.
Plumieri Planch. Ann. Sci. Nat. III. 19: 44. 1853. Ninfa; Nohoch
naab, Nape (Pete"n, Maya).

Floating on quiet pools, 500 meters or less; Pete"n; Alta Verapaz;
Izabal; Huehuetenango. Southern Texas; Mexico; British Hon-
duras to Salvador and Panama; West Indies; South America.

Plants large and coarse, from thick rootstocks; leaves long-petiolate, thick,
suborbicular, 15-45 cm. broad, with a deep narrow sinus at the base, coarsely
sinuate-dentate or the teeth often acutish, green above, red-purple beneath and
often with purplish black blotches, the veins conspicuously elevated and reticulate;
flowers diurnal, raised above the water, 8-16 cm. broad, white; sepals oblong-
lanceolate, obtuse or subacute, green marked with purple-black lines, little if at
all broadened at the base; petals 12-21, oblong-lanceolate, obtuse; stamens 90-190,
the outer ones long-appendaged at the apex; carpels 14-23, free from one another
at the sides, the styles short, stiff, fleshy.

The Maya name "sachab" sometimes is given the plant in Yuca-
tan. The large and handsome flowers are fragrant. This plant is
common in some parts of the Central American lowlands, being
found in almost every open swamp, but it seems to be infrequent
in Guatemala.

Some imported species with blue (probably Nymphaea zan-
zibarensis Casp.) or white flowers are planted occasionally for
ornament in Guatemala, notably in pools in the Central Park of
Guatemala City.

242 FIELDIANA: BOTANY, VOLUME 24

Nymphaea blanda G. F. W. Mey. Prim. Fl. Esseq. 201. 1818.

Floating in quiet, open or shaded water, sometimes probably in
brackish pools, at or near sea level; Izabal (collected only about
Puerto Barrios, where it is found in Manicaria swamps). British
Honduras, along the Atlantic coast to Panama; northern South
America.

Plants small, arising from short thick tubers; leaves small, long-petiolate, thin,
entire, green above and beneath, the basal sinus extending to the center of the
blade, the blades mostly 5.5-11 cm. wide, the petiole covered with long septate
hairs or often glabrous; flowers opening at night, 8-9.5 cm. broad, white; sepals
3.5-4.5 cm. long, 1.5-2 cm. wide, lance-ovate, much broadened below, green;
petals about 16, the outer ones almost 4 cm. long and 1.5 cm. wide, the inner ones
smaller; stamens about 65, the largest outer ones 2.5 cm. long; carpels about 26.

The Guatemalan material is referable to var. Fenzliana (Lehm.)
Caspary, in which the petioles and peduncles are glabrous rather
than hairy. The plants about Puerto Barrios grow mostly in very
small pools in swamps under tidal influence, and at low tide the leaves
often are stranded upon the mud.

Nymphaea Rudgeana G. F. W. Mey. Prim. Fl. Esseq. 198.
1818.

Floating in lake, about 500 meters; Jutiapa (Lago de Giiija,
southeast of Asuncion Mita, Steyermark 31828). West Indies;
South America.

Plants rather large, from a thick short rootstock; leaves long-petiolate,
rounded, 15-30 cm. wide, coarsely but shallowly sinuate-dentate, the teeth
unequal and distant, green above, usually reddish brown beneath, the narrow
basal sinus extending to the center of the blade, the petioles glabrous; flowers
opening at night, 7-15 cm. broad, white; sepals oblong-ovate, 6-7 cm. long, obtuse,
green, much broadened at the base; petals 12-32, elliptic to oblong-lanceolate,
sometimes yellowish, the flowers fragrant; stamens 40-80, the outer ones not long-
appendaged; carpels 11-24, united by their sides; fruit depressed-globose, truncate
at the apex.

The plants growing at Lake Giiija were remarkable for their
long peduncles that were spirally coiled. We have not observed
coiled petioles in other Central American species, although it is not
impossible that they exist.

CERATOPHYLLACEAE. Hornwort Family

Slender branched aquatic herbs, usually submersed in the water for all or
most of their length; leaves sessile, verticillate, very numerous, finely dissected
into rather stiff lobes; flowers almost minute, monoecious, solitary and sessile in

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 243

the leaf axils; involucre 8-12-cleft; perianth none; stamens numerous, crowded
on a flat or convex receptacle; anthers sessile or nearly so, linear-oblong, the
connective produced into a fleshy appendage, this often 2-3-dentate; pistillate
flower consisting of a sessile, 1-celled, 1-ovulate ovary; ovule pendulous; style
filiform, stigmatic at the apex; fruit a small indehiscent nutlet; endosperm none;
radicle very short, the cotyledons thick, oval.

The family consists of a single genus with perhaps 2 species.

GERATOPHYLLUM L. Hornwort

Represented in Central America by a single species of almost
worldwide distribution*

Ceratophyllum demersum L. Sp. PI. 992. 1753.

In lakes or shallow pools or ponds, 1,800 meters or less; Alta
Verapaz; Izabal; Jutiapa; Guatemala; Solola. Mexico; Honduras;
widely distributed in temperate and tropical regions of both hemi-
spheres.

Plants often forming large and dense masses in the water, often a meter long
or more; leaves 6-12-verticillate, the segments almost filiform, generally 1-2.5
cm. long, rather stiff and not collapsing when removed from the water; fruit oval,
4-5 mm. long, with a slender, straight or curved, spinose beak 5-6 mm. long, smooth
and ecalcarate, or sometimes with a long basal spur on each side, or tuberculate
and with narrowly winged, spiny margins, sometimes broadly winged and without
spines.

In temperate North America this often is an abundant plant,
filling lakes and streams, but in Central America it is sporadic in
occurrence and seldom plentiful.

RANUNCULACEAE. Buttercup Family

Annual or perennial herbs, rarely woody vines, usually with acrid sap ; leaves
mostly alternate, except in Clematis, simple or compound; stipules none, but the
leaf base often clasping or sheathing; plants glabrous, or with pubescence of simple
hairs; sepals 3-15, usually caducous, often petal oid, imbricate except in Clematis;
petals as many as the sepals or more numerous, sometimes none; flowers regular or
irregular; stamens numerous, hypogynous, the anthers introrse; carpels of the
ovary numerous or rarely solitary, 1-many-ovulate, 1-celled, the ovules anatro-
pous; fruit generally of achenes or follicles; seeds without endosperm.

About 35 genera, widely distributed in both hemispheres, the
species most numerous in temperate and arctic regions; in the tropics
found principally in the higher mountains. No other genera are
known in Central America.

Plants woody vines with white flowers; leaves opposite Clematis.

Plants annual or perennial herbs; leaves alternate, or sometimes all basal.

244 FIELDIANA: BOTANY, VOLUME 24

Flowers irregular, the posterior sepal calcarate. Cultivated plants, the leaves

much dissected; flowers blue, purple, pink, or white Delphinium.

Flowers regular, none of the sepals calcarate.

Fruit capsule-like, many-seeded; flowers pale blue; cultivated plants.

Nigella.
Fruit of 1-seeded achenes; flowers not blue; native plants.

Peduncles bearing an involucre-like whorl of leaves below the flower;

petals none, the sepals petaloid, pinkish white Anemone.

Peduncles not bearing an involucre; petals present or absent.

Petals usually present, bright yellow; leaves simple or with 9 or fewer

leaflets Ranunculus.

Petals none, the flowers greenish or whitish; leaves decompound, with
very numerous leaflets Thalictrum.

ANEMONE L.

Usually erect perennial herbs; basal leaves generally long-petiolate, divided
or dissected; cauline leaves opposite or verticillate and forming a single involucre
near or remote from the pedunculate flower or flowers; sepals 4-20, commonly
colored and resembling petals; petals none; stamens numerous, shorter than the
petals; carpels of the ovary numerous, 1-ovulate, the ovule pendulous; fruit of
few or often numerous achenes, these capitate, the style persistent.

About 80 species, in temperate and arctic regions of both hemi-
spheres; in the tropics found only in the high mountains, and then
few in number. Only the following has been found in Central
America.

Anemone mexicana HBK. Nov. Gen. & Sp. 5: 33. 1821.

Dry slopes, 2,400-2,500 meters; Huehuetenango (just above
Soloma, Sierra de los Cuchumatanes, Steyermark 48448). Southern
Mexico.

Plants erect from a dense cluster of fleshy-fibrous roots, the stems often
several, very slender, 20-40 cm. high, sparsely pilose with long weak hairs, bearing
at the apex 2-3 involucral leaves and 2-5 flowers; involucral leaves sessile or short-
petiolate, 3-parted, the segments irregularly lobate and serrate; basal leaves on
very long, slender petioles, 3-foliolate, the divisions mostly 4-7 cm. long, lobate
and serrate, green and sparsely pilose above, paler beneath, sparsely long-pilose;
peduncles 5-10 cm. long; sepals usually 5 but sometimes more numerous, white
tinged with pink, petaloid, sparsely appressed-pilose outside, oval or broadly
ovate, rounded at the apex, 14-18 mm. long, finely veined; filaments glabrous;
achenes about 10, obliquely obovoid, subcompressed, glabrous; style long and
slender, 2-3 times as long as the ovary.

The species was described originally as having pubescent achenes
but in all material we have seen they are glabrous. Probably the
hairs on the receptacle were taken to be pubescence on the achenes
themselves. The single Guatemalan station is an isolated one, far

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 245

removed from the nearest locality at which the plant is known to
occur in Mexico.

Anemone japonica Sieb. & Zucc., a native of eastern China, is
grown for ornament infrequently in Guatemalan gardens, as at
Momostenango. It is a rather coarse perennial, 50 cm. high or more,
with pubescent leaves pale beneath, each stem bearing several large,
pure white flowers.

Aquilegia Skinneri Hook, was reported from Guatemala by
Hemsley. It is a Mexican species and there is no reason to suppose
that it ever has been collected in Guatemala. The Old World
columbine, Aquilegia vulgaris L., was observed in cultivation in
Guatemala City, and doubtless is found rarely in other regions of the
country.

CLEMATIS L.

Usually woody vines (all Central American species) ; leaves opposite, petiolate,
pinnately compound, with 3 or more leaflets; flowers mostly white or creamy white,
dioecious or monoecious; flowers small or large, commonly cymose-paniculate;
sepals 4-5, valvate, petaloid, spreading in an thesis; petals none; stamens numerous,
the filaments slender, elongate, glabrous or pubescent, the anthers small, short,
obtuse; carpels of the ovary numerous, each with a long slender plumose style;
fruit a head of hard achenes, these terminated by the much elongate, long-hairy,
persistent style.

Species more than 100, in tropical and temperate regions of both
hemispheres. Some, especially those of Asiatic origin, often are
grown for ornament and most of the species are showy in flower.
Only the following ones are native in Central America.

Leaflets coarsely dentate, the teeth usually numerous and large, usually very

densely tomentose or sericeous beneath C. grossa.

Leaves entire or with an occasional tooth, or sometimes finely and evenly serrate-
dentate, glabrous beneath or sparsely and inconspicuously sericeous.
Leaflets usually very lustrous beneath, most of them evenly serrate-dentate
with numerous teeth on each side, usually 5-7, the venation elevated

beneath and conspicuously reticulate C. caleoides.

Leaflets not notably lustrous, usually entire, sometimes with an occasional
large tooth, usually 3 in most of the leaves, the venation neither conspicu-
ously elevated nor reticulate beneath C. dioica.

Clematis caleoides Standl. & Steyerm. Field Mus. Bot. 23:
52. 1944.

Moist or wet, mountain forest or thickets, sometimes in Cupressus
forest, 1,400-3,800 meters; endemic, but to be expected in Chiapas;

246 FIELDIANA: BOTANY, VOLUME 24

El Progreso; Chimaltenango (type from Cerro de Tecpam, region
of Santa Elena, Standley 58732); Solola; Quiche"; Huehuetenango ;
San Marcos.

A glabrous woody vine, sometimes climbing over trees, the stems glabrous,
the younger parts sparsely short-pilose but quickly glabrate; leaflets generally 5-7,
long-petiolulate, thick-chartaceous or thin-coriaceous, lustrous, especially beneath,
ovate, mostly 6-12 cm. long and 3.5-7.5 cm. wide, acuminate or acute, broadly
rounded at the base or rather deeply cordate, denticulate or crenate-dentate
throughout, with usually numerous teeth on each side, the teeth small and often
appressed, glabrous above or pubescent on the nerves, slightly paler beneath,
sparsely pilosulous, especially on the nerves, or in age almost wholly glabrous;
flowers dioecious, laxly cymose-paniculate, white, long-pedicellate, the pedicels
laxly tomentulose; sepals elliptic or oblong-elliptic, about 8 mm. long, densely
sericeous-tomentose.

Although the plant is rather widely distributed in the highlands
of central and western Guatemala and locally plentiful, we have
found it in flower but once. The leaflets often blacken in drying.
In their numerous, small, evenly distributed teeth and almost com-
plete lack of pubescence they are very unlike those of the other
local species.

Clematis dioica L. Syst. ed. 10. 1084. 1759. Barba de viejo;
Zepit (Pete'n, Maya, fide Lundell); Barba de chivo; Corona de angel;
Crespillo (fide Aguilar) ; Chilpat (Pete'n, apparently a Nahuatl name) ;
Cabellos de Angel; Rabo de chivo; Chimecate (Escuintla); Barbilla;
Barba de venado.

Moist thickets or open forest, often in second growth, frequent
in roadside hedges, 2,250 meters or less, most common at low eleva-
tions; Pete'n; Alta Verapaz; Baja Verapaz; El Progreso; Izabal;
Zacapa; Chiquimula; Jalapa; Jutiapa; Santa Rosa; Escuintla;
Guatemala; Quiche1; Huehuetenango; San Marcos. Mexico; British
Honduras to Salvador and Panama; West Indies; South America.

A slender, large or small, woody vine, sometimes climbing over small trees,
the stems and leaves glabrous or thinly sericeous, especially on the lower leaf
surface; leaves mostly 3-foliolate, the leaflets usually ovate and 3-10 cm. long,
acute or acuminate, rounded or subcordate at the base, 3-6-nerved, long-petiolu-
late, entire or sometimes with 1-2 large coarse teeth on either side; sepals white,
oblong or elliptic, 6-9 mm. long, sericeous-tomentose; achenes 4 mm. long, sparsely
or densely pubescent, the feathery, plumose styles or "tails" 3-5 cm. long.

Called "tietie" in British Honduras, the stems doubtless used
there and elsewhere as a substitute for cordage; "cabeza de vieja"
(Chiapas); "mexnuxib" (Yucatan, Maya). In Salvador the plant
is sometimes called "hierba de mendigo." This refers to the fact

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 247

that the juice of the leaves will produce blisters and even sores on
the skin, and it is stated that the leaves are sometimes utilized for
this purpose by professional beggars, who thus make a more pitiful
appeal to the public. The fluffy seed heads sometimes are used for
stuffing cushions. A gum that exudes from the stems is utilized for
gluing pieces of wood and is said to have as good properties as the
best glue. The vines are rather handsome when in flower, and when
covered with the fluffy seed heads they are even more conspicuous.
This species is a common weedy plant through much of the Central
American lowlands. It is somewhat variable in several respects
but no one has yet discovered any practical means of separating
several species. In fact, C. dioica and C. grossa are not too well
differentiated.

Clematis grossa Benth. PL Hartweg. 33. 1840 (type from San
Bartolo, Chiapas). C. sericea HBK. ex DC. Syst. 1: 144. 1818, not
C. sericea Michx. 1803. C. polycephala Bertol. Fl. Guat. 424. 1840
(type from Volcan de Agua, Sacatepe"quez, Velasquez^ Barba de
viejo; Ichac (Soloma, Huehuetenango) ; Crespillo; Ratichuli (San
Juan Sacatepe"quez) ; Cabello de angel; Rismachi ig (Baja Verapaz);
Usmachima (Chimaltenango) ; Angel quen (Alta Verapaz); Bejuco
de crespillo (San Marcos); Biskicam (Coban, Quecchi); Tusup
(Quezaltenango) ; Colchillo (Santa Rosa) ; Bejuco de algoddn.

Moist or wet, sometimes dry thickets or forest, frequently in
hedges, 3,000 meters or less, most frequent at middle and rather
high elevations; Alta Verapaz; Baja Verapaz; Izabal; Jalapa; Santa
Rosa; Escuintla; Guatemala; Suchitepe"quez; Retalhuleu; Quezal-
tenango; San Marcos; Totonicapan; Huehuetenango. Mexico;
southward to Panama; South America.

A large or small, woody vine, the stems usually densely long-pilose or some-
what villous, at least when young; leaflets 3 or often 5, long-petiolate, membrana-
ceous, ovate or broadly ovate, mostly 4-10 cm. long, acuminate to obtuse, broadly
rounded or often cordate at the base, coarsely crenate with few crenations on each
side, often shallowly 3-lobate, pilose or glabrate above, beneath usually densely
tomentose or sericeous, sometimes merely thinly sericeous, but the pubescence
more often abundant; flowers usually dioecious, very numerous, cymose-panicu-
late, on long or short pedicels, white; sepals elliptic or oblong, 7-9 mm. long, obtuse
or rounded at the apex, sericeous-tomentose; stamens numerous, the filaments
stout, glabrous; achenes numerous, about 3 mm. long, the plumose styles densely
hairy and greatly elongate in age.

The leaves are reported to be applied sometimes as poultices to
produce blisters or local irritation. Such properties seem to be com-
mon to many or perhaps all species of the genus. The Guatemalan

248 FIELDIANA: BOTANY, VOLUME 24

plant has been reported under the name C. dioica var. brasiliana
Eichler. The proper name for it is uncertain. It is undoubtedly
Clematis grossa Benth., but since it has so wide a range, extending
far southward in South America, it is likely that an earlier specific
name can be found for it when once the tropical American species
are properly monographed.

DELPHINIUM L. Larkspur

Erect, usually branched, annual or perennial herbs; flowers showy, racemose or
paniculate, mostly blue, purple, pink, or white; leaves palmately lobed or divided;
sepals 5, the posterior one prolonged into a spur; petals 2 or 4, small, the 2 posterior
ones calcarate, the lateral ones, when present, small; carpels of the ovary few,
sessile, many-ovulate, follicular at maturity.

Species perhaps 150, in the north temperate zone. Several are
native in Mexico but none in Central America. A large number are
found in temperate North America.

Spur equaling or longer than the petals; flowers violet, pink, or white; floral bracts
short, not exceeding the pedicels D. Ajacis.

Spur shorter than the petals; flowers always violet; floral bracts conspicuous,
long, foliaceous, equaling the flowers D. orientale.

Delphinium Ajacis L. Sp. PI. 531. 1753. Espuela; Espuela de
caballero.

• Native of southern Europe but grown for ornament in many
other parts of the earth; planted frequently in the gardens of Guate-
mala, at almost all elevations, sometimes escaping from cultivation
to cornfields and other cultivated ground, as in the mountains of
Quezaltenango.

An erect annual, usually 75 cm. high or less, sparsely or much branched, finely
pubescent; leaves much dissected into narrowly linear segments; lower leaves
petiolate, the upper ones sessile or nearly so; racemes short or elongate, often
25 cm. long, many-flowered, the flowers pedicellate, blue, violet, white, or pink;
spur slender, somewhat curved; follicle only 1, erect, pubescent, rostrate.

This is one of the commonest flowers of Guatemala, in gardens
pf rich and poor alike. Large bunches of the blossoms are common
in most of the larger markets, and in the Coban market, at least at
some seasons, it is the commonest of all flowers. It is much used in
the Easter processions of Coban, especially tied about the lighted
candles. In Guatemala the leaves crushed in water are employed
to relieve toothache and also to kill head lice. Some Delphinium
species native in the western United States have been found poison-
ous to stock, and they contain several toxic alkaloids, which may

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 249

well explain the local medicinal applications of the cultivated
larkspur.

Delphinium orientale J. Gay in Desm. Cat. PL Dord. 12. 1840.
Espuela; Espuela de caballero.

Native of western Asia; grown for ornament in many other
regions of the earth; grown frequently in Guatemalan gardens, but
not distinguished ordinarily from D. Ajacis.

An erect annual similar to the preceding, the pubescence sparse and crisp;
racemes long and dense, the flowers always intense violet or purple; sepals broader
than in D. Ajacis, the spur shorter than the petals; follicle 1, cylindric, abruptly
mucronate at the apex.

There is grown also in Guatemalan gardens a tall perennial
Delphinium with long dense spikes of light blue flowers. Specimens
of it have not been obtained, and the species name is uncertain, but
it is one of the species cultivated commonly in the United States.
The flowers often are on sale in the markets of Guatemala and Coban
but were not noted elsewhere.

NIGELLA L.

Erect annuals; cauline leaves alternate, subpinnately dissected into filiform
segments; flowers whitish, bluish, or yellowish, sometimes falsely involucrate by
the sessile floral leaves; sepals 5, regular, petaloid, deciduous; petals 5, unguiculate,
the blade small, 2-fid; carpels of the ovary 3-10, sessile, more or less connate,
several-ovulate, dehiscent at maturity at the apex; seeds angulate, the testa
crustaceous or subcarnose, usually granulate.

About 10 species, in the Mediterranean region and western Asia.

Nigella damascena L. Sp. PI. 584. 1753. Estrella del mar.

Native of the Mediterranean region; often grown for ornament
in other parts of the earth; a rather frequent ornamental plant in
gardens and parks of the Guatemalan highlands.

Plants glabrous, erect, more or less branched, usually 40 cm. high or less;
leaves dissected into numerous filiform soft segments, the solitary terminal flowers
each surrounded by a whorl of dissected leaves; sepals ovate-oblong, mucronate;
petals subsessile, pale blue; capsule membranaceous, ovate, smooth, 1.5-2 cm.
long, tipped by the erect-spreading styles; seeds triquetrous, transversely corrugate.

RANUNCULUS L. Buttercup

Usually perennial herbs, rarely annuals; leaves entire or dissected; flowers
usually bright yellow, small or medium-sized, terminal, solitary or paniculate;

250 FIELDIANA: BOTANY, VOLUME 24

sepals 3-5, caducous; petals as many as the sepals or numerous, with a nectarifer-
ous pit or scale at the base; stamens shorter than the sepals and petals, generally
numerous; carpels of the ovary numerous, 1-ovulate, the ovule ascending from the
base of the cell; achenes capitate or in short spikes, apiculate by the persistent
style or often long-rostrate, compressed or subglobose, smooth or variously
roughened.

Perhaps 200 species, chiefly in temperate and arctic regions, in
the tropics found only in the mountains. One or two other species
are found in southern Central America. Most of our Guatemalan
collections have been determined by Dr. Lyman Benson.

Leaves compound.

Stems usually glabrous; divisions of the leaves linear or nearly so.R. dichotomus.
Stems pilose or hirsute; divisions of the leaves much broader than linear.
Basal leaves pinnately compound, the terminal segment long-stalked, the

lateral ones mostly sessile or nearly so R. geoides.

Basal leaves ternate, the divisions all long-stalked, often ternate. . .R. pilosus.
Leaves simple.

Leaves densely hispidulous beneath; plants acaulescent, the peduncles usually

not exceeding the leaves, 1-flowered; petals minute R. Donianus.

Leaves glabrous or nearly so (if not, the petals large and conspicuous) ; plants
usually with well-developed stems or stolons, the stems often several-
flowered; petals usually well developed and conspicuous.
Leaves much longer than wide, ovate to lanceolate or linear-lanceolate,
entire or nearly so, obtuse to attenuate at the base . . . R. hydrocharoides.
Leaves all as broad as long or nearly so, often dentate or crenate, most of

them cordate at the base.

Leaves all entire or nearly so; plants with elongate stolons. R. flagelliformis.
Leaves coarsely dentate or crenate; plants without stolons. . .R. peruvianus.

Ranunculus dichotomus Mocifio & Sess£ ex DC. Reg. Veg.
Syst. 1: 288. 1818.

Wet meadows or in muddy places, sometimes a weed in cultivated
ground, 1,500-2,400 meters; Sacatepe"quez ; Chimaltenango; Totoni-
capan; Quezaltenango. Mexico.

Perennial from a cluster of fleshy-fibrous roots, the stems erect or decumbent,
usually glabrous, mostly 35 cm. long or shorter, 1-few-flowered; basal leaves often
numerous and large, long-petiolate, as much as 30 cm. long, pinnately 3-5-foliolate,
the segments dissected into linear lobes, pilose beneath ; flowers bright yellow, long-
pedunculate; sepals appressed-pilose, reflexed in age; petals usually 5, oval or
rounded-obovate, rounded at the apex, 10-13 mm. long, conspicuously veined;
fruit heads subglobose, 7-10 mm. broad; achenes long-rostrate, glabrous, strongly
compressed.

Ranunculus Donianus Pritzel in Walp. Repert. 2: 740. 1843.
R. humilis G. Don ex Walp. Repert. 1: 44. 1842, not R. humilis
Pers. 1807.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 251

Open rocky ridges, with Pinus and Juniperus, 2,600-3,800
meters; Huehuetenango (Sierra de los Cuchumatanes). Mountains
of central Mexico.

Perennial from a cluster of very thick, fleshy roots, acaulescent, the plants
mostly 4 cm. high or less; radical leaves few or numerous, long-petiolate, the
petioles with dilated basal sheaths, pilose above with ascending or appressed hairs;
leaf blades ovate to rounded-ovate, 5-12 mm. long, obtuse, obtuse to truncate
at the base, shallowly crenate or sublobate, hispidulous on both surfaces, often
very densely so beneath, the hairs often appressed; peduncles naked or sometimes
with a few reduced bractlike leaves, scarcely if at all exceeding the leaves, usually
1-flowered; sepals small, appressed-pilose; petals minute or none, yellow; fruit
heads subglobose, 3 mm. in diameter; achenes few, turgid, glabrous, apiculate.

Ranunculus flagelliformis J. E. Smith in Rees, Cycl. no. 13.
1819.

Swampy meadows, sometimes floating in shallow open pools,
1,350-3,000 meters; Jalapa; Chimaltenango; San Marcos. Central
Mexico; Costa Rica; western South America.

Plants perennial, glabrous throughout, the stems weak, usually creeping and
rooting at the nodes, very slender and somewhat succulent; leaves long-petiolate,
cordate-orbicular or reniform-orbicular, mostly 5-15 mm. broad, rounded at the
apex, shallowly or deeply cordate at the base, entire or nearly so; peduncles
opposite the leaves, the flowers white, 2-4 mm. broad; petals 2-3, minute; achenes
few, apiculate, turgid, the fruiting heads about 3 mm. in diameter.

Ranunculus geoides HBK. Nov. Gen. & Sp. 5: 47. 1821.
Hierba de pozo (fide Aguilar).

Mostly in moist or wet, open meadows, chiefly in alpine situa-
tions, rarely a weed in cultivated ground, 2,500-4,000 meters;
Chimaltenango; Solola; Totonicapan; Huehuetenango; Quezalte-
nango. Mexico.

Perennial from a cluster of fleshy-fibrous roots, the stems elongate, ascending
or procumbent, mostly 35 cm. long or less, sparsely or densely hirsute or pilose with
appressed or ascending hairs; basal leaves few or numerous, long-petiolate, abund-
antly hirsute or pilose with appressed or spreading hairs, sometimes glabrate,
pinnately compound, usually 5-foliolate but many of the upper leaves 3-foliolate,
the terminal segment long-stalked, the lateral ones sessile or nearly so, shallowly
or deeply lobate or crenate; stems 1-several-flowered, the flowers bright yellow,
long-pedunculate; sepals appressed-pilose, less than half as long as the petals;
petals usually about 10, oblong or cuneate-oblong, 6-10 mm. long; fruit heads
subglobose, about 1 cm. broad; achenes numerous, glabrous, slender-rostrate,
compressed.

Ranunculus hydrocharoides Gray, Mem. Amer. Acad. II. 5:
306. 1855. Sanguijuela.

252 FIELDIANA: BOTANY, VOLUME 24

In shallow pools in alpine meadows, 3,180-3,500 meters; Hue-
huetenango (Sierra de los Cuchumatanes). Southern and central
Mexico.

Perennial from a cluster of fleshy-fibrous roots, glabrous throughout, the stems
erect, simple, thick and somewhat fistulous; leaves all on very long, spongy petioles,
the basal ones with mostly ovate blades 1-4 cm. long, narrowed to an obtuse apex,
rounded to acute at the base; cauline leaves few, petiolate, lanceolate or linear-
lanceolate, entire or nearly so; flowers few, yellow, long-pedunculate but the stout
peduncles usually shorter than the subtending leaves; sepals oblong-elliptic, 2-3
mm. long; petals 5-6, oblong-elliptic, 2.5-3 mm. long; heads of achenes 3-4 mm.
in diameter, ovoid-globose; achenes glabrous, somewhat compressed, short-
rostrate.

In the Cuchumatanes there is a belief that if stock eat this plant,
the liver is affected and the animals die. There is probably no true
basis for this belief. The Guatemalan material is referable to the
aquatic form of the species with elongate, often floating stolons,
R. hydrocharoides var. natans (Nees) Benson.

Ranunculus peruvianus Pers. Syn. PL 2: 103. 1807. R.
Salasii Standl. Field Mus. Bot. 11: 154. 1936 (type from El Choi,
Sierra de los Cuchumatanes, Huehuetenango, J. Garcia Salas 1410).

Moist or wet, alpine meadows, often about the margins of pools,
3,100-3,700 meters; Totonicapan; Huehuetenango. Mexico; Costa
Rica; mountains of western South America.

Perennial from a cluster of fleshy-fibrous roots, the stems erect or ascending,
mostly 30 cm. long or less, glabrous or sparsely pilose, especially at the nodes,
1-few-flowered, the flowers long-pedunculate, bright yellow; cauline leaves bract-
like, divided into linear segments, 1-2 bracts on each stem; basal leaves few or
numerous, on very long, slender, usually glabrous petioles, the shredded bases of
the petioles persisting as a cluster of fiber^ at the summit of the rootstock; leaf
blades 1-5 cm. wide, orbicular or reniform in outline, shallowly or deeply and
narrowly cordate at the base, evenly and deeply dentate with numerous, narrowly
to broadly triangular or ovate-triangular teeth, glabrous, or rarely pilose beneath
or at the base of the blade; peduncles very long and slender, appressed-pilose
above; sepals broadly elliptic or suborbicular, sparsely appressed-pilose; petals 5,
suborbicular, 6-10 mm. long, broadly rounded at the apex.

Part of the Guatemalan material is treated by Benson as a local
variety of the species, R. peruvianus var. Salasii (Standl.) Benson,
but it is now believed by the senior author that R. Salasii does not
deserve any special nomenclatorial designation, and it is more
practical and sensible to treat all the Guatemalan specimens as
R. peruvianus.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 253

Ranunculus pilosus HBK. ex DC. Reg. Veg. Syst. 1: 287. 1818.
R. Amarillo Bertol. Fl. Guat. 424. 1840 (type from Guatemala,
Vel&squez). Gengibre (Baja Verapaz fide Garcia Salas); Hierba de
pozo (fide Aguilar); Asuchel (Huehuetenango).

Moist or wet meadows, thickets, or open forest, often on open
banks, sometimes in oak forest, 1,200-2,100 meters; Alta Verapaz;
Baja Verapaz; Zacapa; Jalapa; Guatemala; Chimaltenango; Solola;
Quich^ ; Huehuetenango; Totonicapan. Mexico; Costa Rica;
western South America.

Perennial, from dense clusters of fleshy-fibrous roots, the stems erect and as
much as 75 cm. high, sometimes procumbent or prostrate and rooting at the lower
nodes, stout or slender, usually densely pilose or hirsute with spreading or ascend-
ing hairs; cauline leaves numerous, the basal leaves long-petiolate, ternate, the
segments all long-stalked, the segments ternate or deeply lobate, coarsely dentate
or again lobate, hirsute or appressed-pilose; stems usually several-flowered, often
many-flowered, the flowers bright yellow; sepals broad, pilose or hirsute; petals
often about 10, sometimes fewer, oblong or obovate, 6-10 mm. long, rounded at
the apex; fruit heads globose, 1 cm. broad; achenes numerous, glabrous, slender-
rostrate, compressed.

There is sometimes seen in Guatemalan gardens, as an ornamental
plant, a Ranunculus with rather large, very double, bright yellow
flowers. This is probably R. repens L. var. flore-pleno DC., a plant
of Old World origin, occasionally found in gardens of the United

States.

THALICTRUM L.

Reference: Bernard Boivin, American Thalictra and their Old
World allies, Rhodora 46: 337-377; 391-445; 453-487. 1944.

Perennial herbs, usually tall, with simple or branched, generally leafy stems,
the roots usually yellow; leaves small or large, basal and cauline, the cauline ones
alternate, with sheathing petioles, the blades ternately decompound; flowers
small, mostly polygamous, green or yellowish, sometimes purplish or whitish,
paniculate, not involucrate; sepals 4-5, petaloid; petals none; stamens long-
exserted, the anthers mostly large and conspicuous; carpels of the ovary few or
numerous, inserted on a small receptacle, 1-ovulate; ovules pendulous; fruit of
achenes, these often stipitate, not caudate, generally compressed, the sides 1-3-
nerved; style deciduous or none.

Species about 80, chiefly in temperate regions of the northern
hemisphere, in the American tropics found only in the higher
mountains. A few other species occur in southern Central America.
Most of the Guatemalan material we have studied has been examined
and in some cases determined by Mr. B. Boivin.

254 FIELDIANA: BOTANY, VOLUME 24

Leaflets peltate, the petiolule attached well above the base of the blade.

T. guatemalense.
Leaflets not peltate.

Leaflets acute to long-acuminate T. Standleyi.

Leaflets rounded or very obtuse at the apex.

Principal leaflets 2-3 cm. long, yellowish beneath when dried, the terminal

ones usually deeply lobate T. Steyermarkii.

Principal leaflets 1-2 cm. long, not at all yellowish beneath, shallowly crenate-
lobate T. Johnstonii.

Thalictrum guatemalense C. DC. & Rose, Contr. U. S. Nat.
Herb. 5: 188. 1899 (type Heyde 164, without locality, from Guate-
mala). T. peltatum var. hirsutum Loes. Bull. Herb. Boiss. II. 3: 89.
1903 (type from Zaculeu near Huehuetenango, Huehuetenango,
Seler 3153). T. hondurense Standl. in Yuncker, Field Mus. Bot. 17:
362. 1938. Supote (Huehuetenango).

Moist to rather dry thickets and forest, most frequent in oak or
pine forest, 900-2,100 meters; Baja Verapaz; Zacapa; Jalapa;
Jutiapa; Guatemala; Sacatepe"quez ; Chimaltenango; Solola; Hue-
huetenango. Southern Mexico; Honduras.

A tall slender herb, commonly about a meter high, often much branched above,
the stems hispid or puberulent; leaves generally large, 2-4 times ternate; leaflets
numerous, mostly 1-2 cm. long, rounded or broadly ovate, very obtuse or rounded
at the apex, rounded at the base, peltately attached a short distance above the
base, thick and firm, coarsely crenate-lobate, petiolulate, usually scaberulous or
at least roughened on the upper surface, paler beneath and densely puberulent or
glandular-pubescent; inflorescence small and few-flowered or large, much branched
and many-flowered, the flowers on slender but short pedicels; anthers slender-
rostrate, the cells about 4 mm. long; achenes very oblique, broadly clavate, 4 mm.
long, short-stipitate, coarsely costate, minutely puberulent.

The plant rises from a dense cluster of rather slender but fleshy,
bright yellow roots. Similar roots are found in the other local
species of Thalictrum. Plants reported from Guatemala as T. pel-
tatum DC. belong to this species, and probably also the Guatemalan
records of T. strigillosum Hemsl. and T. lanatum Lecoyer.

Thalictrum Johnstonii Standl. & Steyerm. Field Mus. Bot.
22: 229. 1940. Culantrillo de zorra (fide Aguilar).

Moist or wet, mixed, mountain forest, 2,100-3,000 meters;
endemic; El Progreso; Solola; Totonicapan (type from Desconsuelo,
John R. Johnston 1643); Quiche"; Huehuetenango (?).

An erect glabrous perennial herb commonly 1-1.5 meters high, the stems
slender, simple or branched; cauline leaves long-petiolate, the leaflets numerous,
epeltate, membranaceous, slender-petiolate, suborbicular or irregularly rhombic,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 255

mostly 1-1.5 cm. long, shallowly 2-3-lobate, with very obtuse or rounded, apicu-
late lobes, green and glabrous on the upper surface, scarcely paler beneath, glabrous
but sparsely and very minutely glandular; flowers dioecious, laxly paniculate, the
panicles mostly small and few-flowered, the pedicels almost filiform, elongate;
sepals purplish, oval or broadly elliptic, 2.5-3 mm. long; filaments about 7 mm.
long, the anthers linear, 2 mm. long; achenes unknown.

Guatemalan records of T. Galeottii Lecoyer are referable to this
species.

Thalictrum Standleyi Steyerm. Field Mus. Bot. 22: 229. 1940.

Moist or wet, mixed forest, sometimes in Abies forest, often on
wooded slopes of loose white sand, 2,500-3,000 meters; endemic;
Huehuetenango (Sierra de los Cuchumatanes) ; Quezaltenango; San
Marcos (type from Rio Vega, near San Rafael and the Mexican
boundary, Volcan de Tacana, Steyermark 36258).

An erect herb 1-2.5 meters tall, sometimes rather weak and supported on other
vegetation, the stems somewhat fistulous, striate, sparsely villous with lax hairs,
more densely villous at the nodes; leaves large, decompound, on short or elongate
petioles, the petiolules very unequal, 1-6 cm. long; leaflets numerous, large, thick-
membranaceous, epeltate, ovate or broadly ovate, mostly 4-10 cm. long and 2.5-
6.5 cm. wide, acuminate or long-acuminate, deeply cordate at the base or some-
times merely truncate, deeply and coarsely crenate, rarely somewhat 3-lobate,
the crenations sometimes again crenate or shallowly 3-lobate, deep green and
glabrous above, usually lustrous when dried, the nerves and veins prominent,
paler beneath, almost glabrous but bearing a few small gland-tipped hairs at
least on the nerves near the base of the blade, the nerves and veins elevated and
closely reticulate; flowers polygamous-monoecious, rather large, in large lax
leafy-bracteate panicles; sepals broadly ovate, obtuse, 6 mm. long, sparsely viscid-
villosulous or almost glabrous; stamens numerous, the slender filaments 5 mm.
long or more, the anthers linear, 4.5 mm. long, subulate-apiculate; young achenes
strongly asymmetric, substipitate, obliquely rostrate, the filiform style 1 cm. long
or more.

This is one of the most distinct members of the genus, easily
recognized by the leaflets alone, which are noteworthy for their
combination of large size, deeply cordate bases, acuminate or long-
acuminate apices, coarsely crenate or doubly crenate margins, and
elevated reticulate venation. The foliage suggests that of some
species of Clematis.

Thalictrum Steyermarkii Standl. Field Mus. Bot. 22: 230.
1940.

Moist or wet thickets or forest, sometimes in Alnus forest, often
on wooded stream banks, 1,800-2,600 meters; endemic; Quezalte-

256 FIELDIANA: BOTANY, VOLUME 24

nango; San Marcos (type from Volcan de Tajumulco, barrancos
south and west of town of Tajumulco, Steyermark 36575).

An erect perennial herb about 2.5 meters high, almost glabrous, the stems
thick, somewhat fistulous; leaves very large, pinnately decompound, long-petiolate;
leaflets very numerous, the terminal ones on slender petiolules as much as 2 cm.
long, the lateral ones on shorter petiolules, membranaceous, broadly oblong to
broadly ovate or cuneate-obovate, mostly 2-3 cm. long and 1-2 cm. wide, obtuse
and apiculate, rounded or truncate at the base, sometimes entire but usually
shallowly or deeply 3-lobate, deep green on the upper surface and glabrous or
very minutely granular-puberulent, slightly paler beneath, yellowish green, at
least when dry, glabrous or sparsely and almost microscopically puberulent, the
nerves and veins very slender, prominent, laxly reticulate; flowers apparently
dioecious, forming a large lax many-flowered leafy panicle, the pedicels capillary,
greatly elongate and mostly 4-6 cm. long; follicles strongly asymmetric, sessile,
about 6 mm. long and 2.5-3 mm. broad, acute at the base, apically attenuate into
a style as much as 9 mm. long, minutely puberulent or almost glabrous, coarsely
costate.

This plant is noteworthy for its greatly elongate, almost capillary
pedicels. We refer to this species one collection determined by
Boivin as T. Hintonii Boivin, a Mexican species. The specimen is
sterile and is obviously a small plant of T. Steyermarkii.

BERBERIDACEAE. Barberry Family

Herbs, shrubs, or small trees; leaves alternate, simple or compound, the
petioles dilated at the base or stipulate; flowers perfect, solitary or in racemes,
cymes, or panicles; sepals and petals imbricate, usually in whorls of 3, rarely of
2 or 4; stamens free, as many as the petals and opposite them, the filaments short,
the anthers opening by 2 valves or rarely by longitudinal slits; ovary superior,
1-carpellate; ovules few to many, rarely only 1, borne on the ventral surface of
the cell or at its base; style short or none, the stigma usually peltate; fruit baccate
or follicular; seeds anatropous, with endosperm; embryo usually small, straight.

Ten genera, in the northern hemisphere, only Berberis extending
southward to the Straits of Magellan, along the Andes. One other
genus (Berberis) is represented by one species in Central America,
in the high mountains of Costa Rica and Panama.

MAHONIA Nuttall

Reference: Friederich Fedde, Versuch einer Monographic der
Gattung Mahonia, Bot. Jahrb. 31: 30-133. 1901.

Unarmed shrubs or small trees; leaves persistent, coriaceous, alternate, odd-
pinnate, rarely 3-foliolate, the leaflets often spinose-dentate, the lateral ones sessile;
stipules minute, subulate; flowers small, yellow, in many-flowered racemes or
panicles springing from the axes of bud scales; sepals 9; petals 6; ovary commonly

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 257

few-ovulate; fruit baccate, dark blue, usually with a glaucous bloom, rarely red or
whitish.

About 50 species in North America and eastern and central
Asia. No other species are known from Central America. Most
species of the genus are said to be susceptible to black stem rust or
wheat rust of cereals, and therefore are a dangerous pest in the
vicinity of grain fields. The fruit is acid and edible. By many
authors the genus has been combined with Berberis.

Leaflets entire M. Johnsionii.

Leaflets spinose-dentate.

Leaflets 5-13, rounded or very obtuse at the apex; flowers in short racemes

mostly much shorter than the leaves M. volcania.

Leaflets mostly 15-17, attenuate-acuminate; flowers in large lax panicles, these
often as long as the leaves M. paniculata.

Mahonia Johnstonii Standl. & Steyerm. Field Mus. Bot. 23:
6. 1943. Berberis Johnstonii Standl. & Steyerm. op. cit. 22: 140. 1940.

Dry, brushy, often rocky hillsides, 1,300-1,650 meters; endemic;
Baja Verapaz (Santa Rosa); Zacapa (Sierra de las Minas); Saca-
tepe"quez (type collected near Parramos, John R. Johnston 1525).

A shrub 2-6 meters high, glabrous, sparsely branched; leaves large, the petiole
and rachis slender, naked; leaflets 5-9, usually 7, coriaceous, sessile, entire or
subundulate, elliptic-oblong, sometimes oblong or oval-oblong, mostly 3-5 cm.
long and 1.5-2.5 cm. wide, rounded or very obtuse at the apex, sometimes retuse,
broadly cuneate at the base, lustrous above, paler beneath, the nerves and veins
prominulous and closely reticulate; inflorescence and fruit unknown.

Apparently of very local distribution, for we have found it in
only three widely separated localities.

Mahonia paniculata Oerst. Vid. Medd. Kjoebenhavn 1856:
36. 1857. Berberis paniculata Hemsl. Biol. Centr. Amer. Bot. 1 : 24.
1879. B. Hemsleyi Bonn. Smith in Pittier, Prim. Fl. Costar. 2:
17. 1898. Yema de huevo; Anchix (Huehuetenango).

Moist forest, 1,800-2,800 meters; Zacapa; Guatemala (Volcan
de Pacaya); Huehuetenango. Volcan de Irazu, Costa Rica.

A glabrous shrub or tree 1.5-9 meters high, slender, with few branches, some-
times epiphytic; leaves large, the slender rachis naked, the petioles short; leaflets
mostly 13-17, oblong-lanceolate, 5-9 cm. long, attenuate-acuminate, truncate or
rounded at the base, subsessile, coriaceous, spinose-serrate with bristle-tipped
teeth, somewhat lustrous above, paler beneath, the veins prominulous and laxly
reticulate; panicles larger than in most species of the genus, about equaling the
leaves, copiously branched and many-flowered, the bracts rather conspicuous;
flowers yellow, 7 mm. long, slender-pedicellate; berry 3-seeded.

258 FIELD IANA: BOTANY, VOLUME 24

The name "yema de huevo" (egg yolk) alludes to the bright
yellow wood, characteristic of this genus. That of this species is
used in Huehuetenango and perhaps elsewhere for imparting a yellow
dye to sacks, petates, and other articles.

Mahonia volcania Standl. & Steyerm. Field Mus. Bot. 23:
6. 1943.

Moist or wet, often dense, pine or coniferous forest, sometimes
in Juniperus forest, 3,000-3,700 meters, sometimes on limestone;
endemic; Sacatepe"quez (type from Volcan de Agua, 3,000 meters,
Standley 65221); Huehuetenango (Che'mal).

A slender glabrous shrub 1-5 meters high with few branches; leaves short-
petiolate, the leaflets 5-13, close together, rigid-coriaceous, sessile, oval to oblong-
oval or broadly oblong, 2-4 cm. long, 10-17 mm. wide, rounded or broadly obtuse
at the apex and spine-tipped, rounded or broadly obtuse at the base, appressed-
spinose-serrate along the whole margin, lustrous above, the costa impressed, the
veins prominulous, pale, closely reticulate, pale beneath; flowers bright yellow,
racemose, the racemes dense and many-flowered, 3 cm. long, the slender but rigid
pedicels 10 mm. long or less; outermost sepals broadly ovate, 2 mm. long; petals
6-7 mm. long.

MENISPERMACEAE. Moonseed Family

Reference: L. Diels, Menispermaceae, Pflanzenreich IV. 94. 1910.

Mostly scandent shrubs, sometimes erect trees or shrubs, without tendrils;
leaves alternate, without stipules, petiolate, penninerved or usually palmate-
nerved, entire or palmate-lobate, the petiole articulate at the base and often also
at the apex; flowers dioecious, in small cymes, these racemose or paniculate, the
flowers small, greenish, whitish, or yellowish; sepals variable in number, usually in
whorls of 3, free or rarely coalescent, imbricate or valvate, the outer ones usually
smaller than the inner; petals commonly in 2 series of 3, sometimes reduced to 1 or
none, usually free, imbricate or valvate; stamens numerous or as many as the
petals and opposite them, often 3 or 6, free or variously connate; carpels usually 3,
sometimes 6 or more, inserted on a short torus or rarely on an elongate gynophore,
free; styles terminal or subterminal, commonly recurved, the stigmas entire, lobate,
or cleft; ovules usually 2 at first but soon reduced to 1, amphitropous, affixed to the
ventral suture; fruits drupaceous, the carpels free, sessile or stipitate, the exocarp
membranaceous or subcoriaceous, the mesocarp more or less fleshy; endocarp
chartaceous or osseous, usually rugose, tuberculate, or variously costate; seed
often hippocrepiform, the testa membranaceous; endosperm copious, scant, or
none, ruminate or continuous; embryo usually curved, the radicle minute; cotyle-
dons pale and foliaceous or thick and semiterete.

Diels recognizes 63 genera, which are widely distributed, mostly
in tropical regions. Two genera besides those treated here are
represented in southern Central America, and two others have
Mexican species.

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 259

Sepals 4; anthers dehiscent by transverse slits. Plants scandent; leaves often or
usually peltate, commonly almost as broad as long or broader; endosperm

present Cissampelos.

Sepals 6; anthers opening by longitudinal slits. Leaves conspicuously longer than
broad.

Leaves conspicuously peltate; vines; endosperm present Disciphania.

Leaves not peltate or obscurely so; vines or trees; endosperm present or absent.

Endosperm present in the seed; petals none Abuta.

Endosperm none; petals present '. . .Hyperbaena.

ABUTA Aublet

Large woody vines, glabrous or often densely pubescent; leaves coriaceous or
thinner, usually long-pedicellate, entire, generally palmate-nerved; staminate
inflorescence usually paniculately compound, the pistillate flowers racemose or
spicate; staminate sepals 6, the outer 3 bract-like, the 3 inner ones larger, generally
pubescent outside and sometimes also within; petals rarely few and minute, com-
monly none; stamens 6, connate at the base or free, the anthers extrorse, introrse,
or lateral; carpels of the ovary 3, the stigmas sessile, simple or 2-fid, subulate,
recurved; drupes short-stipitate or attenuate at the base, ovoid; endocarp with a
septiform condyle above the middle; seed induplicate above the condyle and hip-
pocrepiform; endosperm ruminate; embryo hippocrepiform, the cotyledons accum-
bent, equal.

About 15 species, all except the following in South America.

Abuta Steyermarkii Standl. Field Mus. Bot. 23: 156. 1944.
Hyperbaena Steyermarkii Standl. op. cit. 22: 232. 1940.

Moist or wet, mixed forest, 900 meters or less; endemic; Alta
Verapaz (between Chirriact£ and Semococh) ; Izabal (type collected
along Rio Dulce above Livingston, Steyermark 39454).

A large woody vine, the stems terete, densely pilose-tomentose with long,
ochraceous or fulvescent hairs; leaves firm-coriaceous, on slender petioles 2-9 cm.
long; leaf blades suborbicular to broadly elliptic or oblong-elliptic, 9-16 cm. long,
4-14 cm. wide, rounded at the apex and shortly cuspidate-acuminate, or gradually
or abruptly acuminate, narrowly rounded to subcordate at the base, more or less
pilose above, at least along the nerves, usually brownish beneath, densely velutin-
ous-pilose, 5-nerved from the base; pistillate flowers spicate, the spikes usually
dense, few-many-flowered, sessile, axillary, 5 cm. long or less, the rachis densely
pilose, the flowers closely sessile; inner sepals rounded-ovate, 3 mm. long, obtuse,
densely tomentose outside, glabrous within; carpels of the ovary densely tomentose.

In general appearance this species resembles the South Ameri-
can A. rufescens Aubl., but in that the pistillate flowers are long-
pedicellate.

CISSAMPELOS L.

Scandent shrubs (in Central America), rarely erect herbs or shrubs; leaves on
long slender petioles, all or most of them peltate in Guatemalan species, glabrous

260 FIELDIANA: BOTANY, VOLUME 24

or pilose, mostly ovate-rounded, orbicular, or rounded-cordate; staminate inflores-
cences axillary, the cymes many-flowered, with very slender branches; pistillate
cymes mostly simple and few-flowered, in the axils of leaves or bracts, the bracts
often accrescent; staminate sepals 4, usually pilose, obovate; petals connate to
form a patelliform or cupular corolla, rarely 2-4 and free; stamens connate to form
a column; pistillate sepal 1, obovate, pilose dorsally; petal 1, shorter than the
sepal; carpel 1, villous; drupes usually pilose, with juicy fleshy epicarp; endocarp
crustaceous-osseous, costate dorsally and with transverse costules.

Species 20, generally dispersed in tropical regions. Two other
species are known from southern Central America.

Stems thinly pilose with very long and slender, spreading hairs; leaves peltate,
very sparsely pilose or sometimes almost glabrate; bracts long-ciliate, those
of the staminate inflorescence similar to the pistillate ones . . . C. tropaeoli folia.
Stems glabrate or often very densely pilose with short hairs; leaves peltate or
epeltate, often very densely pubescent; bracts not long-ciliate, those of the
staminate inflorescence often reduced or absent.

Leaves peltate or epeltate, usually copiously pubescent or tomentose; inflores-
cence of corymb-like cymes in the axils of the leaves, mostly shorter than

the leaves C. Pareira.

Leaves conspicuously peltate, sparsely short-pilose or almost glabrous; inflores-
cence of elongate panicles composed of numerous small cymes.

C. grandifolia.

Cissampelos grandifolia Triana & Planch. Ann. Sci. Nat. IV.
17: 44. 1862. Alcotdn.

Wet thickets, 1,250-2,000 meters; Alta Verapaz; Baja Verapaz;
Quezaltenango; San Marcos(?). Southern Mexico; Costa Rica;
Panama; southward to Peru.

A large or small, woody vine, the slender branches puberulent or short-pilose,
sometimes with a few longer spreading hairs, often glabrate; leaves thin, conspicu-
ously peltate, rounded-ovate to suborbicular, 5-15 cm. long and often almost as
wide, acuminate to very obtuse and mucronate, green on the upper surface,
pubescent or almost glabrous, paler beneath, thinly or densely pilose; staminate
panicles large and much branched, lax, often 15-20 cm. long, the bracts usually
much reduced, or absent, sometimes well developed, the branches short-pilose;
sepals obovate, 1-1.5 mm. long; corolla 1.5 mm. broad, green; drupes obovoid,
compressed 5-6 mm. long, tuberculate, pilose.

Called "curarina" in Veracruz, where the plant is said to be
used as a remedy for snake bites.

Cissampelos Pareira L. Sp. PI. 1031. 1753. Alcotan; Tamagds;
Curarina; Curarina de monte; Ixcatu-can (San Juan Sacatepe"quez) ;
Cuxogui, Cuxba (Quecchi) ; Guaco (fide Aguilar) ; Bejuco de la prenada,
Estrella de la prenada (Pete'n) ; Curarina (Huehuetenango) .

STANDLEY AND STEYERMARJC: FLORA OF GUATEMALA 261

Common in dry to wet thickets or forest, often in second growth,
sometimes in pine-oak forest, ascending to about 1,800 meters but
most plentiful at low elevations, chiefly below 1,000 meters; Pete"n;
Alta Verapaz; Baja Verapaz; Izabal; Zacapa; Chiquimula; Jalapa;
Santa Rosa; Escuintla; Guatemala; Suchitepe"quez ; Retalhuleu;
Huehuetenango; San Marcos; Quezaltenango; Quiche". Mexico;
British Honduras to Salvador and Panama; West Indies; South
America; Old World tropics.

A small or large vine, climbing over shrubs or small trees, the stems slender,
usually densely short-pilose or puberulent, often tomentose; leaves long-petiolate,
firm, rounded-ovate to reniform, peltate or epeltate, 3-10 cm. long, rounded and
mucronate at the apex, sometimes emarginate, broadly rounded or cordate at the
base, commonly tomentose or sericeous-tomentose but often glabrate; staminate
inflorescence corymbose, borne in the axils of normal leaves and usually shorter
than the leaves, the bracts small and inconspicuous or none, the pedicels mostly
filiform and pilose; flowers green, the sepals 1-1.5 mm. long; fruit red or orange-red,
obovoid or suborbicular, compressed, 4-5 mm. long, pilose.

"Peteltun," "tsutsuc" (Yucatan, Maya). The usual name in
Guatemala is "alcotan," and the plant is well known in most of
Central America by this name since important medicinal properties
are attributed to it. The roots are hard, tortuous, brown, and rugose,
with a bitter flavor. Throughout much of tropical America they
have a high reputation as a remedy for bites of snakes or other poison-
ous animals. Dieseldorff states that about Coban an extract of the
root is employed in treating fevers, and in Pete"n it is a domestic
remedy for erysipelas. The species is a highly variable one, which
is not unnatural considering its wide distribution.

Cissampelos tropaeolifolia DC. Reg. Veg. Syst. 1: 532. 1818.
Aspirina (Huehuetenango) ; Alcotan.

Moist or wet forest or thickets, ascending from sea level to about
2,000 meters; Izabal; Suchitepe"quez; Retalhuleu; Quezaltenango;
San Marcos; Huehuetenango. Veracruz to Oaxaca and Chiapas;
British Honduras; Honduras; Costa Rica; Panama. Western South
America.

Stems slender, thinly pilose with long, spreading, white, rather lax hairs, often
glabrate in age; leaves peltate, usually well above the base, thin, rounded-ovate to
suborbicular, mostly 5-13 cm. long, rounded to acute at the apex, mucronate,
truncate or broadly rounded at the base, somewhat paler beneath and often
glaucescent, sparsely pilose with long spreading hairs or glabrate; staminate
inflorescence cymose-paniculate, sometimes 15 cm. long, lax, the branches very
slender, usually with large green bracts similar to those of the pistillate inflores-
cence, the branches long-pilose, the flowers pale green, slender-pedicellate; sepals

262 FIELDIANA: BOTANY, VOLUME 24

1.2 mm. long; corolla scarcely 1 mm. broad; pistillate inflorescences with large,
green, cordate-orbicular or reniform bracts, these accrescent in fruit, long-ciliate;
drupes dull red, sparsely long-pilose, 6-7 mm. long.

Easily recognized by the rather sparse, very long, spreading
hairs of the stems. This species is not so "weedy" as C. Pareira,
being found chiefly in more or less primeval forest, or along its
borders, and never in such dry situations as are normal for C. Pareira.
It is used in domestic medicine likeC. Pareira, and in Huehuetenango
is administered in decoction as a remedy for colds.

DISCIPHANIA Eichler

Scandent shrubs, glabrous or pubescent; leaves variable in shape, entire or
lobate, often peltate; inflorescence spicate, simple, the bracts minute; sepals 6 in
the staminate flower, subequal, elliptic, membranaceous or carnose; petals 6,
much smaller than the sepals, carnose; stamens 3 or rarely 6, free, the filaments
short or obsolete, the anthers dehiscent by longitudinal slits; carpels 3, free, the
style very short or obsolete, the stigmas simple, discoid; drupes usually by abortion
solitary, straight, the exocarp juicy and fleshy; endocarp ligneous, more or less
compressed, with longitudinal wings or angles, these sometimes erose or fimbriate;
seed ovoid, straight, with endosperm; cotyledons foliaceous.

Eleven species, all except two Mexican ones and the following in
South America.

Disciphania calocarpa Standl. Field Mus. Bot. 4: 305. 1929
(type from Lancetilla Valley near Tela, Honduras). D. coriacea
Standl. Carnegie Inst. Wash. Publ. 461: 55. 1935 (type from Rio
Grande, British Honduras, Schipp S458).

Wet forest, often on limestone, 1,500 meters or less; Alta Verapaz
(near Chirriacte") ; Izabal; Huehuetenango. British Honduras;
Honduras; Costa Rica.

Usually a small vine with few branches, but sometimes 18 meters long and
climbing over trees, the older branches covered with thick corky ridged bark,
glabrous throughout; leaves long-petiolate, coriaceous or membranaceous, peltate,
with the petiole attached far above the base, broadly ovate to ovate-oblong or
broadly oblong, 8-17 cm. long, acute or abruptly short-acuminate with obtuse tip,
rounded at the base, palmate-nerved, blackening when dried, lustrous; flowers
short-pedicellate or subsessile, in very long and slender, usually interrupted
racemes, these pendent in fruit; fruits oval, 1.5-2 cm. long, rounded at base and
apex, turning yellow and then bright red, glabrous.

The fruits are pretty and showy, somewhat suggesting cherries,
but they are not produced in much abundance, unless exceptionally.
A decoction of the plant is used in domestic medicine in Huehue-
tenango, as a remedy for kidney diseases and as a "blood purifier."

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 263

HYPERBAENA Miers

Scan dent shrubs or sometimes erect shrubs or small trees; leaves coriaceous,
entire or angulate, palmately or pinnately nerved; staminate flowers in small
paniculate cymes with slender branches, the pistillate racemose, the bracts and
bractlets minute, pilose; staminate sepals membranaceous, glabrous or pilose, the
3 outer ones small, the 3 inner concave, imbricate; petals 6, subcarnose, obovate;
stamens 6, the filaments dilated at the apex, the cells dehiscent by vertical lateral
slits; carpels 3, free, gibbous, the style ex centric, extrorsely reflexed; drupes sessile,
the rudiment of the style near the base much curved; endosperm ligneous or
crustaceous-coriaceous; seed hippocrepiform, without endosperm; cotyledons
thick-carnose or subcorneous, semicylindric, often unequal, the radicle very short.

About 40 species, mostly in the West Indies but ranging from
southern Mexico to Brazil; 4 or 5 other species are known in
Central America.

Leaves broadest near the apex, obtuse, some or all of them shallowly 3-lobate or

3-angulate at the apex H. Winzerlingii.

Leaves broadest at or below the middle, not at all angulate or lobate.

Leaves palmately 3-5-nerved, the nerves arising from the very base of the blade;

woody vines.

Branchlets glabrous or puberulent; leaves glabrate or glabrous, the hairs
mostly confined to the nerves of the lower surface.

Pistillate pedicels short, 2-4 mm. long H. hondurensis.

Pistillate pedicels 6-15 mm. long H. vulcania.

Branchlets densely pilose with spreading hairs or tomentose; leaves densely

short-pilose or tomentose beneath H. brunnescens.

Leaves penninerved, or triplinerved but the basal nerves arising far above the

base of the blade; erect shrubs or trees.

Leaves short-pilose beneath with spreading hairs, often glabrate in age but
some of the pubescence persistent beneath along the costa.

H. guatemalensis.
Leaves glabrous H. mexicana.

Hyperbaena brunnescens Standl. Field Mus. Bot. 22: 21. 1940.

Moist or wet thickets or forest, often on limestone, 1,600 meters
or less; Alta Verapaz; Izabal (type from Puerto Barrios, Standley
73091); endemic.

A small woody vine, the branches densely pilose with short, spreading, golden
brown hairs; leaves on petioles 4-6.5 cm. long, subcoriaceous, ovate-oblong, 13-16
cm. long, 5-8 cm. wide, narrowly acuminate, subtruncate at the base or obtuse,
lustrous above and almost glabrous, brownish beneath, densely velutinous-pilose
with short, spreading, golden brown hairs, 5-nerved, the nerves arising at the base
of the blade.

Known only from sterile specimens, which do not show typical
leaves of fertile branches, and these may be substantially different
in shape.

264 FIELDIANA: BOTANY, VOLUME 24

Hyperbaena guatemalensis Standl. Journ. Wash. Acad. Sci.
15: 475. 1925. Granadilla (Chiquimula) ; Bailador (El Progreso);
Canchijd (fide Aguilar).

Dry brushy hillsides or along stream beds, 250-1,300 meters;
endemic; Zacapa; Chiquimula; El Progreso (type from Barran-
quillo, Wilson Popenoe 965) ; Jalapa (Guastatoya) ; Quiche".

A tree of 9-12 meters, the branchlets densely puberulent; leaves on stout
petioles 1.5-2.5 cm. long, oblong or elliptic-oblong, 10-14 cm. long, 3.5-8 cm. wide,
acute to almost rounded and apiculate, rounded or obtuse at the base, thick-
coriaceous, drying pale green, with somewhat wavy or undulate margins, sparsely
and finely puberulent above or almost glabrous, beneath rather densely and softly
short-pilose or in age glabrate, the nerves prominent on both surfaces, penninerved,
the lateral nerves 6-7 pairs; fruit subglobose, glabrous, 2 cm. long, broadly rounded
at the apex, slightly contracted at the base.

Hyperbaena hondurensis Standl. Field Mus. Bot. 4: 305. 1929.

Dense wet forest or thickets, sometimes in Liquidambar forest,
1,600 meters or less; Alta Verapaz; Huehuetenango. British Hon-
duras; Honduras (type from Lancetilla Valley near Tela, Atlantida).

A large or small vine, the stems sometimes 15 meters long and 3.5 cm. in
diameter, the slender branches puberulent or glabrate; leaves on long slender
petioles, coriaceous, ovate to oblong-elliptic or rarely oblong, mostly 11-20 cm.
long and 5-8 cm. wide, acute or acuminate, sometimes obtuse, usually broadest
near the base, obtuse to subcordate at the base, glabrous above, slightly paler
beneath and glabrous or nearly so, palmately 3-5-nerved, the nerves arising at
the base of the blade; pistillate inflorescences simple, racemose, solitary or fascicu-
late in the leaf axils, half as long as the leaves or often much shorter, densely and
minutely grayish-puberulent, the stout pedicels mostly 2-3 mm. long; sepals 3-3.5
mm. long, minutely sericeous; carpels densely short-pilose; style obsolete.

Hyperbaena mexicana Miers, Ann. Nat. Hist. III. 19: 94.
1867. H. nectandrifolia Standl. Field Mus. Bot. 8: 11. 1930 (type
from Izamal, Yucatan, G. F. Gaumer).

Damp thickets, sometimes on brushy stream banks, 120-1,500
meters; Pete*n; Alta Verapaz (below Tamahu); Santa Rosa (near
Chiquimulilla) ; Retalhuleu (Nueva Linda); Quezaltenango; Hue-
huetenango. Southern Mexico, the type from Cututepeque,
Oaxaca, also in Tabasco and perhaps other states; Yucatan; British
Honduras.

A shrub or small tree with short thick trunk, sometimes as much as 10 meters
high with a trunk 25 cm. in diameter, glabrous throughout, or the young parts
sometimes minutely short-pilose; leaves pale when dried, thick-coriaceous, often
lustrous, on stout petioles 1-2.5 cm. long, oblong to narrowly lance-oblong, 10-22

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 265

cm. long and 2.5-7 cm. wide, acute or sometimes obtuse, acute or obtuse at the
base, penninerved, the lateral nerves 4-7 pairs; staminate inflorescences much
branched, with almost capillary branches, 6-7.5 cm. long, the flowers yellowish;
fruit subglobose, 2.5 cm. long.

It is quite possible that two species may be represented by the
Guatemalan specimens referred here, but they are mostly sterile
and on that account must all be referred for the present to H.
mexicana.

Hyperbaena vulcania Standl. & Steyerm. Field Mus. Bot. 23:
8. 1943.

Moist or wet forest or thickets, 200-1,500 meters, Pacific boca-
costa; Escuintla (type from Barranco Hondo above Las Lajas,
Standky 63878); Guatemala; Sacatepe"quez; Chimaltenango; Solola;
Retalhuleu; Quezaltenango; San Marcos; endemic, so far as known,
but probably extending into Chiapas.

Usually a large vine, climbing over good-sized trees, the branches puberulent
or densely short-pilose; leaves long-petiolate, subcoriaceous, variable in shape and
size, entire, oval or rounded-oval to ovate or rounded-ovate, mostly 9-20 cm. long
and 5.5-13 cm. wide, sometimes even larger, obtuse or rounded at the apex and
apiculate or short-acuminate, or often acute or acuminate, rounded or shallowly
cordate at the base, lustrous and glabrous above, usually fuscous when dried,
brownish beneath, pilosulous or puberulent on the nerves or almost wholly gla-
brous, 5-nerved from the very base, the costa emitting usually 3 nerves on each
side above the base; pistillate flowers racemose, the racemes sessile or pedunculate,
lax and few-flowered, 7-12 cm. long, the rachis and pedicels densely pilosulous or
brown-puberulent, the stout pedicels 6-15 mm. long; inner sepals 5 mm. long,
broadly ovate, obtuse, densely puberulent, the apex recurved, the outer sepals
minute, ovate; carpels densely short-pilose.

Hyperbaena Winzerlingii Standl. Trop. Woods 9: 10. 1927.
Tcansic (British Honduras, Maya).

In chicle forest, 200 meters or less; Pete"n (Carmelita, F. E. Egler
42-239). Northern British Honduras, wet thickets or limestone
forest, little above sea level; type from Orange Walk District,
Winzerling V.12. Yucatan; Campeche.

A densely branched tree 6 meters high with a trunk 15-30 cm. in diameter,
the stiff branchlets puberulent or hispidulous; leaves rigid-coriaceous, the stout
petioles 1 cm. long or shorter, cuneate-oblong, cuneate-oblanceolate, or obovate,
4-13 cm. long, 1-4 cm. wide, very variable in size and outline, even on the same
branch, rounded to subacute at the apex, usually cuneate-attenuate to the base
but sometimes merely acute or even obtuse, usually or often dilated at the apex
and angulate or somewhat trilobate but often entire, glabrous, penninerved;
staminate inflorescences paniculate, axillary, solitary or fasciculate, mostly less

266 FIELDIANA: BOTANY, VOLUME 24

than half as long as the leaves, the almost capillary branches and pedicels hispidu-
lous; pistillate inflorescences 3 cm. long or less, the flowers pedicellate; fruit sub-
globose, somewhat oblique, 1.5 cm. long or somewhat larger, broadly rounded at
the apex, glabrous.

MAGNOLIACEAE. Magnolia Family

Shrubs or more often large trees, glabrous or pubescent; leaves alternate,
membranaceous or coriaceous, entire, penninerved; stipules large, deciduous,
enclosing the young buds; flowers large and showy, solitary, terminal or axillary,
perfect, most often white; sepals and petals often similar, hypogynous, several-
seriate, imbricate, deciduous; stamens numerous, hypogynous, free, the filaments
often thick or dilated; anthers elongate, 2-celled, introrsely dehiscent by longi-
tudinal slits; carpels of the gynoecium numerous, 1-celled, spirally arranged on an
often elongate axis and forming a cone-like spike; ovules 2 or more in each cell,
horizontal; stigmas sessile; fruit dry or fleshy, opening by the abaxial suture, in
age the whole fruit spike often hard and more or less woody; seeds large, the
endosperm abundant, oily, the embryo very small.

Six genera, in temperate and tropical regions of America, Asia,
and Malaysia. The family, at least as represented in America, is
characteristic of warm-temperate regions, in the tropics being
represented only in mountains. Only the following genera are
found in Central America.

Mature carpels of the fruit dehiscent along the dorsal suture; stipules free, the
petioles not scarred Magnolia.

Mature carpels of the fruit circumscissile; stipules adnate to the petiole, leaving
a conspicuous scar near the apex of the petiole after their fall .... Talauma.

MAGNOLIA L. Magnolia

Shrubs or sometimes large trees, glabrous or pubescent; stipule buds terete,
the stipules membranaceous, in bud enclosing the young leaves, free from the
petiole, deciduous; leaves persistent and coriaceous, or membranaceous and decidu-
ous; flowers mostly large and showy, terminal, solitary, sessile or short-pedicellate;
sepals 3; petals 6-12, in 2-4 series, imbricate; anthers linear, the cells introrsely
adnate; gynophore sessile, the carpels numerous, forming an oblong spike, 2-ovu-
late, coriaceous at maturity, persistent, dorsally dehiscent; seeds often pendulous
on a long slender funicle from the opened carpel, drupe-like, the testa fleshy out-
side, crustaceous within.

Species about 35, in Mexico, Central America, southeastern
United States, and Asia. Three additional species are native in
Costa Rica and Panama. A few of the Asiatic species with colored
flowers and thin deciduous leaves are cultivated occasionally for
ornament about Guatemala City. M. Yoroconte Dandy, described
from Copan, Honduras, is to be expected in eastern Guatemala.
Its local name is "yoroconte."

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 267

Leaves sericeous beneath; cultivated tree M. grandiflora.

Leaves glabrous; native species M. guatemalensis.

Magnolia grandiflora L. Syst. Nat. ed. 10. 1082. 1759. Mag-
nolia.

Cultivated rather frequently in Guatemala, in parks and gardens
chiefly of the uplands and highlands, as at Guatemala, Antigua,
Coban, Jalapa, Retalhuleu, Solola, and many other places. Native
of southeastern United States, but introduced into cultivation in
many other parts of the world.

A medium-sized or often large tree with dark bark; leaves short-petiolate,
coriaceous, elliptic to oval or oblong-elliptic, acute or acuminate at each end,
10-30 cm. long, glabrous and lustrous above, covered beneath with lustrous brown
hairs; flowers large and showy, fragrant, the petals creamy white, 5-10 cm. wide;
fruit cone-like, large, oval, the seeds 1.5-2 cm. long.

Wherever known, this tree is esteemed for its beautiful flowers
and leaves, the latter often used in the United States for making
funeral wreaths. It must have been introduced into Guatemala
long ago, for in such places as Antigua there are numerous giant
trees, larger than those seen in cultivation in the United States,
where this magnolia is hardy as far north as Washington, D.C. Trees
at Coban were noted in flower in early April.

Magnolia guatemalensis Bonn. Smith, Bot. Gaz. 47: 253.
1909. Mamey (Zacapa; probably an erroneous name); Magnolia.

Known certainly only from the great swamp east of Tactic,
Alta Verapaz, about 1,450 meters, the type being Tuerckheim
11.2165; trees on the divide along the road from Tactic to Santa
Rosa (Baja Verapaz) perhaps are of the same species although they
may be Talauma (specimens were not obtainable); sterile material
from Sierra de las Minas, Zacapa, probably is referable here.

A glabrous tree 6-15 meters high with a low trunk and a dense, dark green
crown; leaves on petioles 1.5-2 cm. long; elliptic or oval, mostly 12-16 cm. long
and 5.5-8.5 cm. wide, subacute to almost rounded at the apex, obtuse or rounded
at the base, coriaceous, somewhat lustrous, concolorous; pedicels 4 cm. long or less;
sepals about 6 cm. long and 6-8 mm. wide; petals white, 6.5-7 cm. long, 3 cm.
wide, obtuse; stamens almost 100; gynophore 2.5 cm. long and 1 cm. thick, the
carpels about 25; fruiting cones 5 cm. long and 2 cm. thick, or probably larger
in age.

The stipules and sepals are often bright red. The tree is a hand-
some one, although its flowers are smaller and less conspicuous than
those of M. grandiflora nor is the foliage quite so handsome. The

268 FIELDIANA: BOTANY, VOLUME 24

leaves are curious in that they are very concave, with incurved
sides. The tree is abundant in the Tactic swamp, forming dense
groves or thickets. Some of the planted trees in the gardens of
Coban are believed to be of this species. Sterile material collected
in the region of Chelae, Alta Verapaz, perhaps represents an addi-
tional and undescribed species. The leaves are much narrower than
those of M. guatemalensis.

TALAUMA Jussieu

Mostly tall trees, similar to Magnolia, glabrous or nearly so; leaves persistent,
coriaceous, petiolate; flowers terminal, solitary, large and showy, white, sessile
or short-pedicellate; stipules at first united with the petiole, finally deciduous
and leaving a transverse scar at the apex of the petiole; sepals 3; petals 6-many,
in 2 or numerous series, imbricate; anthers linear, the cells introrsely adnate;
gynophore sessile; carpels numerous, capitate or spicate, 2-ovulate, in fruit form-
ing a cone-like structure, thick-coriaceous or woody, at maturity not dehiscent
dorsally but circumscissile near the base, falling off separately or in masses; seeds
like those of Magnolia, often pendulous from the receptacle by long funicles.

Twenty species or more, in tropical America and Asia. Two
other species are known from Central America, in Costa Rica and
Panama.

Talauma mexicana (DC.) G. Don, Hist. Dichl. PI. 1: 851. 1831.
Magnolia mexicana DC. Reg. Veg. Syst. 1: 451. 1818. Palo de pena.

In forest, about 1,500 meters; Huehuetenango (Maxbal);
reported from Alta Verapaz and Baja Verapaz. Southern Mexico;
Honduras.

A large tree, sometimes 30 meters tall with a trunk a meter in diameter,
glabrous or nearly so; leaves long-petiolate, oval or elliptic, mostly 15-30 cm. long,
acute or obtuse at each end, lustrous, the ultimate venation reticulate and promi-
nent; flowers pedicellate, sweet-scented, with an odor suggestive of apple blossoms,
white, sometimes tinged with purple; sepals very broad, about 6 cm. long, thick
and leathery; petals obovate; fruit large and woody, the seeds bright red, with a
juicy outer testa or aril.

Called "anonilla" in Yucatan, where cultivated, the powdered
cones (more probably the petals) said to be used like nutmeg for
flavoring chocolate and other articles of food. Elsewhere in Mexico
the tree is called "flor de corazon" and "yoloxochitl." The tree was
highly esteemed by the original inhabitants in that country because
of the sweet odor of the blossoms, a single flower being sufficient to
perfume a whole house. The flowers were reserved for the exclusive
use of the nobility. The plant was prized also for its medicinal

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 269

properties, and is still used in Mexico in domestic medicine. The
bark is employed as a remedy for fevers, and is said also to have an
effect upon the heart similar to that of digitalis. The Nahuatl name
"yoloxochitl" (heart flower) is an allusion to the shape of the
unopened flower buds.

WINTERACEAE

Trees or shrubs, often with acrid sap; leaves alternate, generally coriaceous,
penninerved, entire; stipules none; flowers relatively small, usually cymose or
fasciculate, perfect or rarely polygamous; sepals 2-6, free and imbricate or united;
petals in 2 or more series, commonly conspicuous in bud, imbricate; stamens
several, hypogynous, the filaments thick or dilated; anthers introrse, 2-celled,
opening by longitudinal slits; carpels of the gynoecium several or only 1, more or
less forming a single verticel, free or partially united; ovules 1-many in each carpel;
stigmas sessile, or distinct styles present; fruit capsular or baccate; seeds with
copious endosperm, the embryo minute.

Six genera, all except the following in southeastern Asia, Malay-
sia, and Australasia. The family has been united by most authors
of the past with the Magnoliaceae.

DRIMYS Forster

Reference: A. C. Smith, The American species of Drimys, Journ.
Arnold Arb. 24:1-33. 1943.

Shrubs or trees with persistent leaves, glabrous, aromatic; leaves pellucid-
punctate, usually whitish beneath; flowers small, perfect or polygamo-dioecious,
the peduncles bearing 1 or several flowers, sometimes appearing pseudo-terminal;
sepals 2-3, membranaceous, in bud united and subglobose, in anthesis irregularly
cleft or ruptured, deciduous; petals 6-many, in 2-many series, imbricate; filaments
stout, the anther cells lateral, parallel or divergent; carpels usually numerous and
forming a single whorl, sometimes few or only 1, many-ovulate, at maturity
baccate, indehiscent; stigmas sessile; testa of the seed crustaceous, lustrous.

About 40 species, 4 American, the others in Australia and Malay-
sia. Only one is found in North America.

Drimys granadensis L. f. Suppl. PI. 269. 1781. D. mexicana
DC. Reg. Veg. Syst. 1: 444. 1817. D. granadensis var. mexicana
A. C. Smith, Journ. Arnold Arb. 24: 23. 1943.

Moist or wet, mixed, mountain forest, sometimes in Liquidambar
forest, 1,600-3,000 meters; Zacapa; El Progreso; Huehuetenango.
Southern Mexico; Costa Rica; Panama; northwestern South
America.

270 FIELDIANA: BOTANY, VOLUME 24

Usually a large shrub or small tree, in Guatemala sometimes 12 meters tall,
with grayish bark; leaves petiolate, narrowly oblanceolate-oblong to oblong or
oblong-obovate, mostly 8-16 cm. long, obtuse or acute, attenuate to the base,
entire, coriaceous, bright green and often lustrous above, usually very glaucous
beneath; flowers solitary or umbellate, long-pedicellate, white, about 1.5 cm. broad;
petals rather few, obtuse or subacute, lance-oblong; stamens bright yellow; berries
subglobose, 5-6 mm. long, at first greenish yellow, at maturity dull black.

In Costa Rica called "muelo" and "quiebra-muelas" ; in Mexico,
"chilillo," "chachaca," "palo picante," and "palo de chile." The
wood is light brown or pinkish, the sapwood grayish, somewhat
suggesting beech (Fagus) ; when freshly cut it has a slight odor sug-
gestive of apples. In regions where abundant (including also the
three related South American species) it has been used for boxes,
cases, interior woodwork, and miscellaneous articles in which great
strength or durability is not required. The tree, as it grows in
southern South America (chiefly D. Winteri Forst.), has had an
interesting history and was formerly of considerable economic
importance. Known in commerce as "Winter's bark," it was first
obtained by Winter, captain of one of the ships of Sir Francis
Drake's expedition of 1577. The three vessels of the fleet were
damaged by storm and Winter's ship was driven to the Straits of
Magellan, where several weeks were spent to recuperate the health
of the crew. Drimys attracted the commander's attention, and he
tried the bark as a preventive of scurvy, then so common among
ships' crews on long voyages. Specimens of the bark were presented
to the famous botanist Clusius, who gave it the name of Cortex
Winteranus. It became a favorite remedy in Europe, but as it was
difficult to obtain the bark from southern South America, that of
Canella alba, a West Indian tree of a different family, often was
substituted for it. At the present time Winter's bark is little used
except in domestic medicine in regions where it is native. It is
aromatic and pungent and has toxic and antiscorbutic properties.
In Costa Rica the bark is chewed to relieve toothache. When
the fresh bark or the leaves are chewed, they burn the tongue
almost like chile. Most of the American species of Drimys are
much alike and in recent years usually they have all been combined
with D. Winteri Forst., under which name the present species has
been reported from southern Central America and from Mexico.
The Guatemalan material is referable to var. mexicana.

ANNONACEAE. Custard-apple Family

Trees or shrubs, the leaves alternate, entire, without stipules; flowers mostly
perfect and 3-parted; sepals 3, rarely 2, valvate or imbricate; petals commonly 6

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 271

and biseriate, valvate or imbricate, the inner often rudimentary or absent; stamens
numerous, the anther cells adnate, the connective usually expanded and truncate
above the anther; carpels of the ovary numerous, rarely few, generally free; ovules
1 or more in each cell; fruiting carpels sessile or stipitate, free (monocarps) or
united to form a fleshy, sometimes very large multiple fruit; seeds with or without
an aril, with copious ruminate endosperm and a minute embryo.

About 75 genera, in the tropics of both hemispheres. A few
additional ones are represented in southern Central America.

Carpels of the fruit more or less completely fused at maturity, forming a usually

very large, globose or ovoid fruit. Petals valvate in bud.
Outer petals with vertical wings; carpels of the fruit united only below.

Rollinia.
Outer petals not winged; carpels of the fruit completely or almost completely

fused Annona.

Carpels of the fruit distinct, often stipitate.
Outer petals imbricate in bud.

Pedicels not bracteate Sapranthus.

Pedicels bracteate.

Flowers axillary . . Guatteria.

Flowers opposite the leaves Malmea.

Outer petals valvate in bud.

Outer petals erect and connivent in flower, oblong or linear. Leaves dis-
tichous; monocarps splitting open at maturity Xylopia.

Outer petals separated in flower and often spreading.

Carpels of the fruit lopsided, dehiscent along one edge; petals mostly

linear-lanceolate Anaxagorea.

Carpels of the fruit indehiscent, not lopsided.

Petals linear-oblong; introduced and cultivated tree Cananga.

Petals broad; native trees or shrubs.

Pedicels without bracts; inner petals somewhat saccate, broad, with

conspicuously incurved edges Cymbopetalum.

Pedicels bracteate; inner petals not at all saccate, plane.

Flowers opposite the leaves Desmopsis.

Flowers axillary, or sometimes produced at leafless nodes.

Unonopsis.

ANAXAGOREA St. Hilaire

Reference: R. E. Fries, Acta Hort. Berg. 12: 6-27. pis. 1, 2. 1934.

Chiefly shrubs, sometimes trees; flowers perfect, axillary, short-pedicellate,
solitary or fasciculate, yellowish green; sepals 3, valvate, united at the base;
petals 6, biseriate, valvate, spreading, plane, subequal, rather thin; stamens numer-
ous, linear, the connective apiculate beyond the anther; torus slightly convex;
carpels numerous or sometimes few, the style subglobose or oblong, the ovules 2
in each cell, basal, erect; mature carpels stipitate, clavate, bivalvate along the
inner edge; seeds not arillate.

Twenty species or more, in Malaysia and tropical America.
Three other Central American species are known from Nicaragua,

272 FIELDIANA: BOTANY, VOLUME 24

Costa Rica, and Panama. A. crassipetala Hemsl. is reported from
Guatemala by Fries (op. cit. 25) on the basis of a Friedrichsthal
specimen from "St. Juan," which doubtless is rather Nicaraguan.

Anaxagorea guatemalensis Standl. Trop. Woods 7: 4. 1926;
Fries, Acta Hort. Berg. 12:26. /. 2, f-g. 1934. Palanco.

Izabal, the type collected between Los Andes and Entre Rios,
S. J. Record 41.

A medium-sized tree; leaves on petioles 7-15 mm. long, papyraceous, obovate,
22-35 cm. long, 9-16 cm. wide, rounded and cuspidate at the apex, acute or
rounded at the base, the adult leaves glabrous, paler beneath; inflorescences about
5-flowered, the pedicels 5-10 mm. long, ferruginous-tomentulose; flower buds
conic; sepals ovate-oblong, obtuse, ferruginous-tomentulose, recurved, 7-8 mm.
long; outer petals ferruginous-tomentulose, 13 mm. long or larger; fruits few,
minutely puberulent, on stipes 15-18 mm. long, the body of the fruit 10-12 mm.
long; seeds 12-14 mm. long and 7-8 mm. broad, black.

This is the most northern species known in the genus, the
majority of whose representatives are South American.

ANNONA L.

Reference: R. E. Fries, Acta Hort. Berg. 10: 197-315. pis. 10-25.
1931.

Trees or shrubs, the pubescence of simple or stellate hairs; flowers usually
perfect, solitary or in few-flowered inflorescences, these terminal, opposite the
leaves, or more or less concrete with the branch and appearing internodal; sepals
3, small, valvate; petals 6, free or connate at the base, biseriate, the inner ones
sometimes rudimentary or none, the outer ones carnose, valvate, concave at the
base or throughout, connivent or somewhat spreading, the inner ones imbricate
or valvate; stamens numerous, extrorse, the connective produced above the cells
into a dilated-truncate disk, rarely attenuate-apiculate or semiorbicular; carpels
numerous, often connate, the ovules solitary, basal, erect; fruit fleshy, consisting
of the concrete carpels.

About 100 species, all natives of America. Several additional
species grow wild in southern Central America. The generic name
has often been written Anona. It is derived from "anon," an
Indian name of the Greater Antilles.

Flowers globose or very broadly pyramidal in bud.

Leaves copiously pubescent beneath; fruit covered with a felt-like tomentum.

A. pur pur ea.

Leaves glabrous beneath or essentially so; fruit not or scarcely tomentose.
Peduncles glabrous; leaves rounded or very obtuse at the base, without
depressions beneath in the axils of the nerves; mature fruit smooth.

A. glabra.

Peduncles sericeous; leaves mostly acute or acutish at the base, with minute
depressions beneath in the axils of the nerves; mature fruit covered with
curved spines A. muricata.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 273

Flowers oblong or narrowly oblong in bud, more or less triquetrous.

Leaves densely velutinous-pubescent beneath, even in age A. Cherimola.

Leaves glabrous or glabrate in age, when young sometimes pubescent but the

hairs chiefly appressed, not velutinous.
Lower leaves of the floriferous branches bract-like, rounded, clasping the

branch; testa of the seed thick.
Leaves 8-14 cm. long, the petiole 1 cm. long or more; peduncles 3-5 cm.

long; basal bract-like leaves soon glabrous A. diversifolia.

Leaves 6 cm. long or less; petiole 5 mm. long; peduncles 1-2 cm. long;
basal bract-like leaves with persistent hairs on the margins and lower

surface A. macroprophyllata.

Lower leaves of the floriferous branches not rounded and clasping; testa of

the seed thin.

Mature fruit with a hard thick shell, the areoles usually somewhat de-
pressed A. scleroderma.

Mature fruit with a thin soft rind, the areoles not depressed, often elevated

or rounded and separated by depressions.
Carpels of the mature fruit free at the apex, the whole fruit covered with

rounded tubercles or projections A. squamosa.

Carpels of the mature fruit completely united, the fruit smooth or
nearly so.

Leaves elliptic, about twice as long as broad A. lutescens.

Leaves lance-elliptic or narrowly lance-oblong, usually 3 times as long

as wide or longer.

Fruits large, commonly 8-12 cm. in diameter, or often much larger.

A. reticulata.

Fruits small, 1.5-3 cm. in diameter A. primigenia.

Annona Cherimola Mill. Gard. Diet. ed. 8. No. 5. 1768.
Anona; Pac (Cacchiquel) ; Pap (Poconchi, Quecchi); Tsumuy,
Tzumux (Quecchi).

Cultivated commonly at 900-1,800 meters and sometimes even
to 2,400 meters, producing best between 1,200 and 1,800 meters;
frequently wild in pastures, hedges, thickets, oak forest, or on open
slopes, 1,200-2,500 meters; Alta Verapaz; Baja Verapaz; Jalapa;
Jutiapa; Santa Rosa; Guatemala; Sacatepe"quez; Chimaltenango;
Solola; Quich^ ; Huehuetenango; Totonicapan; San Marcos. Mex-
ico to British Honduras and Panama; West Indies; Colombia to
Bolivia.

A shrub or small tree, commonly 5-9 meters tall, the branchlets ferruginous-
tomentose; leaves on petioles 8-12 mm. long, membranaceous, commonly elliptic,
rarely lance-elliptic, 8-15 cm. long, 4-9 cm. wide, rounded to obtuse or rarely
acute at the apex, cuneate to rounded at the base, sericeous above at first, soon
glabrate, velutinous-tomentose beneath; flowers opposite the leaves, solitary or
binate, the pedicels tomentose, 8-12 mm. long; sepals triangular, tomentose, 2-4
mm. long; petals linear, obtuse, ferruginous-tomentose outside, 1.5-2.5 cm. long,

274 FIELDIANA: BOTANY, VOLUME 24

greenish inside; fruit globose or ovoid, large, the surface with round protuberances
and marked with U-shaped areoles, sometimes smooth, the pulp white, slightly
acidulous; seeds black.

Popenoe has expressed some doubt as to whether this species is
native in Guatemala but if not, it must have been in cultivation
for a long time, and now is extensively naturalized in many regions
of the highlands. It is the highland anona of Guatemala, its place
being filled in the lowlands by A. reticulata. The fruits of A. Cheri-
mola sometimes are carried down to the lowland markets for sale,
as at Retalhuleu. The fruit is of excellent quality, much liked by
some foreigners residing or traveling in Central America, while
others find it insipid and unattractive. The individual fruits in
Guatemala sometimes weigh six pounds or even more, but ordinarily
they are a good deal smaller. The crushed seeds mixed with lard
are sometimes applied as a paste to the human body to kill lice or
other parasites. In Salvador this species is sometimes called "anona
poshte"; Maya names reported are "pox" (Yucatan) and "tukib"
(British Honduras). The name "chirimoya" (whence the specific
name), probably of Quechua origin, is applied to the species in
Mexico.

Annona diversifolia Safford, Science n. ser. 33: 471. 1911;
Journ. Wash. Acad. Sci. 2: 122. /. l-4a. 1912; Contr. U. S. Nat.
Herb. 18: 19. /. 27-29a, pi. 5. 1914. Anona blanca (Oriente) ; Papauce
(San Marcos).

Cultivated occasionally in the Pacific coast region at 600 meters
or less; wild in thickets in Chiquimula and probably also Jutiapa;
said to be cultivated about Chimaltenango (1,800 meters). Southern
Mexico; Salvador.

A small tree, the branchlets glaucous, quite glabrous; leaves on petioles 8-18
mm. long, membranaceous, obovate, 8-14 cm. long, 4-6 cm. wide, rounded or
subacute at the apex, acute or rounded at the base, glabrous, glaucous beneath;
lower leaves of the flowering shoots orbicular and cordate-clasping, 2-4 cm. long;
flowers solitary, the pedicels slender, glabrous, recurved or pendulous, 3-5 cm.
long, minutely bracteolate below the middle; sepals rounded-triangular, ferrugi-
nous-pilose above, 2-3 mm. long; outer petals linear-oblong, obtuse, minutely
pubescent outside, 2.5 cm. long, about 6 mm. wide at the base, the inner petals
rudimentary; fruit broadly ovoid, tomentulose, generally 13-15 cm. long and
12-15 cm. broad, covered with low rounded protuberances; seeds oblong-ovoid,
2 cm. long, 1 cm. broad.

The flesh is cream-colored or slightly tinged with pink and of
delicious flavor. In Central America, wherever known, this is
usually considered the best of all anonas. It is said to be cultivated

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 275

abundantly in Chiapas about Tapachula. In some parts of Mexico
the fruit is called "ilama," a name of Nahuatl derivation.

Annona glabra L. Sp. PI. 537. 1753. A. palustris L. Sp. PI. ed.
2. 757. 1762. Anonillo (Izabal).

Wet thickets or usually in swamps, often in mangrove swamps,
at or near sea level; Izabal. Southern Mexico to British Honduras
and Panama; southern Florida; West Indies; widely distributed in
South America; western Africa.

A shrub or small tree, sometimes 10 meters tall, the trunk rarely 50 cm. in
diameter, often somewhat enlarged or buttressed at the base, the bark thin, reddish
brown; branchlets glabrous; leaves short-petiolate, papyraceous, bright green,
ovate-elliptic to oblong-elliptic, 7-14 cm. long, 3-8 cm. wide, short-acute or some-
times obtuse, rounded or obtuse at the base, glabrous; flowers solitary, arising
below the petioles, the pedicels 1.5-2 cm. long, glabrous, bracteolate above the
base; sepals rounded, apiculate, glabrous, 3-5 mm. long; petals glabrous outside,
the outer ones ovate, 2.5-3 cm. long, the inner ones somewhat smaller; fruit
globose-ovoid, 5-12 cm. long, smooth, yellowish at maturity, the pulp cream-
colored.

Names applied to the species in neighboring regions are "cork-
wood," "alligator apple," "bobwood" (British Honduras); "anona"
(Honduras); "corcho" (Tabasco); "xmaac," "xmac" (Yucatan,
Maya). The wood is brown, soft, and weak. It is often utilized
along the Atlantic coast of Central America for bottle stoppers and
floats for fishing nets and lines. The fruit is insipid and seldom
eaten by people but there is a popular belief, perhaps correct, that
it is eaten commonly by alligators.

Annona lutescens Safford, Contr. U. S. Nat. Herb. 18: 41.
/. 49-52, pi. 23. 1914. Anona amarilla.

Alta Verapaz, cultivated and perhaps also wild; type collected
near Cahabon, 0. F. Cook 93. Chiapas; reported by Fries from the
Province of Habana, Cuba.

A small tree, the branchlets fulvous-sericeous, becoming glabrate; leaves on
petioles 8-15 mm. long, membranaceous, ovate to elliptic or obovate, 7-14 cm.
long, 3.5-7.5 cm. wide, short-acuminate or obtuse, rounded or subacute at the
base, somewhat sericeous when young but soon glabrate, with only a few hairs
persistent beneath along the nerves; inflorescences opposite the leaves or arising
from the middle of an internode, several-flowered, the pedicels 12-18 mm. long,
sericeous; sepals triangular, 2-3 mm. long; petals linear-oblong, obtuse, puberulent
outside, 1.5-2 cm. long, the inner petals rudimentary; fruit globose-ovoid, smooth,
yellow, 8-9 cm. in diameter or larger, the areoles scarcely perceptible.

This is presumably the pale yellow anona offered for sale in the
Coban market, but we have not found it growing in the vicinity of

276 FIELDIANA: BOTANY, VOLUME 24

that town, and the fruits probably are brought from the lowlands.
The species, although recognized by Fries as a valid one, is based
upon rather slight characters and whether it is more than a form of
A. reticulata can only be determined by further study. Here perhaps
belongs a sterile collection from Alta Verapaz, whose vernacular
name was given as "mecate." The bark is employed for tying
frames of huts.

Annona macroprophyllata Bonn. Smith, Bot. Gaz. 49: 453.
1910; Safford, Contr. U. S. Nat. Herb. 18: 47. pi. 26. 1914.

Type collected near Fiscal, Dept. Guatemala, 1,100 meters,
C. C. Deam 6191. Chiapas (near Tapachula); Salvador.

A shrub of 3-4 meters according to description, but doubtless attaining a larger
size, the branchlets glabrous, glaucous; leaves on petioles 2-3' mm. long, mem-
branaceous, elliptic to obovate or oblong, 4-6 cm. long, 2-3.5 cm. wide, rounded
and often emarginate at the apex, rounded or subacute at the base, glaucous,
glabrous from the first; basal leaves of the branchlets cordate-orbicular and clasp-
ing, 1-2.6 cm. long, at first ferruginous-pilose, later glabrate; flowers solitary, the
pedicels glabrous, 1-2.5 cm. long; sepals ovate, ferruginous-villous, 3-4 mm. long;
outer petals oblong, obtuse, minutely pubescent outside, about 20 mm. long
and 5-7 mm. wide, the inner ones oblong, rudimentary; ovaries glabrous; fruit
unknown.

Annona muricata L. Sp. PI. 536. 1753. Guanaba; Guanabana
(name of Antillean origin) .

Not common in Guatemala but planted in the lowlands, rarely
above 900 meters; occasional in the lower regions of Alta Verapaz
and Izabal, and in the lowlands of the Pacific slope; not known wild
in Guatemala unless occasionally persisting about settlements.
Generally cultivated in tropical America, the native region unknown.

A small tree, 8 meters tall or less, the foliage ill-scented, the young branchlets
ferruginous-sericeous, soon glabrate; leaves on petioles 5 mm. long, papyraceous,
lustrous, obovate to oblong, 8-15 cm. long, 3-6 cm. wide, obtusely short-acute,
short-acute at the base, glabrous above, beneath sericeous at first but soon glabrate,
domatiate in the axils of the nerves; flowers solitary, terminal or opposite the
leaves, the pedicels 1.5-2 cm. long, sericeous; outer petals rounded-ovate, con-
tracted-acute at the apex, cordate at the base, very thick, 2.5-3.5 cm. long, yellow-
ish, the inner petals somewhat smaller; ovaries ferruginous-strigose; fruit ovoid
or oblong-ovoid, 15-20 cm. long or larger, green, covered with curved flexible
spine-like tubercles; seeds black, 1.5 cm. long.

The English name is "soursop." The Maya name of Yucatan is
"tacob." No Indian name for the fruit is known in Guatemala;
hence we suspect that it may be of comparatively recent introduc-
tion, perhaps from the Antilles after the Conquest. The rind of the

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 277

fruit has an unpleasant odor, but the white flesh is agreeably acidu-
lous. Although sometimes eaten as a dessert fruit, the guanaba is
used mostly for flavoring ices and beverages of various kinds, includ-
ing bottled carbonated drinks. The flavor is a popular one and very
agreeable. If quantities of the juice could be preserved and exported
to the United States, there is every reason to believe that it would
become popular there for the same purposes. While the trees are
far from plentiful in Guatemala, the fruits often are available in
quantity in the markets of Guatemala City, to which they are taken
from the lowlands, and in smaller numbers in the market of Coban.
They often weigh five or six pounds or even more. The wood is light-
colored and soft. It is used sometimes in Salvador for making ox
yokes, because the wood is considered fresca, and does not cause the
hair of the oxen's necks to fall out. In Salvador there are distin-
guished two varieties of the fruit: the Guanaba azucaron, that has
sweet flesh and is eaten raw or made into refrescos, and the Guanaba
acida, that is very sour and is used only for preparation of refrescos.
A decoction of the leaves sometimes is applied to the hair to kill
head lice. In the American Virgin Islands the fruit is said to be used
as bait in fish traps.

Annona primigenia Standl. & Steyerm. Field Mus. Bot. 23: 7.
1943. Anonillo.

Moist or wet thickets or forest, 1,000 meters or" less; Pete"n (type
from Gavilan, Fallabon-Yaxha road, Lundell 2213; collected also
at Uaxactun); Alta Verapaz; Zacapa. British Honduras (San
Antonio; San Agustin); Campeche.

A tree as much as 10 meters tall, the trunk to 15 cm. in diameter, the branch-
lets at first sparsely short-pilose, soon glabrate; leaves on slender petioles 7-14
mm. long, membranaceous, darkening when dried, elliptic to lance-oblong or
obovate-oblong, 8-14 cm. long, 3-6 cm. wide, acute or subacuminate, rounded
to subacute at the base, glabrous above, glabrous beneath in age, with small pits
in the axils of the nerves; inflorescences several-flowered, arising from the middle
of the internodes, the fruiting pedicels glabrous, 1.5-3 cm. long; fruit subglobose,
1.5-3 cm. in diameter, almost smooth or sometimes obviously areolate, russet-
colored, sparsely puberulent or almost glabrous; seeds few or rather numerous,
lustrous, dark brown, 8 mm. long.

The fruit is said to be edible, but it can provide little pulp. The
species is noteworthy in having the smallest fruits of all Central
American species. Otherwise it is closely related to A. reticulata.
Possibly it may represent a wild ancestor of the cultivated forms of
A. reticulata.

278 FIELDIANA: BOTANY, VOLUME 24

Annona purpurea Mocifio & Sess£ ex Dunal, Monogr. Anon.
64. pi. 2. 1817. Sencuyo; Sincuyo; Cabeza de muerto; Soncoya;
Suncuyo; Chincuya; Matacuy (name reported, its application ques-
tionable).

Frequent in wet or dry forest, often in second growth or in
thickets, common in cultivation, chiefly at low elevations but some-
times ascending to about 1,200 meters; Alta Verapaz; Izabal;
Chiquimula; Jutiapa; Santa Rosa; Retalhuleu; San Marcos. South-
ern Mexico; British Honduras to Panama; Trinidad; Venezuela.

A tree, often 10 meters high or more, with broad spreading crown, the young
branchlets densely ferruginous-tomentose; leaves large, deciduous, membrana-
ceous, on petioles 3-5 mm. long, broadly obovate to elliptic-obovate, mostly 12-30
cm. long and 6-14 cm. wide, short-acuminate, rounded at the base, green and
glabrate above, paler beneath, brownish-villous even in age; flowers extra-axillary,
solitary, subsessile; sepals triangular-ovate, acuminate, 1-2 cm. long; outer petals
valvate, thick and rigid, ovate-lanceolate, as much as 5 cm. long and 2 cm. wide,
ferruginous-sericeous outside, the inner petals imbricate, thinner, elliptic-oblong,
rounded at the apex, 2.5 cm. long, 1.5 cm. wide; fruit subglobose, 10-12 cm. in
diameter or larger, covered with a rusty felt-like tomentum and with very numer-
ous pyramidal hard pointed projections; seeds obovoid, castaneous, 3 cm. long.

The Maya names "pox," "chacoop," and "polbox" are reported
from Yucatan, and "oop" from British Honduras. The term for
the fruit appears in the name of a caserio of Jutiapa, called Cin-
cuya. The pulp is orange-colored, fragrant, and rather fibrous.
The fruit is often eaten when nothing better is available, but it is
poor in flavor and there is a popular belief that it is "unhealthy."
It does appear at times in the markets.

Annona reticulata L. Sp. PI. 537. 1753. Anona; Anonillo;
Anona colorada; Tzumuy (Quecchi, Poconchi); Pac (Poconchi);
Cahuex (Quiche") ; Oopchi (Pete"n, Maya).

Moist or dry thickets and forest, often in second growth, common
in cultivation, chiefly at 1,200 meters or less, rarely grown at slightly
higher elevations; Pete"n; Alta Verapaz; Baja Verapaz; El Progreso;
Zacapa; Jutiapa; Santa Rosa; Escuintla; Guatemala; Quiche";
Suchitepe"quez; Retalhuleu; Huehuetenango; San Marcos. Mexico;
British Honduras to Salvador and Panama; West Indies; South
America.

A small tree, sometimes 12 meters tall, the trunk 30 cm. or less in diameter,
the crown rounded or spreading, the young branchlets grayish-sericeous, soon
glabrate; leaves on petioles 8-12 mm. long, membranaceous, lanceolate to oblong-
lanceolate, mostly 10-20 cm. long and 2-5 cm. wide, mostly long-acuminate, acute
to rounded at the base, often blackening when dried, at first appressed-pilose on

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 279

both sides but soon glabrate, somewhat paler beneath; inflorescences arising from
the middle of the internodes, rarely opposite the leaves, several-flowered, the
pedicels grayish-sericeous, 1.5-2.5 cm. long; sepals rounded-triangular, acuminate,
2-3 mm. long; petals linear-oblong, obtuse, somewhat dilated at the base, puberu-
lent outside, 1.5-2.5 cm. long; fruit globose-ovoid, 8-12 cm. in diameter or even
larger, usually dark reddish green or reddish brown, almost smooth, the areoles
faint; pulp sweet, rather insipid, somewhat tallow-like.

Maya names reported are "tsulipox," "op," "pox" (Yucatan).
Called "anona colorada" in Yucatan and Salvador. The name
"anona" appears geographically in such place names as Las Anonas,
a caserio of Guatemala, and El Anonal, a caserio of Huehuetenango.
This custard apple is one of the favorite fruits of all Central America,
and large quantities are consumed in its season. It is too sweet and
insipid to please the northern palate, although some foreigners do
become fond of it in time. Apparently the tree is native in Guate-
mala, as in many other parts of Central America. The bark is
chocolate-colored, sapwood whitish, heartwood pale yellow. In
Salvador and probably also in Guatemala the wood is used for
making ox yokes. In Mexico the leaves and branches sometimes are
employed for tanning, and they are said to give a blue or black dye.

Annona scleroderma Safford, Journ. Wash. Acad. Sci. 3: 105.
/. 1. 1913; Contr. U. S. Nat. Herb. 18: 18. /. 22-23. 1914. A. testu-
dinea Safford, op. cit. 106. /. 2, 3. 1913 (type from Tela, Honduras).
Anona del monte; Poxte (Quecchi).

Moist or wet, mixed forest, 1,800 meters or less, chiefly near sea
level; Pete"n; Alta Verapaz (type from Cahabon, 0. F. Cook 89);
Izabal; Huehuetenango. British Honduras; Atlantic coast of
Honduras.

A tree 25 meters high or less, the trunk to 30 cm. in diameter, the young
branchlets ferruginous-puberulent or glabrate; leaves on petioles 1.5-2.5 cm. long,
subcoriaceous, narrowly oblong to oblong-elliptic, 15-35 cm. long, 5.5-9 cm. wide,
acuminate, short-cuneate or rounded at the base, glabrous; flowers greenish yellow,
extra-axillary, often fasciculate on the older branches, the pedicels 1.5 cm. long,
sericeous; sepals connate, sericeous outside, 6 mm. long; petals 3, contracted and
linear above the broad base, ferruginous-sericeous outside; fruit globose or
depressed-globose, 8-10 cm. in diameter, sometimes excavate at the base, con-
spicuously areolate, reddish green, the areoles somewhat depressed and separated
by slightly elevated lines, the rind becoming hard and shell-like; seeds 2 cm. long,
castaneous, lustrous.

Fries considers A. scleroderma and A. testudinea distinct species,
but the characters by which he separates them can hardly be con-
sidered important or likely to be constant. The tree is known only

280 FIELDIANA: BOTANY, VOLUME 24

in the wild state. It is common in wet forest of the Honduran low-
lands, and is reported by Popenoe as occasional in forests of Alta
Verapaz at middle elevations. The fruit has an agreeable flavor,
but the seeds are very large. The leaves and fruit have the odor
characteristic of A. muricata.

Annona squamosa L. Sp. PI. 537. 1753. Anono (tree), Anona
(fruit); Saramuya, Chirimoya (Pete"n).

Infrequent in Guatemala, but cultivated in Pete"n, also in Zacapa,
and well naturalized in some regions of Zacapa, chiefly on low dry
hills. Widely cultivated in tropical America, although usually rare
in Central America; native region unknown.

A shrub or small tree, usually 3-6 meters tall, the crown rounded or spreading,
the branchlets at first grayish-sericeous; leaves on petioles 6-12 mm. long, mem-
branaceous, elliptic or lance-elliptic, 5-11 cm. long, 2-5 cm. wide, subacute,
cuneate at the base, usually blackening when dried, grayish-sericeous when young,
soon glabrate, usually glaucescent beneath; flowers opposite the leaves, pale
yellow, solitary or in few-flowered inflorescences, the pedicels glabrous or pubes-
cent, 1-2 cm. long; sepals rounded-triangular, acute, glabrous or pubescent outside,
1.5-2 mm. long; petals linear-oblong, obtuse, glabrate outside or tomentulose,
1.5-3 cm. long; inner petals rudimentary; fruit globose or cordate-ovoid, glabrous,
glaucous, 8-9 cm. in diameter, the carpels not completely fused but projecting as
rounded protuberances; pulp yellowish white, creamy or custard-like, very sweet,
pleasantly flavored.

The English name is "sugar-apple" or "sweetsop." Among the
various Central American anonas this is easily recognized by its
distinctive fruit, always with more or less pale bloom, and consisting
of incompletely fused, round-tipped carpels, which give it an appear-
ance quite unlike that of other species. Popenoe states that the fruit
is often ruined by insect larvae, but trees observed about Zacapa
were yielding a heavy crop of fine fruit. Lundell reports that in
Pete"n the leaves are placed in bath water of children to refresh
them when they are fretful. In some parts of its range, leaves of
this species are rubbed over floors or placed in hens' nests to keep
away vermin, and the seeds are said to have insecticide properties.

CANANGA Hooker & Thomson

Trees, the leaves petiolate, membranaceous; peduncles arising in the leaf
axils or from defoliate nodes, usually in umbelliform clusters, the flowers large;
sepals 3, valvate; petals 6, biseriate, valvate at first, subequal, elongate, plane;
stamens numerous, linear, the connective produced beyond the anther cells and
acute; torus convex, concave in the middle, the carpels numerous, attenuate to an

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 281

oblong style bearing a capitate stigma; ovules numerous, biseriate; fruit consisting
of stipitate berries, the seeds surrounded by pulp.

About three species, natives of Asia, Malaysia, and Australia.

Cananga odorata (Lam.) Hook. & Thorns. Fl. Ind. 1: 130. 1855.
Uvaria odorata Lam. Encycl. 1: 595. 1785. Canangium odoratum
Baill. ex King, Journ. As. Soc. Beng. 61, pt. 2: 41. 1892. Ilang-ilang.

Native of Burma and Java, but grown for its sweet-scented
flowers in many other tropical regions; introduced rather recently
into Central America, probably by way of the Canal Zone, now
frequent in Panama and occasional elsewhere; planted in Guate-
mala at Zacapa beside the railroad hotel, also at Puerto Barrios, and
said to be in cultivation at various places of the North Coast.

A large shrub or small tree, the slender branches puberulent; leaves on petioles
1-2 cm. long, lance-oblong or ovate-oblong, 10-15 cm. long or larger, acuminate,
broadly rounded or even subcordate at the base, glabrate above, sparsely pubes-
cent beneath; flowers greenish yellow, very fragrant, the petals linear-lanceolate,
long-attenuate; berries oval or oblong, on long slender stipes.

The tree is noted for its intensely fragrant flowers whose odor is
strongest at night, when it can be detected at a long distance. The
fine large tree at Zacapa attracts the attention of many passing
travelers, especially tourists. The flowers yield a fragrant volatile
oil known in commerce as oil of ilangilang, much used in perfumes.

GYMBOPETALUM Bentham

Reference: R. E. Fries, Acta Hort. Berg. 10: 180-194. pis. 6-9.
1931.

Trees or shrubs, the leaves usually large, papyraceous-chartaceous; flowers
large, perfect, solitary, the peduncles terminal or arising between the nodes, some-
times apparently axillary, ebracteate, articulate at the base; sepals 3, short, val-
vate; petals 6, biseriate, valvate, the outer ones sessile, subovate, plane, the inner
larger, very thick, involute-cymbiform, with an inflexed mucro, narrowed at the
base and often short-stipitate; torus convex; stamens numerous, linear-cuneate,
the anthers long, linear, the connective truncate-dilated beyond the cells; carpels
numerous, the ovules 4-14, ventral; fruits stipitate, baccate, oblong-cylindric,
finally dehiscent laterally; seeds ovoid, with a bilobate aril.

Nine species, in tropical America. One other species of Central
America occurs in Costa Rica.

Leaf blades very acute at the base C. stenophyllum.

Leaf blades rounded or very obtuse at the base C. penduliflorum.

282 FIELDIANA: BOTANY, VOLUME 24

Cymbopetalum penduliflorum (Dunal) Baill. Adansonia 8:
268. 1867-68. Unona penduliflora Moc. & Sess£ ex Dunal, Monogr.
Anon. 100. pi. 28. 1817. Orejuela; Muc (Coban, Quecchi); Anon de
montana (Izabal).

Usually in wet forest, at 800 meters or less, sometimes cultivated;
Pete"n; Alta Verapaz; Izabal; Huehuetenango. Veracruz and Oaxaca
to Tabasco; British Honduras.

A tree, often 10-23 meters tall, the trunk 25 cm. or more in diameter, the
bark light or dark gray; young branchlets softly and densely short-pilose; leaves
almost sessile, narrowly oblong to oblong, 10-25 cm. long, 3-8 cm. wide, short-
acuminate, at the base obtuse to subcordate and somewhat unequal, lustrous
above, glabrous except beneath along the costa, there sparsely pilose; flowers
pendulous, the pedicels pilose, 10 cm. long; sepals ovate-triangular, short-acumi-
nate, tomentulose, 7-8 mm. long; petals yellowish green, very thick and fleshy,
grayish-tomentulose, the outer ones plane, broadly ovate, 2.5 cm. long, the inner
ones cymbiform, rounded, short-stipitate, 3 cm. long, the margins strongly
involute; berries short-stipitate, very hard and heavy, 5-8 cm. long, 2.5-3 cm.
thick, reddish brown, rounded at the apex, subterete, containing 9-10 seeds, these
oblong-ellipsoid.

The crown of the tree is pyramidal or spreading; inner bark
whitish ; wood white throughout, turning cream color after exposure,
susceptible to stain, not used so far as known. The curious large
pendent flowers are very fragrant. "Orejuelas," as the dried petals
are called, are well known in many parts of Central America distant
from all places where the tree is known to grow. These petals must
be produced and gathered in great quantities somewhere, to judge
by their occurrence in almost every market, large or small. In
Salvador and Honduras the market people state that they come from
Guatemala, which is doubtless true. In Guatemalan markets it is
invariably stated that they come from Coban, but when one reaches
Coban it is found that the source is somewhere farther on, probably
in the lowlands of Alta Verapaz. There are a few trees planted in
fincas in the city of Coban. The dried petals are employed in Guate-
mala principally for flavoring pinol and other beverages. They were
one of the favorite spices that the ancient Mexicans used for flavor-
ing chocolate and they still are so used in some regions of Mexico,
and probably also in Central America (see W. E. Safford, Science
n. ser. 33: 470. 1911; Smithson. Kept. 1910: 428. 1911; Journ. Wash.
Acad. Sci. 2: 234. 1912). The Nahuatl name was "xochinacaztli,"
signifying "ear-flower," the petals having a fancied resemblance
to the human ear. The bark of this tree is sometimes employed for
making rope.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 283

Cymbopetalum stenophyllum Donn. Smith, Bot. Gaz. 20: 2.
1895; Fries, Acta Hort. Berg. 10: 189. pi. 8. 1931.

Known only from Retalhuleu; type from Caballo Blanco, Rio
Ocosito, 75 meters, J. D. Smith 1491; collected also by Bernoulli
and Cario (no. 3291) in the same department.

A shrub 3.5-4.5 meters high, the young branchlets minutely sericeous, soon
glabrate; leaves on petioles 2-3 mm. long, membranaceous, lanceolate or oblance-
olate, 11-16 cm. long, 3.5-5 cm. wide, rather long-acuminate, acute and unequal
at the base, densely pellucid-punctate; flowers opposite the leaves, the pedicels
glabrous, 3-4.5 cm. long; sepals very broad, 2.5 mm. long; petals grayish-tomentu-
lose, the outer ones broadly ovate, membranaceous, flat, subacute, 1.5-2 cm. long,
13-15 mm. wide, the inner ones fleshy, rounded-obovate, obtuse-apiculate, with
the whole margin involute, 2.5-3 cm. long, 17-20 mm. wide; ovules 6-9 in each
carpel.

DESMOPSIS Safford
Reference: R. E. Fries, Acta Hort. Berg. 10: 16-28. 1930.

Shrubs or trees, the pubescence of simple hairs; flowers perfect, yellow-
green, the inflorescences 1-2-flowered, sometimes arising from the trunk or large
branches, the pedicels commonly elongate, 2-bracteate, the lower bract foliaceous;
sepals 3, valvate, triangular-ovate; petals 6, subequal, biseriate, valvate, linear
to linear-oblong or lanceolate, thick, not nerved; stamens numerous, short, cuneate,
subsessile, the anthers extrorse, linear-oblong, the connective truncate-dilated
beyond the cells; torus convex or subcylindric, pilose; carpels 7-20, the ovaries
setose-pilose; ovules 2-8 in each carpel, parietal, 1-2-seriate; stigmas depressed-
globose or clavate-capitate, sessile; fruits stipitate or rarely subsessile, globose,
ovoid, or short-cylindric, 1-few-seeded; seeds discoid to subglobose.

About 12 species, in tropical America from southern Mexico to
Venezuela. Five other species are known from southern Central
America.

Flowers arising on the trunk of the tree; leaves densely and softly pubescent

beneath D. stenopetala.

Flowers borne on the young branchlets; leaves glabrate beneath or sparsely

pubescent.
Leaves large, all or mostly 3-9 cm. wide, usually much more than 3 cm.

Petals oblong, 10-18 mm. long, 4-6 mm. wide D. bibracteata.

Petals linear, about 25 mm. long D. Schippii.

Leaves small, all or most of them 1.5-2.5 cm. wide.

Leaves conspicuously punctate beneath D. guatemalensis.

Leaves not evidently punctate beneath D. izabalensis.

Desmopsis bibracteata (Robinson) Safford, Bull. Torrey Club
43: 190. pi. 9. 1916. Unona bibracteata Robinson, Amer. Journ. Sci.
III. 50: 175. 1895.

284 FIELDIANA: BOTANY, VOLUME 24

Perhaps occurring in Guatemala, the basis for the report being
Friederichsthal 1176, from San Rafael, which may or may not be a
Guatemalan locality of that name; described from Nicaragua and
known also from Costa Rica and Panama.

A shrub or small tree, the young branchlets sparsely pilose with golden sub-
appressed minute hairs; leaves on petioles 2-3 mm. long, rigid-membranaceous,
oblong-lanceolate to elliptic or rhomboid, 5.5-14 cm. long, 2.5-5.5 cm. wide,
lustrous, glabrous above, beneath hirsute at first but soon glabrate, obtuse or
rounded at the apex, cuneate at the base; flowers solitary, pale yellow, fragrant,
the pedicels 1.5-2.5 cm. long, minutely appressed-pilose; sepals broadly ovate,
obtuse, 2-3 mm. long; petals oblong, 10-18 mm. long, 4-6 mm. wide, sericeous
outside, obtuse; carpels 14-20, the fruits on stipes 5 mm. long, subglobose or
short-cylindric, rounded at each end, constricted between the seeds, glabrous in
age, 5-10 mm. long, 6-8 mm. thick.

Desmopsis guatemalensis Standl. & Steyerm. Field Mus. Bot.
23: 156. 1944.

Moist or wet, mixed, mountain forest, 1,300-1,500 meters;
endemic; Quezaltenango (type from Montana Chicharro, lower
southeastern slopes of Volcan de Santa Maria, Steyermark 34304);
San Marcos (above Finca El Provenir, Volcan de Tajumulco).

A shrub or small tree of 4-6 meters, the branchlets shortly and densely his-
pidulous or pilosulous; leaves small, short-petiolate, firm-membranaceous, some-
what lustrous, the petioles 2-4.5 mm. long, brownish-hirtellous; leaf blades lance-
oblong, 4-6 cm. long, 1.2-1.8 cm. wide, gradually attenuate to the subobtuse apex,
subacute at the base, conspicuously punctate, especially beneath, glabrous
above or puberulent only on the costa, almost concolorous beneath, at first
appressed-pilose but in age pilose only along the costa, the lateral nerves 11-13
on each side; flowers opposite the leaves, the slender peduncle 2 cm. long or in
fruit 2.5 cm. long, appressed-pilose, 2-bracteate, the bracts 1.5-2 mm. long;
sepals ovate, subacute, 2.5-4 mm. long, sericeous outside, glabrous within; petals
fleshy-subcoriaceous, yellowish, linear-lanceolate, gradually attenuate to the apex,
20 mm. long, 1.5-3 mm. wide, sparsely pilosulous outside, glabrous within; berries
on slender stipes 8-9 mm. long, globose, red, 12-15 mm. long, 10 mm. broad,
glabrate; seeds subglobose, brown.

Related to D. lanceolata Lundell which was described from Mount
Ovando, Chiapas, and may well occur in Guatemala. That, how-
ever, has much broader, obtuse petals and larger leaves.

Desmopsis izabalensis Standl. & Steyerm. Field Mus. Bot. 23 :
157. 1944.

Known only from the type, Izabal, on ridge top, along Rio Frio,
Cerro San Gil, 75-150 meters, Steyermark 41543.

A tree of 6 meters, the slender branchlets very densely hispidulous with spread-
ing, brownish or sordid hairs; leaves small, short-petiolate, firm-membranaceous,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 285

more or less lustrous, the petioles about 3 mm. long, densely hispidulous; leaf
blades narrowly oblong-lanceolate, 6-9.5 cm. long, 1.5-2.5 cm. wide, narrowly
long-attenuate to the subacute apex, obtuse or subacute at the base, epunctate,
glabrous above except on the subimpressed costa, there short-hispidulous, almost
glabrous beneath but in age sparsely pilose along the costa; flowers opposite the
leaves, apparently pendulous, the peduncle very slender, in fruit about 3 cm. long,
sparsely hispidulous or almost glabrous; berries on stipes 5-6 mm. long, globose,
9 mm. in diameter, rounded at base and apex, glabrate but when young apparently
appressed-pilose.

Flowers of this species are not known, but probably they will
provide additional characters for separating it from D. guatemalensis,
to which it appears to be closely related.

Desmopsis Schippii Standl. Field Mus. Bot. 11: 130. 1932.

Type from British Honduras, Nineteen Mile, Stann Creek
Valley, growing on creek bank, 75 meters, W. A. Schipp 960; doubt-
less extending into Pet£n or Izabal. Known also from Honduras
(Lake Yojoa, Comayagua, 600 meters).

A tree of 9-18 meters, the trunk 25 cm. or more in diameter, the young branch-
lets appressed-pilosulous, soon glabrate; leaves on petioles 4-6 mm. long, rigid-
membranaceous, elliptic or oblong-elliptic, 12-16 cm. long, 4.5-7 cm. wide,
abruptly cuspidate-acuminate, acute to rounded at the base, glabrous in age;
inflorescences mostly 1-flowered, the slender pedicels 2 cm. long, glabrous or
glabrate; sepals obtuse, 2.5 mm. long; petals yellow or yellow-green, 2.5-3 cm. long,
2.5 mm. wide, sparsely and minutely sericeous.

Desmopsis stenopetala (Bonn. Smith) R. E. Fries, Acta Hort.
Berg. 10: 26. 1930. Porcelia stenopetala Bonn. Smith, Bot. Gaz. 40:
1. 1905. Sapranthus stenopetalus Safford ex Standl. Field Mus. Bot.
4: 206. 1929. Cacao-te.

Moist or wet forest, 500 meters or less; Alta Verapaz, the type
from Cubilgiiitz, 350 meters, Tuerckheim 8496; Huehuetenango.
British Honduras, 630 meters.

A small or medium-sized tree, reported as 9 meters tall with a trunk 20 cm.
in diameter, the young branchlets densely brownish-tomentose; leaves on petioles
4-5 mm. long, oblanceolate or oblong, 18-30 cm. long, 6-9 cm. wide, subcaudate-
acuminate, usually obtuse or rounded at the base, lustrous above and almost
glabrous, densely velutinous-pilose beneath; flowers usually arising on the trunk,
fasciculate, salmon-pink, the pedicels 12-15 mm. long; sepals broadly ovate,
subobtuse, tomentulose outside, 3 mm. long; petals thick, linear from a broad
base, obtuse, about 2 cm. long and 2.5-3 mm. wide, tomentulose outside; carpels
8-12.

Imperfect berries seen are borne on very short thick stipes, oval
or globose, 1-2-seeded, rounded at the apex, glabrous, about 2 cm.
long and 12 mm. broad.

286 FIELDIANA: BOTANY, VOLUME 24

GUATTERIA Ruiz & Pavon

Reference: R. E. Fries, Acta Hort. Berg. 12: 291-549. pis. 1-40.
1939.

Shrubs or trees, the pubescence of simple hairs; flowers axillary, solitary or
few, the pedicels articulate and bracteate below the articulation, perfect, sericeous
outside or sometimes villous or velutinous; sepals 3, valvate; petals 6, biseriate,
imbricate, subequal or the outer ones smaller, erect or spreading; stamens numer-
ous, linear-cuneate, the filaments very short, the connective produced beyond the
anthers into a truncate disk; torus semiglobose-conic or short-cylindric, the carpels
numerous, the ovules solitary, basal, erect; fruits usually stipitate; seeds not
arillate.

About 215 species, all in tropical America. Several additional
Central American species occur in southern Central America.

Inflorescences often several-flowered, or 1-flowered, terminal or arising near the

middle of an internode.

Inflorescences 1-flowered, terminal; leaf blades obtuse or subacute at the base.

G. grandiflora.
Inflorescences arising near the middle of an internode, several-flowered; leaf

blades cuneate-attenuate at the base G. anomala.

Inflorescences 1-flowered, arising from the leaf axils G. amplifolia.

Guatteria amplifolia Triana & Planch. Ann. Sci. Nat. Bot. IV.
17: 35. 1862 (type from Chagres, Panama). G. diospyroides Baill.
Adansonia 8: 269. 1868 (type from Chinantla, Oaxaca). G. dios-
pyroides subsp. hondurensis R. E. Fries, Acta Hort. Berg. 12: 378.
/. 12b. 1939 (type from Lancetilla, Honduras). G. platypetala R. E.
Fries, op. cit. 381. /. llb-c, 12c. 1939 (type from Puerto Barrios,
C. C. Deam 50). Anona.

Moist or wet, dense forest, sometimes in second growth, 400
meters or less; Alta Verapaz; Izabal; Quiche". Southern Mexico;
British Honduras to Panama; probably extending to Colombia.

A shrub or small tree, sometimes 6 meters high, the branchlets sparsely seri-
ceous or almost glabrous; leaves on petioles 4-6 mm. long, elliptic to elliptic-oblong,
mostly 15-30 cm. long and 6-12 cm. wide, usually obtuse or rounded and shortly
cuspidate-acuminate, rounded to subacute at the base, when young sparsely
hirsute but soon glabrous or nearly so or the pubescence more persistent beneath;
flowers solitary or 2 in an axil, the pedicels 8-15 mm. long, sericeous; sepals
rounded-ovate, 5 mm. long, sometimes reflexed, sericeous outside; petals green
•or yellowish green, sericeous outside, oblong-obovate, obtuse, subequal, 14 mm.
long, 7-9 mm. wide; fruits on slender stipes 17-22 mm. long, ellipsoid-fusiform,
narrowed at each end, 10-12 mm. long, 6 mm. thick, turning red and at maturity
black.

We are quite unable to agree with Fries in his division of the
material of this alliance into species, the characters upon which he

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 287

relies for separating them seeming to us fantastically unimportant.
In his section Macrophyllum of Guatteria he recognizes six species
and two varieties (!) or subspecies, four of which are Central Ameri-
can. We have not studied the South American ones, but we strongly
suspect that all six represent a single remarkably uniform unit.
The numerous specimens we have studied are so uniform that it is
hard to imagine how any one ever would have attempted to divide
them into "species." Guatteria amplifolia is one of the characteristic
and often abundant shrubs of the whole Atlantic coast of Central
America.

Guatteria anomala R. E. Fries, Acta Hort. Berg. 12: 524. /.
la-f. 1939. G. grandiflora Bonn. Smith, Enum. PI. Guat. 6: 2. 1903;
Trees & Shrubs 1: pi 26. 1903; not Donn. Smith, 1889.

Known only from the type, Tuerckheim 7816, from Cubilgiiitz,
Alta Verapaz, 350 meters.

Branchlets glabrous; leaves on petioles 3-4 mm. long, obovate or oblong-
obovate, 10-17 cm. long, 5-7 cm. wide, obtusely short-acute, cuneately decurrent
to the base, glabrous; inflorescences arising from the middle of the internodes,
few-several-flowered, the pedicels slender, grayish-puberulent, 1-2 cm. long;
sepals ovate-triangular, finally reflexed, puberulent, 6 mm. long; petals divergent,
subequal, oblong or oblanceolate, obtuse, grayish-pulverulent, about 25 mm. long
and 7 mm. wide; fruits ellipsoid, obtuse, 15-18 mm. long, 10-12 mm. thick, the
stipes 7-8 mm. long; seeds castaneous, strongly rugose.

Guatteria grandiflora Donn. Smith, Bot. Gaz. 14: 25. 1889.

Moist or wet forest, 900-1,200 meters; endemic; type, Tuerckheim
1235, from Pansamala, Alta Verapaz, 1,100-1,200 meters; Huehue-
tenango.

Branchlets glabrous; leaves on petioles 3-6 mm. long, chartaceous, oblong-
obovate or oblong-elliptic, 12-20 cm. long, 4.5-6 cm. wide, abruptly short-cuspi-
date, acute or obtuse at the base, glabrous above, somewhat verruculose beneath,
almost glabrous; flowers terminal, solitary, the pedicels glabrous, 3-3.5 cm. long;
sepals ovate, acute, 7-9 mm. long, reflexed, puberulent; petals fleshy, oblong,
obtuse, tomentulose, 2.5 cm. long, 8-10 mm. wide; fruits 10-12, glabrous, ellipsoid,
obtuse at each end, 2 cm. long, 1 cm. broad, stipitate; seeds corrugate.

MALMEA R. E. Fries
Reference: R. E. Fries, Acta Hort. Berg. 10: 37-46. /. 5. 1930.

Trees or shrubs, glabrous or with pubescence of simple hairs; leaves distichous;
short-petiolate, membranaceous-chartaceous; flowers perfect, medium-sized, the
inflorescences 1-few-flowered, terminal or opposite the leaves; sepals 3, imbricate

288 FIELDIANA: BOTANY, VOLUME 24

in bud; petals 6, biseriate, spreading, subequal or the inner ones slightly larger,
rounded-elliptic, fleshy, blackish when dried, imbricate in bud, with thin margins;
stamens numerous, short, cuneate, the connective truncate-dilated beyond the
anthers; torus hemispheric-columnar; carpels numerous, the ovule 1, basal, erect;
berries numerous, 1-seeded.

Nine species are known, two of them native in Panama and Costa
Rica.

Malmea depressa (Baill.) R. E. Fries, Acta Hort. Berg. 10:
43. 1930. Annona depressa Baill. Adansonia 8: 267. 1868. Guatteria
depressa Safford ex Standl. Contr. U. S. Nat. Herb. 23: 278. 1922.
G. leiophylla (Bonn. Smith) Safford ex Standl. Field Mus. Bot. 3:
268. 1930, nomen nudum.

Wet forest, at or little above sea level; Pete*n; Izabal. Veracruz
to Campeche and Yucatan; British Honduras; Atlantic coast of
Honduras.

A shrub or tree, usually 10 meters high or less, with smooth gray bark, the
trunk 20 cm. or less in diameter, the young branchlets minutely appressed-pilose,
soon glabrous; leaves on petioles 3-4 mm. long, lanceolate to elliptic, mostly 7-12
cm. long and 2-5 cm. wide, acute to attenuate-acuminate, usually acute and
unequal at the base, lustrous above, somewhat pilose when young but in age
glabrous, the veins prominulous; inflorescences terminal or opposite the leaves,
1-few-flowered, the pedicels 1-2 cm. long, glabrous or sparsely hirsute; sepals
rounded-ovate, obtuse, glabrous, 2-3 mm. long; petals broadly ovate or elliptic,
glabrous, greenish, 18-23 mm. long; berries on stipes 1.5 cm. long or shorter,
ellipsoid, red, obtuse, glabrous, 11-13 mm. long and 8 mm. broad.

Known in British Honduras by the names "lancewood" and
"wild soursop." The Maya name of Yucatan is "elemuy." The
fruits are eaten by birds and sometimes by people. The wood is
described as fragrant. In the Forests and flora of British Honduras
(Field Mus. Bot. 12: 137. 1936) a specimen of Malmea depressa was
listed as Oxandra sp. on the basis of a determination by Fries. The
specimen is sterile but there is no doubt that it is really referable
here.

ROLLINIA St. Hilaire

Reference: R. E. Fries, Acta Hort. Berg. 12: 112-190. pis. 9-20.
1934.

Trees or shrubs, the pubescence of simple or rarely stellate hairs; flowers
perfect, solitary or in few-flowered inflorescences, the pedicels bracteate at the
base, articulate above the bract; sepals 3, small, valvate, free or connate at the
base; petals 6, biseriately valvate, connate at the base to form a short globose
tube, the outer 3 petals provided with a spur-like process or with a vertical,
laterally compressed wing, the inner petals minute; stamens numerous, extrorse,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 289

the connective dilated above the anther; torus convex; carpels numerous, 1-ovu-
late, the ovule basal, erect; fruits coalescent to form an often large, globose or
ovoid syncarp.

Fries recognizes 55 species, all in tropical America and mostly in
South America. Four other species are described from Costa Rica
and Panama. Little is known about the fruits of the Central Ameri-
can species, but in general the fruits in this genus are somewhat
similar to those of Annona squamosa and more or less edible. Those
of some of the South American species are reported to be of good
quality, comparable with those of Annonas.

Leaves densely and softly pubescent beneath with lax spreading hairs.

R. Rensoniana.
Leaves sparsely pubescent beneath with wholly or chiefly appressed hairs.

R. Jimenezii.

Rollinia Jimenezii Safford, Journ. Wash. Acad. Sci. 6: 378. /. 3.
1916. Anona; Chirimoya; Anonillo.

Moist or wet forest or thickets, sometimes in dry areas, common
in many parts of the lowlands, chiefly at little above sea level but
ascending to about 1,400 meters; Alta Verapaz; Izabal; Jutiapa;
Santa Rosa; Escuintla; Sacatepe*quez; Suchitepe"quez ; Solola;
Retalhuleu; Huehuetenango; Quezaltenango; San Marcos. Oaxaca
to Tabasco; Honduras; Costa Rica.

A large shrub or usually a small tree, sometimes 10 meters high or even more,
with a trunk 30 cm. in diameter, the bark light gray to pale brown, the trunk
sometimes with small buttresses; young branchlets densely ferruginous-pubescent,
the hairs subappressed; leaves on petioles 7-10 mm. long, membranaceous, obovate
to oblong-elliptic or lance-oblong, 10-24 cm. long, 4-8.5 cm. wide, cuspidate-
acuminate, rounded to acute at the base, pilose above at first but soon glabrate,
slightly paler beneath, pilose along the nerves and veins with rather long, whitish-
ferruginous, mostly subappressed hairs; inflorescences opposite the leaves or
arising slightly below the nodes, 1-3-flowered, the pedicels 1-3 cm. long, bracteate
above the base; sepals rounded-triangular, subacute, ferruginous-sericeous, 2-3
mm. long; corolla green or reddish green, ferruginous-tomentose, about 2 cm.
broad, the wings horizontal or slightly recurved, oblong, not contracted at the
base, 9-10 mm. long, 5-6 mm. high; fruit subglobose, 6-10 cm. long, the carpels
laxly coherent, gibbous, obtuse.

The fruit is edible, with acidulous flavor, but it appears to be
little esteemed in Guatemala, and some informants said it was not
eaten at all. It is reported to be yellow when mature. The inner
bark is dark chocolate-brown; the wood is white or pale yellow.
This is one of the commonest small trees on the low hills and plains
from Retalhuleu to Escuintla, often growing abundantly in fence

290 FIELD IAN A: BOTANY, VOLUME 24

rows. The species has been reported from Guatemala as R. Sieberi
A. DC. and R. pulchrinervia A. DC., species not occurring in Central
America. The material referred here is variable in leaf characters,
and it may well be that when ampler flowering material has been
collected it will be found to represent several species.

Rollinia Rensoniana Stand!. Journ. Wash. Acad. Sci. 13: 351.
1923. R. mexicana Standl. Field Mus. Bot. 11: 155. 1936.

Santa Rosa (Mataquescuintla, 1,500 meters). Salvador (type
from Santa Tecla) ; Veracruz.

A tree about 6 meters high, the young branchlets densely ferruginous-tomen-
tose; leaves on petioles 7-12 mm. long, elliptic or oblong-elliptic, 10-20 cm. long,
4-8.5 cm. wide, acuminate or cuspidate-acuminate, usually rounded at the base,
membranaceous or chartaceous, green above, at first whitish-pilose but soon
glabrate, beneath rather densely covered with long soft yellowish hairs; inflores-
cences 1-2-flowered, the pedicels ferruginous-tomentose, 1-4 cm. long; sepals and
corolla ferruginous-tomentose, the sepals 3 mm. long; corolla 2-2.5 cm. broad, the
wings oblong or obovate, horizontal, not or but slightly contracted at the base;
immature fruit 2.5 cm. in diameter, the carpels acutish, pyramidal, very prominent.

This species was once reported from Guatemala as R. puberula
A. DC. In Salvador the tree is called "churumuyo," and the fruits
are eaten. The wood is employed there for making ox yokes.

Reference: R. E. Fries, Acta Hort. Berg. 10: 3-15. /. 1, 2. 1930.

Shrubs or medium-sized trees, the pubescence of simple hairs; flowers medium-
sized or often very large, ill-scented, dark brown-purple, solitary and opposite the
leaves or arising from the trunk and older branches; sepals 3, imbricate; petals
biseriate, imbricate, subequal, membranaceous, linear-oblong to elliptic; torus
subglobose; stamens numerous, short, sessile, the anthers oblong-linear, extrorse,
the connective truncate-dilated beyond the cells; carpels numerous, sericeous, the
stigmas sessile, globose-disciform; ovules 5 or more, biseriate; fruits sessile or
short-stipitate, mostly oblong-cylindric; seeds commonly numerous.

About 7 species, in Mexico and Central America. One other
species, S. Palanga R. E. Fries, is known from Nicaragua and Costa
Rica.

Petals large, mostly 6-19 cm. long; leaves velutinous-pilose beneath.

Sepals 2-2.5 cm. long; petals 17-19 cm. long S. megistanthus.

Sepals 1-1.5 cm. long; petals 6-8 cm. long S. nicaragiwnsis.

Petals relatively small, 1.5-4 cm. long.

Pedicels mostly less than 1 cm. long; petals 2.5-4 cm. long; leaves velutinous-
pilose beneath S. campechianus.

Pedicels 1.5-2 cm. long; petals 15-22 mm. long; leaves in age glabrate.

S. microcarpus.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 291

Sapranthus campechianus (HBK.) Standl. Contr. U. S. Nat.
Herb. 23: 279. 1922. Asimia campechiana HBK. Nov. Gen. & Sp. 5:
61. 1821. Asimina insularis Hemsl. in Hook. Icon. 16: pi. 1514-
1886. Nitxmaxche (Pete"n, Maya).

Wet thickets or forest, little above sea level, Pete"n. Tabasco to
Yucatan and British Honduras; Honduras.

A large shrub or a tree 6 meters high, the trunk seldom more than 8 cm. in
diameter, the young branchlets pilose; leaves on petioles 2-4 mm. long, mem-
branaceous, oblanceolate to elliptic or obovate-oblong, 5-17 cm. long, 2-7 cm.
wide, acuminate, cuneately narrowed to the acute or obtuse base, in age glabrate
and green above, beneath usually copiously short-pilose; flowers solitary, the
pedicels 5-10 mm. long, bracteate below the middle; sepals triangular-ovate,
subobtuse, pilose outside, 6-7 mm. long; petals linear-oblong, obtuse, 5-7-nerved,
pilose outside, 2.5-4 cm. long, 5-7 mm. wide; fruits subglobose, sessile, densely
tomentulose or in age glabrate, at maturity almost 2 cm. in diameter, usually
several and forming a dense head.

Called "palanco" in Honduras; names reported from British
Honduras are "sufricaya" and the Mayan terms "boytob" and
"elemuy"; Maya names of Yucatan are "chacnixmax," "chacmax,"
and "chac-elemuy."

Sapranthus megistanthus Standl. & Steyerm. Field Mus.
Bot. 23: 7. 1943.

Known only from the type, collected along roadside near Estancia
Grande, Dept. Guatemala, 600 meters, Standley 59219.

A tree of 9 meters, the young branchlets densely tomentose with ochraceous,
soft, mostly spreading hairs; leaves on stout petioles 5-6 mm. long, membrana-
ceous, oblong-elliptic, 10-14 cm. long, 5.5-7 cm. wide, acute or obtuse, obtuse at
the base, green above, softly and densely velutinous-pilose with short whitish
hairs, beneath more densely pilose with longer hairs; peduncles thick, 1.5 cm. long,
tomentose; sepals tomentulose, narrowly lance-oblong, 2-2.5 cm. long, 7 mm. wide
at the base; petals dark brown-purple, sparsely puberulent within, tomentulose
outside, oblanceolate-oblong, 17-19 cm. long, 7 cm. wide, subobtuse, narrowed
to the base.

The pendent flowers have a strong offensive odor of carrion, such
as is found in most or all other species. They are twice as large as
in any other member of the genus.

Sapranthus microcarpus (Bonn. Smith) R. E. Fries, Svensk.
Vet. Akad. Handl. 34, No. 5: 12. 1900. Porcelia microcarpa Donn.
Smith, Bot. Gaz. 20: 1. 1895. Asimina Purpusii Brandeg. Univ.
Calif. Publ. Bot. 4: 375. 1913 (type from Veracruz).

292 FIELDIANA: BOTANY, VOLUME 24

Moist or wet forest, chiefly on the Pacific slope, 100-1,400 meters;
Chiquimula; Santa Rosa; Solola; Quezaltenango (type from Rio
Ocosito, J. D. Smith 1484). Veracruz; Honduras; Salvador.

Often only a shrub of 2 meters but sometimes a tree 12 meters high, the slender
young branches pubescent; leaves on petioles 2-4 mm. long, membranaceous,
obovate to oblong, 6-10 cm. long, 3-5 cm. wide, glabrous above or nearly so,
minutely short-pubescent beneath, especially on the nerves, or often almost
glabrous; flowers solitary, the peduncle 15-18 mm. long; sepals pubescent, lanceo-
late, acute, 6-7 mm. long; petals dark brown-red, puberulent or glabrate, linear-
lanceolate or lance-oblong, obtuse, 15-22 mm. long, 4-5 mm. wide; fruits cylindric,
orange, short-stipitate, 8-9 mm. thick.

Known in Salvador by the names "palanco," "chufle," and
"canjuro." The fruits have a very disagreeable flavor.

Sapranthus nicaraguensis Seem. Journ. Bot. 4: 369. pi. 54-
1866. Porcelia nicaraguensis Benth. & Hook. Gen. PL 1 : 956. 1867.
Cabeza de padre, Guinea de mico (fide Aguilar); Cojon de venado
(Izabal; perhaps S. campechianus).

Moist or rather dry thickets or forest, sometimes in pine forest,
1,400 meters or less; Zacapa; Chiquimula; Jutiapa; Santa Rosa;
Guatemala; Sacatepe*quez; Quiche"; Retalhuleu. Salvador; Hon-
duras; Nicaragua (type collected between Leon and Granada).

A large shrub or small tree, sometimes 7 meters high, the branchlets tomen-
tose; leaves on petioles 5-10 mm. long, membranaceous, oval or elliptic, 10-22
long, 5-10 cm. wide, acute or obtuse at each end, sometimes rounded at the base,
velutinous-pilose on both surfaces or glabrate above; peduncles tomentose; sepals
tomentulose, ovate, subobtuse, 1-1.5 cm. long; petals at first green, turning dark
brown-purple, more or less tomentulose, oblong-lanceolate, obtuse, 6-8 cm. long,
2-3 cm. wide; carpels sericeous at first; fruits sessile, oval, about 5 cm. long and
3.5 cm. broad, rounded at each end.

Called "palanco" and "poshte" in Salvador. Some of the Guate-
malan specimens referred here are sterile and may be referable
rather to S. megistanthus. The flowers are curious because of their
lurid coloring, but repulsive because of their intense and disagree-
able odor.

UNONOPSIS R. E. Fries

Reference: R. E. Fries, Acta Hort. Berg. 12: 231-264. pis. 1-5.
1937.

Shrubs or trees, the pubescence of simple hairs; flowers small, perfect, the
inflorescences axillary, often at defoliate nodes, mostly several-flowered, the
pedicels articulate above the basal bract; flower buds globose; sepals 3, minute,
valvate; petals 6, thick and rigid, subequal, biseriately valvate, ovate or rounded,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 293

concave; torus short-cylindric, truncate at the apex; stamens numerous, cuneate,
the filaments very short; anthers extrorse, the cells linear, the connective dilated
above the anther, disk-like; fruits stipitate; seeds solitary or few, not arillate,
depressed-globose or ellipsoid.

About 22 species, in tropical America from British Honduras to
southern Brazil. Only one species occurs in Central America.

Unonopsis Pittieri Safford, Journ. Wash. Acad. Sci. 15: 102.
1925. U. Schippii R. E. Fries, Acta Hort. Berg. 12: 254. 1937
(type from Jacinto Hills, Schipp 1203).

British Honduras, wet forest, at 60 meters or less; doubtless
extending into Pete"n or Izabal. Atlantic coast of Honduras;
Panama.

A tree of 9-11 meters, the branchlets glabrous; leaves on petioles 3-8 mm.
long, papyraceous, oblong-elliptic to narrowly oblong, mostly 25-35 cm. long and
8-13 cm. wide, rounded or obtuse and cuspidate-acuminate at the apex, rounded
or obtuse at the base, somewhat sericeous at first but in age glabrous or essentially
so; inflorescences arising from defoliate nodes, the branches sericeous, the pedicels
1 cm. long or less; sepals 1.5 mm. long, sericeous outside; petals broadly ovate,
acute, the outer ones sericeous, about 9 mm. long and 7 mm. wide, the inner
slightly smaller; fruits on stipes 5-10 mm. long, black at maturity, globose,
glabrous, 1-1.5 cm. in diameter; seed 1, globose or compressed.

We find no reason for separating U. Schippii, which its author
himself considered rather doubtfully distinct from U. Pittieri.

XYLOPIA L.

Reference: R. E. Fries, Acta Hort. Berg. 10: 86-124. pis. 4-6. 1930.

Trees or shrubs, usually with long slender branches, the leaves coriaceous,
distichous; flowers perfect, solitary or fasciculate in the leaf axils, sessile or short-
pedicellate; sepals 3, connate at the base or higher, valvate; petals 6, biseriately
valvate, the outer ones elongate, thick, narrowly concave, connivent, the inner
ones included; stamens numerous, the connective truncate-dilated above the cells;
torus conic, excavate in the middle; styles elongate; carpels 2-6-ovulate, the ovules
ventral; fruits oblong or elongate, usually at last dehiscent.

About 45 species, in tropical Asia, Africa, and America. Three
other species are known from southern Central America.

Xylopia frutescens Aubl. PI. Guian. 602. pL 292. 1775. X.
frutescens var. glabra Watson, Proc. Amer. Acad. 21: 458. 1886
(type from Lago de Izabal, Watson). Malagueto, Majahua, Capu-
lincillo, Capulin de montana (Pete'n).

294 FIELDIANA: BOTANY, VOLUME 24

Moist or wet thickets, sometimes in pine forest, mostly at 300
meters or less; Pete"n; Izabal; San Marcos. Oaxaca to Chiapas and
Tabasco; British Honduras to Panama; southward to Brazil.

A shrub or tree, said to attain in British Honduras a height of 15 meters and
a trunk diameter of 20 cm. but usually lower, the young branches short-pilose;
leaves on petioles 2-4 mm. long, subcoriaceous, lanceolate, 4-6 cm. long, 8-15 mm.
wide, attenuate-acuminate, acute or obtuse at the base, green and glabrous above,
sparsely or densely sericeous beneath with silvery or ferruginous hairs, or often
glabrate; inflorescences 1-5-flowered; sepals ovate, acute, 2 mm. long; outer petals
white, densely silvery-sericeous, oblong, obtuse, 8-11 mm. long, 2.5 mm. wide,
the inner ones linear-oblong; fruits usually 5-6, subglobose or rhomboid-globose,
on stipes 1-2 mm. long, 10-13 mm. long and 9-10 mm. broad, glabrous, smooth,
orange-red or finally black; seeds 2, obovoid, black, 6-7 mm. long.

Called "polewood" in British Honduras and "palanco" in Hon-
duras. The Indian name "sina" is reported from Honduras (Colon).
Called "tamarindillo" in Oaxaca. The bark is light yellowish brown,
the crown usually depressed and spreading, the terminal branches
very long and slender, with the narrow leaves spreading in two
ranks along the branch. The foliage is handsome, and it is probably
on this account that the tree has been planted along the main street
of Catarina, San Marcos. It is said to grow wild in the lowlands of
San Marcos, but we have seen it wild only in the northern region,
where in some places it is plentiful. The name "palanco" given to
this and some other members of the Annonaceae refers to the fact
that the long straight trunks of small trees are often used as poles
for propelling small boats through shallow water. In Honduras it is
said that the Indians also use the poles for handles of fish spears, and
that oil expressed from the seeds is rubbed on the hair, probably to
give it luster.

MYRISTICACEAE. Nutmeg Family

Reference: A. C. Smith, The American species of Myristicaceae,
Brittonia 2: 393-510. /. 1-9. 1938.

Trees or shrubs; leaves alternate, simple, entire, penninerved, without stipules,
often coriaceous; flowers small, unisexual and monoecious or dioecious, in axillary
or terminal racemes, panicles, or umbels, often fasciculate along the branchlets or
at their ends; perianth simple, usually 3-lobate, the segments valvate; petals none;
anthers 3 or more, extrorse, dorsally adnate to a central stamen column; ovary
superior, 1-celled; ovule 1, basal, anatropous; style short or none, the stigma
disk-like or lobate; fruit normally 2-valvate, often fleshy; seed enclosed in an
entire or laciniate, fleshy aril; endosperm often ruminate.

About 15 genera, in the tropics of both hemispheres. One other
genus, Dialyanthera, is represented in Costa Rica and Panama.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 295

Inflorescence and usually also the lower leaf surface stellate-pubescent, often

densely and conspicuously so Virola.

Inflorescence and leaves glabrous.

Leaves pale beneath, the veins obsolete; staminate flowers not fasciculate.

Myristica.
Leaves not pale beneath; staminate flowers fasciculate Compsoneura.

Dioecious shrubs or trees, the branchlets glabrous; leaves glabrous, petiolate,
entire or slightly undulate, the tertiary nerves subparallel, almost perpendicular
to the costa, often conspicuous; inflorescences 1-2 in the leaf axils or on defoliate
branchlets, racemose, fasciculate-racemose, or narrowly paniculate; bracts sub-
tending the fascicles or lateral branches small or none; bractlets none; flowers
pedicellate; staminate perianth more or less carnose, 3-lobate; anthers 4-10,
oblong, 2-celled, often recurved; ovary subglobose or ellipsoid, the style short,
the stigma peltate or 2-lobate; fruit ellipsoid, glabrous, 2-valvate, smooth or
nearly so, pedicellate, the pericarp very thin; aril entire or nearly so; seed ellipsoid,
irregularly spotted with black or purple.

Eight species, distributed from southern Mexico to Brazil and
Peru. One other Central American species, C. excelsa A. C. Smith,
has been described from Costa Rica.

Compsoneura Sprucei (A. DC.) Warb. Nova Acta Acad.
Leop. Carol. 68: 143. 1897. Myristica Sprucei A. DC. in DC. Prodr.
14: 199. 1856. M. mexicana Hemsl. Biol. Centr. Amer. Bot. 3: 67.
1882. C. costaricensis Warb. Repert. Sp. Nov. 1: 71. 1905. Sangre.

Wet forest, 300 meters or less; Alta Verapaz; Izabal. Tabasco;
British Honduras, along the Atlantic coast to Panama; Venezuela
to Brazil and Peru.

A glabrous shrub or tree, sometimes 14 meters high, the trunk 25 cm. or less
in diameter, the sap red; petioles 1-2.5 cm. long; leaf blades rather thin, elliptic
to narrowly oblong, mostly 10-30 cm. long and 4-10 cm. wide, acute or acuminate
or rounded and cuspidate at the apex, acute to attenuate at the base, lustrous
above, the lateral nerves mostly 4-9 pairs, the tertiary nerves conspicuous; stami-
nate inflorescences 2-8 cm. long, narrowly paniculate or fasciculate-racemose;
flowers in fascicles of 3-15 at the ends of the panicle branches, the slender pedicels
2 mm. long or less; perianth yellow, 1.5-3 mm. long; fruit yellow at maturity,
broadly oval, 2-3.5 cm. long, conspicuously stipitate, rounded at the apex, the
pericarp thin and brittle; aril red.

MYRISTICA L. Nutmeg

Trees; leaves mostly chartaceous, usually whitish or glaucescent beneath;
tertiary nerves mostly obscure or obsolete; inflorescences axillary or supra-axillary,
the peduncles often bifurcate or trichotomous; flowers bracteate, the bractlets

296 FIELDIANA: BOTANY, VOLUME 24

subtending the base of the perianth; flowers rather large for the family, urceolate
or campanulate, pedicellate; anthers 12-30, elongate; style almost none, the
stigmas forming a shallowly bilobate mass; pericarp fleshy-crustaceous, the aril
lace-like, laciniate almost to the base; testa hard, the endosperm ruminate, oily.

Eighty species, in southern Asia, Malaysia, Polynesia, and
tropical Australia.

Myristica fragrans Houtt. Handleid. Hist. Nat. Linn. 2: 333.
1774. M. officinalis L. f. Suppl. 265. 1781. Nuez moscada.

Native of the Moluccas, but grown in many tropical regions for
its seeds, the nutmegs of commerce. Planted upon a small scale in
the lowlands of Alta Verapaz and Izabal, at 350 meters or less;
occasional trees perhaps to be found in other departments.

A tree, generally 9-18 meters high, glabrous throughout or essentially so;
leaves petiolate, subcoriaceous, ovate to elliptic or lanceolate, mostly 8-14 cm.
long, acute or acuminate, acute at the base, the lateral nerves 8-10 pairs; staminate
inflorescence 3-20-flowered, usually bifid, the slender pedicels mostly longer than
the flowers; perianth 5-7 mm. long, urceolate, shallowly 3-lobate at the apex;
pistillate inflorescences usually 1-flowered; fruits short-pedicellate, oval or oval-
obovoid, 3-6 cm. long; aril carmine-red; seeds 1.5-4.5 cm. long, brown.

Nutmeg trees are said to have been planted in Guatemala in
the region of Lake Izabal about 1880, and in recent years they have
been planted with the object of commercial exploitation also in the
lowlands of Alta Verapaz. The seeds are sold everywhere in Guate-
mala, but most of them must be imported from the Old World or
from the Antilles. They are much used in Guatemala for flavoring
desserts and especially such beverages as atol. The dried aril is the
spice known as mace.

VIROLA Aublet

Dioecious trees or rarely shrubs, the inner bark exuding a red sap, the branches
often evidently whorled, the young branchlets tomentose or puberulent; leaves
petiolate, coriaceous to thick-membranaceous, entire or slightly undulate, usually
glabrous above and stellate-pubescent beneath, sometimes glabrate, the tertiary
nerves obscure or obsolete; inflorescences solitary, axillary, broadly paniculate or
almost simple, commonly stellate-pubescent; bracts membranous, enclosing one
or more fascicles of flowers, soon deciduous; bractlets none; flowers sometimes
solitary but usually in fascicles terminating the ultimate branches, pedicellate
or subsessile; staminate perianth pubescent outside, usually 3-lobate; anthers
usually 3; ovary tomentose or puberulent, the style short and stout or obsolete;
fruit globose or ellipsoid, pubescent or glabrous, 2-valvate, the pericarp usually
ligneous; aril laciniate and lace-like; seed globose or ellipsoid.

Species about 38, distributed from Guatemala to Peru and
southern Brazil. Two other species occur in southern Central

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 297

America. The wood is pale brown, light in weight but firm, rather
coarse- textured, easy to work, not durable. It is little used in
Central America, but is said to be suitable for general carpentry
and construction, for boxes, and for other purposes where great
durability is not necessary. It is suitable also for paper pulp. The
trees are abundant in the wet Atlantic coast, often forming a con-
siderable part of the forest. The seeds are beautiful, usually dark
brown and shining, and covered by a lace-like aril. They look very
much like nutmeg seeds but do not seem to have the aromatic
properties of that tree. They are rich in oil, and the oil is reported
to be used in Guatemala for making soap and candles. They often
are found in great quantities under the trees, where they are eaten
by peccaries and other animals. The Panama Indians are said
sometimes to string them on splinters and burn them like candles.
The seeds probably have some aromatic properties, for they are
sold in the country markets, under the name cacao volador and at a
relatively high price, presumably for flavoring beverages or food.
The name "sangre" often given in Central America to Virola alludes
to the red sap and also to the fact that red spots appear on the wood
when it is exposed.

Leaves small, mostly 1.5-3 cm. wide, almost glabrous V. multiflora.

Leaves much larger, mostly 4.5-6.5 cm. wide.

Lateral nerves of the leaves usually 20-30 pairs; leaves persistently stellate-
pubescent beneath, the hairs short-stipitate V. Koschnyi.

Lateral nerves of the leaves 14-21 pairs; leaves glabrate beneath, when young
stellate-pubescent with sessile hairs V. guatemalensis.

Virola guatemalensis (Hemsl.) Warb. Nova Acta Acad. Leop.
Carol. 68: 220. 1897. Myristica guatemalensis Hemsl. Biol. Centr.
Amer. Bot. 3: 66. 1882 (type collected in Guatemala by Skinner,
the locality unknown); 5: pi. 74, /. 5, 6. 1882. V. laevigata Standl.
Field Mus. Bot. 4: 209. 1929 (type from Panama). Chucul (Huehue-
tenango); Palo de sebo; Cacao volador; Cacao cimarrdn.

Moist or wet forest, ascending from sea level to about 1,150
meters; Alta Verapaz; probably in Izabal; Solola; Suchitepe"quez ;
San Marcos; Huehuetenango. Honduras; Costa Rica; Panama.

A tall tree, sometimes 30 meters high, the young branches ferruginous-
tomentulose or cinereous-puberulent; petioles 5-14 mm. long; leaf blades oblong
or narrowly oblong, coriaceous or rather thin, 13-25 cm. long, 4-8 cm. wide,
acuminate or cuspidate, attenuate to broadly obtuse at the base, almost glabrous
when fully developed, when young sparsely puberulent beneath with pale sessile
stellate hairs, the lateral nerves 14-21 pairs; staminate inflorescences 2-3 times
branched, broadly paniculate, many-flowered, 5-12 cm. long and almost as

298 FIELDIANA: BOTANY, VOLUME 24

broad, on a peduncle 1-3 cm. long, the branches ferruginous-puberulent, the
flowers in clusters of 5-10, the pedicels 1 mm. long or less; perianth 2 mm. long,
sparsely stellate-puberulent; fruits on pedicels 5-10 mm. long, ovoid-ellipsoid,
2.5-3.5 cm. long, with a thick pericarp; seed ellipsoid, 2-2.7 cm. long.

Called "sangre" in Honduras. In Guatemala the dry seeds are
much used for flavoring chocolate and other beverages and they are
sold commonly for this purpose in the markets, at a relatively high
price. Oil from the seeds is employed in some quantity for making
soap and candles and for oiling machinery. The young branches
often appear in whorls at the ends of the larger ones and such whorls
are used like egg-beaters for whipping chocolate and for stirring
food in the process of cooking.

Virola Koschnyi Warb. Repert. Nov. Sp. 1: 71. 1905 (type
from San Carlos, Costa Rica). V. merendonis Pittier, Contr. U. S.
Nat. Herb. 20: 453. 1922 (type from Cordillera de Merendon, on
the border between Guatemala and Honduras). Sangre; Drago;
Cedrillo.

Wet forest, 300 meters or less; Alta Verapaz; Izabal. British
Honduras; Honduras; Nicaragua; Costa Rica; Panama.

A tall tree, sometimes 35 meters high with a trunk 1.25 meters in diameter,
the branchlets densely stellate-tomentose; leaves on petioles 7-12 mm. long, thin-
coriaceous or almost membranaceous, narrowly oblong to narrowly elliptic, 13-35
cm. long, 4-13 cm. wide, cuspidate, rounded or obtuse at the base, glabrous above
in age, densely tomentose beneath with stipitate stellate hairs or finally glabrate
and glaucescent, the lateral nerves 18-35 pairs; staminate inflorescences 1-2-
branched, 6-13 cm. long and almost as broad, on peduncles 4 cm. long or less, the
branches and flowers densely tomentose; pedicels 2-5 mm. long; perianth 1.5-3
mm. long; fruits ellipsoid, 2-3 cm. long, densely tomentulose or finally glabrate.

Called "banak" in British Honduras, where the species is con-
sidered the most important of the secondary timbers of the colony.
The trunk usually is supported by small buttresses, and is free of
limbs for most of its length. The wood is used for interior wood-
work and has been exported from British Honduras to the United
States, chiefly for the manufacture of plywood.

Virola multiflora (Standl.) A. C. Smith, Brittonia 2: 499. 1937.
Dialyanthera multiflora Standl. Field Mus. Bot. 8: 12. 1930 (type
from Stann Creek, British Honduras, Schipp 279). V. brachycarpa
Standl. Field Mus. Bot. 11: 131. 1932 (type from Stann Creek
Valley, British Honduras, J. A. Burns 20).

Wet hillside forest, British Honduras, at or near sea level; to be
expected in Izabal.

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 299

A tree of 15 meters with a trunk 30 cm. or more in diameter, the branchlets
sparsely cinereous-puberulent or glabrate; leaves on petioles 5-11 mm. long, thin-
coriaceous, narrowly lance-oblong or oblanceolate-oblong, 6-17 cm. long, 1.5-5.5
cm. wide, acute or acuminate, acute or attenuate at the base, glabrous above or
nearly so, sparsely puberulent beneath at first with sessile stellate hairs, in age
glabrous, the lateral nerves 7-19 pairs; staminate inflorescences 1-2-branched,
3-7 cm. long, the branches minutely puberulent, the pedicels 3 mm. long or less;
perianth puberulent, 1.7-2 mm. long; fruits pedicellate, ellipsoid, 14-17 mm. long,
rounded at the apex, rounded or substipitate at the base.

Known by the names "banak" and "bastard banak."

MONIMIAGEAE

Reference: Janet Perkins & Ernst Gilg, Monimiaceae, Pflanzen-
reich IV. 101. 1901.

Shrubs or small trees, often with resin cells; leaves chiefly opposite, entire or
unequally dentate, membranaceous to coriaceous, penninerved; stipules none;
flowers small, greenish or yellowish, regular, mostly unisexual and monoecious or
dioecious, mostly in axillary or terminal cymes, rarely racemose, paniculate, or
fasciculate; receptacle usually campanulate, globose, or urceolate, membrana-
ceous or carnose, in the pistillate flowers the upper portion often circumscissile after
anthesis, the lower part strongly accrescent, becoming woody or coriaceous and
bearing the carpels, or the whole receptacle accrescent and becoming globose or
urceolate and enclosing the carpels; sepals 4-many, small or minute, often none;
stamens few to very numerous, mostly free, rarely connate into a tube, the fila-
ments filiform or liguliform, equal or unequal, the outer ones often somewhat
petal oid; anthers dehiscent by longitudinal or transverse slits or by valves; ovary
of usually numerous carpels, these free or rarely connate, sometimes immersed in
the receptacle, the carpels 1-celled; ovules solitary, erect or pendulous, usually
anatropous; styles commonly filiform and elongate, generally free; carpels of the
fruit usually distinct and numerous, drupaceous, sometimes enclosed in the
enlarged receptacle; seeds erect or pendulous; endosperm carnose, copious; em-
bryo straight, axial; cotyledons ovate to orbicular, the radicle inferior or superior.

About 30 genera, widely dispersed in tropical regions of both
hemispheres. In North America only two genera are found.

Anthers dehiscent by valves; fruit enclosed in the enlarged, globose or obovoid

receptacle Siparuna.

Anthers dehiscent by longitudinal spits; fruits not enclosed in the receptacle.

Mollinedia.

MOLLINEDIA Ruiz & Pavon

Shrubs or small trees; leaves opposite, entire or dentate, membranaceous or
coriaceous, glabrous or pubescent; flowers unisexual, small, dioecious, in 3-flowered
clusters, arranged in axillary or terminal panicles or racemes, the bracts and
bractlets minute or none; staminate receptacle variable in form, membranaceous
to coriaceous, glabrous or pubescent; sepals 4, in opposite pairs, the 2 outer ones

300 FIELDIANA: BOTANY, VOLUME 24

larger, connivent and imbricate in bud, spreading in anthesis; stamens 8-50,
usually unequal, the filaments very short or none; anthers ovate or oblong, dehis-
cent by longitudinal slits; pistillate receptacle like the staminate one, the sepals
united at the base to form a campanulate cup, the 4 lobes small, subequal, the cup
circumscissile and deciduous after anthesis; carpels 6-35, glabrous or pilose, the
style short; ovule pendulous from the apex of the cell; drupes few or numerous,
inserted on the dilated receptacle, sessile or short-stipitate.

Species 70 or more, all in tropical America, mostly in South
America. Several other species are known from Central America.

Mollinedia guatemalensis Perkins, Bot. Jahrb. 27: 679. 1900
(type, Bernoulli & Carlo 2544, probably from the Pacific bocacosta).
Sakeyen, Anyac (Alta Verapaz); Cafe de montana; Canela de montana.

Usually in dense, moist or wet, mixed forest, 1,700 meters or less;
Pete"n; Alta Verapaz; Izabal; Zacapa; Escuintla; Sacatepe"quez;
Chimaltenango; Suchitepe"quez; Quiche"; Huehuetenango; Que-
zaltenango; San Marcos. British Honduras.

A shrub or tree 2-12 meters high, usually with few branches, the branches
green or ochraceous, sericeous when young, soon glabrate; leaves on petioles about
1 cm. long, elliptic-oval to elliptic-oblong or lance-oblong, mostly 12-18 cm. long
and 3.5-8 cm. wide, acuminate, cuneate-attenuate to obtuse at the base, rather
thick and firm, entire or more often remotely serrate toward the apex, green above,
glabrous, somewhat paler beneath, sparsely pilose with minute appressed hairs;
inflorescences axillary, few-flowered, the flowers yellow or yellowish green, the
pedicels often greatly elongate; receptacle cup-like or ovoid, 6-7 mm. long, the
sepals very short, obtuse or acute, yellowish-strigose outside; stamens about 40;
fruits ellipsoid, green, glabrous, obtuse, about 13 mm. long.

This plant is an inconspicuous one, with no outstanding charac-
ters that may be indicated for its ready recognition, and it is difficult
to place systematically unless one is already familiar with the family.
The available material is somewhat variable and it is possible that
more than one species is represented, but the species described from
Central America and Mexico are already too numerous, and it is
uncertain whether M. guatemalensis is really distinct from some of
the species described from Mexico. One sterile collection from San
Marcos perhaps is referable to one of the narrow-leafed Mexican
species, but until better material of it is collected it cannot be placed
definitely.

SIPARUNA Aublet

Shrubs or small trees, the pubescence often of branched hairs; leaves chiefly
opposite, entire or more often dentate, membranaceous to coriaceous, petiolate;
flowers small, monoecious or dioecious, in axillary cymes, or the inflorescences
sometimes paniculate or racemose; staminate receptacle usually campanulate,

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 301

globose, or urceolate, membranaceous or coriaceous; sepals 4-7, large or small,
sometimes obsolete, usually connate to form a lobate or entire ring, the velum
closing the receptacle often conic, sometimes plane or obsolete; stamens 1-60,
usually unequal, the filaments ligulate to cylindric; anthers dehiscent by valves
on the inner side; carpels of the ovary 4-20, the styles filiform or liguliform, free
or connate; ovule 1 in each carpel; fruits drupaceous, globose or obconic, longi-
tudinally sulcate; seed ascending, with copious endosperm.

About 100 species, all in tropical America and mostly in South
America. Several others are known from southern Central America.

Leaves densely hirsute, the hairs all or mostly simple S. Tonduziana.

Leaves glabrous or glabrate, the hairs minute and stellate S. nicaraguensis.

Siparuna nicaraguensis Hemsl. Biol. Centr. Amer. Bot. 3: 69.
1882. Chuche (Quecchi) ; Kex (San Marcos) ; Hormiguillo (Huehue-
tenango) ; Salvia (San Marcos) ; Cerbatana.

Mostly in moist or wet, dense, mixed forest, sometimes in open
pine forest, 1,800 meters or less, mostly at 600-1,500 meters; Pete"n;
Alta Verapaz; Baja Verapaz; Izabal; Chiquimula; Quiche"; Huehue-
tenango; San Marcos. Southern Mexico; British Honduras to
Honduras, Nicaragua, and Panama.

A shrub or tree, usually 2-6 meters high, sometimes reclining or subscandent,
the older branches ferruginous, the young branches stellate-puberulent, soon
glabrate; petioles very unequal, that of one of a pair of leaves often twice as long
as the other; leaf blades oval to oblanceolate-oblong, often obovate, mostly 7-15
cm. long and 3.5-7 cm. wide, abruptly short-acuminate, usually narrowed toward
the base, the base narrowly rounded to cuneate-attenuate, entire or inconspicu-
ously undulate-dentate, glabrous above or nearly so in age, beneath sparsely
stellate-puberulent or almost glabrous; flowers dioecious, the inflorescences
axillary, few-flowered, equaling or shorter than the petioles, the pedicels mostly
2-4 mm. long but sometimes more elongate; staminate receptacles greenish yellow
or dark red, often orange, cup-like, about 4 mm. broad, minutely stellate-puberu-
lent or glabrate; sepals 4-5, triangular, thickened at the apex, glabrous within;
stamens 5-6; pistillate flowers 4-5 mm. broad, carnose, the sepals 4-6, broadly
oval or rounded; fruiting receptacles 1-1.5 cm. in diameter or larger, usually rose-
colored, rupturing irregularly and exposing the carpels, crimson within.

Called "wild coffee" in British Honduras; "limoncillo" (Hon-
duras); "palo de carabina" (Oaxaca). The leaves have a strong
odor of lemon when crushed. They are used, especially in Alta
Verapaz, for brewing an aromatic tea that is a favorite remedy for
influenza and catarrh. The Quecchi Indians place the leaves on
their foreheads to relieve headache. The wood is soft and white.
The fruits are curious, somewhat suggestive of the pink insect galls
so often found on oak (Quercus) trees. This species has been reported
from Guatemala as S. riparia (Tul.) A. DC., a quite different Mexi-

302 FIELDIANA: BOTANY, VOLUME 24

can plant. A few of the Guatemalan specimens approach S. Sumi-
chrastii (A. DC.) Perkins (S. riparia var. Sumichrastii A. DC.),
which is rather doubtfully distinct from S. nicaraguensis.

Siparuna Tonduziana Perkins, Bot. Jahrb. 31: 746. 1902.
Salvia; Cerbatanero.

Wet forest, at or near sea level; Izabal. Honduras, along the
Atlantic coast to Panama.

A stout shrub 2-3 meters high with few branches, the branches hirsute with
long, simple or stellate hairs; petioles 1 cm. long or less, subequal; leaf blades thin,
oblong to oval-obovate, mostly 12-28 cm. long and 5-13 cm. wide, long-acuminate
or abruptly short-pointed, somewhat narrowed to the obtuse or rounded base,
rather conspicuously serrate, rather densely hirsute, especially beneath, with long,
spreading, simple or stellate hairs, rough to the touch; flowers yellow or greenish
yellow, with an orange velum, cymose, the inflorescences little if at all exceeding
the petioles, the pedicels mostly 3-4 mm. long, hirtellous; flowers 2.5 mm. broad,
the receptacle densely hirtellous; sepals minute, triangular, obtuse; stamens 4-5,
short-exserted.

This plant also has the lemon odor that probably characterizes
all the Central American species.

LAURACEAE. Laurel Family

References: Carolus Mez, Lauraceae americanae, Jahrb. Bot.
Gart. Berlin 5. 1889; Caroline K. Allen, Studies in the Lauraceae,
VI. Preliminary survey of the Mexican and Central American
species, Journ. Arnold Arb. 26: 280-434. 1945.

Trees or shrubs, usually aromatic, rarely parasitic and scandent herbs or
suffrutescent plants, glabrous or pubescent; leaves mostly alternate and petiolate,
simple, entire, penninerved or often triplinerved; stipules none; inflorescences
usually axillary, paniculate, spicate, racemose, umbellate, or rarely capitate, the
bracts deciduous or sometimes forming a more or less persistent involucre; flowers
small, regular, perfect or dioecious, sometimes polygamo-dioecious, often fragrant;
perianth tube small or conspicuous, conic, funnelform, or urceolate, in age generally
accrescent and forming a cupule at the base of the fruit (the berry and cupule
suggestive of an acorn), rarely deciduous; perianth segments 4 or 6, biseriate, the
outer ones sometimes smaller than the inner; stamens usually in 3 or 4 series of 3,
alternate, attached to the perianth tube; stamens of the outer 2 series fertile,
usually eglandular, introrse or rarely extrorse; stamens of the third series usually
fertile, with introrse, lateral, or apical cells, the base of the filament with 2 glands
at the outside; stamens of the fourth (innermost) series usually sterile and reduced
to staminodia, sometimes obsolete; anthers ovate, oblong, rectangular, or tri-
angular, usually with 2 or 4 cells, the cells in 2 vertical rows or in one arcuate row,
opening by valves, usually from the base to the apex, the valves often persistent
and spreading; filaments commonly free, or those of the third series rarely united,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 303

the basal glands mostly sessile and free; ovary free, epigynous, 1-celled, the single
ovule anatropous, pendulous, attached near the apex of the cell; style usually
conspicuous, the stigma obtuse or rarely capitate; fruit a 1-seeded berry or drupe,
usually surrounded at the base by the persistent perianth tube; seed without
endosperm, the testa generally membranous; cotyledons flat-convex.

About 40 genera and 1,000 species, almost confined to the tropics.
A few additional genera are represented in southern Central America.
One tree of the genus Sassafras is found in temperate North
America. Its name is derived from an Indian language of North
America, but this name, in some unknown manner, has become
established in Central and South America for various plants, usually
of other families. In Central America, for instance, the name
sassafras is sometimes applied to species of Croton (Euphorbiaceae) .
The most celebrated and typical plant of the Lauraceae is the Old
World laurel, Laurus nobilis L., native in southern Europe, long a
symbol of victory.

The family is an important one in the tropics as a source of
lumber and in one genus of fruit. It is therefore particularly unfortu-
nate that taxonomically it is perhaps the most difficult group of all
tropical American plants. The American species have not been
monographed as a whole since the time of Mez's monograph, now
long out of date. A good beginning upon a new monograph was
made a few years ago by A. J. G. H. Ostermans of Leiden, but the
work was discontinued long before completion. A complete mono-
graph of all the American Lauraceae is sadly needed, but probably
it will not be of great practical importance when available, nor
will it greatly facilitate determination of material. The flowers
throughout the family are monotonously alike in outward appear-
ance, but of highly varied stamen structure. Trees almost identical
in foliage are found to have quite different flower structure, and
usually it is only by dissection of flowers that the genus can be
determined. In a few genera, such as Litsea and Per sea, it usually
is possible to recognize the genus by general appearance. Because
of the nature of the flowers it is improbable that any simple or easily
workable classification for the family ever can be invented. Fruiting
material of the family usually is quite worthless for purposes of
determination, but often it can be matched by leaf characters with
properly named flowering specimens.

There are given below two general keys, a technical one to the
genera, based chiefly on stamen characters, and a purely artificial
one to the collective species of Beilschmiedea, Licaria, Nectandra,
Ocotea, and Phoebe. With the latter it should be possible to deter-

304 FIELDIANA: BOTANY, VOLUME 24

mine most specimens with fruit, and flowering ones by only the
external characters of the flowers. There is so much variation in
some of the characters used in this key that it, like practically all
other keys, will often be found insufficient. Several of the species
listed on the following pages are very incompletely known, and
in some instances their proper generic status is still doubtful.

KEY TO THE GENERA

Plants small glabrous parasitic vines, without chlorophyll, twining, the stems

herbaceous or nearly so; leaves reduced to minute scales Cassytha.

Large trees or shrubs with normal leaves.

Flowers capitate or umbellate, subtended by an involucre of 4 membranaceous
bracts. Shrubs or small trees with small leaves Litsea.

Flowers not involucrate.

Calyx segments usually very unequal, the outer ones shorter. Anthers

4-celled; fruit usually very large Persea.

Calyx segments equal in length or nearly so; anthers 2-celled or 4-celled.
Anthers 2-celled.

Staminodia (innermost series of stamens) large, ovate or triquetrous.

Beilschmiedea.

Staminodia none or minute and stipe-like Licaria.

Anthers 4-celled.

Staminodia well developed, sagittate or triangular.

Perianth segments persistent Phoebe.

Perianth segments deciduous after anthesis.

Leaves conspicuously triplinerved Cinnamomum.

Leaves penninerved Persea.

Staminodia minute and stipe-like or none.

Anther cells in pairs, one pair above the other Ocotea.

Anther cells all inserted at nearly the same height Nectandra.

ARTIFICIAL KEY TO THE SPECIES OF ALL THE GENERA, EXCLUDING LITSEA

AND PERSEA

Leaves densely tomentose beneath over the whole surface, or densely pilose with
chiefly spreading hairs, or densely spreading-pilose at least along the costa
and nerves, the pubescence persistent wholly or in part, even in age.

Leaves densely covered beneath with a close ferruginous tomentum, this
persistent and conspicuous in age, the leaves bicolored .... Phoebe Salvinii.

Leaves with various pubescence beneath but not as above, not conspicuously
bicolored.

Margins of some or all the leaves conspicuously recurved at the base, often
forming a large basal pocket.

Perianth about 3 mm. long Phoebe amplifolia.

Perianth 5-6 mm. long Nectandra reticulata.

Margins of the leaves not recurved at the base.

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 305

Staminodia or innermost series of stamens minute or none, stipe-like;
leaves very large, mostly 10-18 cm. wide, tomentose beneath with
mostly matted hairs or in age often glabrate; branches usually tomen-
tose, the tomentum mostly close or even appressed . .Nectandra sinuata.

Staminodia well developed, sagittate, or, if minute or aborted, the leaves
6 cm. wide or less; leaves variable in size, usually hirsute beneath,
never tomentose; branches usually hirsute.

Flowers pubescent, at least on the lower part of the perianth, the seg-
ments often densely pilosulous throughout.

Leaves whitish and pruinose beneath Beilschmiedea Anay.

Leaves green beneath.

Branches hirsute Phoebe belizensis.

Branches appressed-tomentulose Ocotea rubriflora.

Flowers glabrous.

Staminodia none or vestigial; flowers and fruit sessile or practically so.

Licaria Peckii.

Staminodia well developed; flowers usually pedicellate, sometimes
long-pedicellate, the cup of the fruit long-stipitate.

Leaves acute or subacute at the base, small, mostly 2-2.5 cm. wide.

Phoebe Bourgeauviana.

Leaves rounded or very obtuse at the base or even subcordate,
usually much wider Phoebe helicterifolia.

Leaves glabrous beneath or sericeous or appressed-pilose, sometimes sparsely
puberulent or barbate in the leaf axils, never with abundant spreading hairs
or tomentose.

Leaves conspicuously triplinerved.
Flowers pubescent.

Leaves small, mostly 7-8 cm. long; inflorescences small, about 3 cm. long,

corymbif orm, few-flowered Phoebe savannarum.

Leaves large, mostly 12-18 cm. long; inflorescences large, paniculate,
many-flowered Phoebe mexicana.

Flowers glabrous, or practically so.

Leaves coarsely and laxly reticulate- veined beneath Phoebe effusa.

Leaves very closely and finely reticulate-veined beneath, the surface
almost pitted.

Leaf blades acute at the base Phoebe Ehrenbergii.

Leaf blades rounded at the base or rounded and abruptly short-acute.

Phoebe areolata.
Leaves not triplinerved.

Leaves finely sericeous or strigillose beneath or puberulent, the pubescence
persistent in age, usually very inconspicuous to the naked eye but
evident under a lens.

Perianth segments united almost to the apex Licaria campechiana.

Perianth segments free almost to the base.
Flowers densely pubescent.

Leaves mostly 7-12 cm. wide Phoebe Gentlei.

Leaves mostly 2.5-5.5 cm. wide.

Leaves acute or subobtuse, mostly elliptic-oblong.

Nectandra surinamensis.
Leaves narrowly long-acuminate, lanceolate or oblong-lanceolate.

Nectandra membranacea.

306 FIELDIANA: BOTANY, VOLUME 24

Flowers glabrous or very sparsely pubescent Phoebe saxchanalensis.

Leaves glabrous beneath or practically so, at least in age, scattered hairs
sometimes persistent along the nerves, and the nerve axils often densely
barbate.
Flowers capitate, the inflorescence simple or compound.

Leaves small, mostly 8-10 cm. long Licaria coriacea.

Leaves large, mostly 12-20 cm. long Licaria capitata.

Flowers not capitate, the inflorescence usually branched.
Flowers glabrous or practically so.

Perianth with a conspicuous tube, the flowers when dry 2-2.5 mm.
broad.

Anthers 4-celled Ocotea Bernoulliana.

Anthers 2-celled.

Basal glands of the stamens free Licaria Cervantesii.

Basal glands of the stamens united in pairs Licaria caudata.

Perianth cleft nearly or quite to the base, the flowers much broader.
Leaves large, mostly 16-25 cm. long.

Veins conspicuously elevated and reticulate on the upper leaf

surface Ocotea verapazensis.

Veins not elevated on the upper leaf surface . . . Ocotea ovandensis.
Leaves relatively small, mostly 8-15 cm. long.

Inflorescence racemiform or narrowly thyrsoid-paniculate; leaves

densely barbate beneath in the leaf axils. .Phoebe padiformis.

Inflorescence broadly paniculate; leaves not barbate beneath or

very obscurely so Nectandra Heydeana.

Flowers conspicuously and usually densely pubescent.

Leaves all or chiefly obtuse to almost rounded at the apex, sometimes
abruptly contracted into a short, very obtuse tip.

Leaf blades rounded or very obtuse at the base, mostly 10-12 cm.
wide Phoebe macrophylla.

Leaf blades acute or subacute at the base, mostly 3-7 cm. wide.
Petioles very thick and broad, marginate almost or quite to the

base Ocotea chiapensis.

Petioles slender, not at all marginate.

Leaves with large pits beneath in the axils of the nerves.

Phoebe may ana.

Leaves not with pits beneath, sometimes barbate in the axils
of the nerves.

Perianth 5-6 mm. long Phoebe ambigens.

Perianth 1.5-3.5 mm. long Ocotea veraguensis.

Leaves very acute to long-acuminate at the apex.

Branches of the inflorescence densely and minutely sericeous with
closely appressed hairs, or covered with a dense, very minute,
closely appressed tomentum.

Leaves subcordate or rounded at the base Ocotea perseifolia.

Leaves acute at the base Nectandra globosa.

Branches of the inflorescence glabrous or pubescent, the pubescence
mostly puberulent or hirtellous, the hairs chiefly spreading and
of appreciable length.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 307

Leaf blades broadest above the middle, long-attenuate to the base.

Ocotea eucuneata.

Leaf blades broadest at or below the middle, not long-attenuate
to the base.

Leaves barbate beneath in the leaf axils, or at least in most of
the axils.

Leaves blackish when dried, mostly 1.5-3 cm. wide.

Ocotea effusa.
Leaves green or brownish when dried, mostly 3-5.5 cm. wide.

Leaves usually with a few spreading hairs beneath along
the costa, the nerves somewhat impressed on the upper
surface, the leaves thus more or less bullate.

Phoebe longicaudata.

Leaves glabrous beneath, the nerves not at all impressed
on the upper surface, the leaves not at all bullate.

Nectandra sanguined.

Leaves not barbate beneath in the leaf axils.

Veins not at all elevated on the upper leaf surface.

Nectandra glabrescens.

Veins conspicuously elevated and reticulate on the upper

leaf surface.

Branches of the inflorescence glabrous or essentially so,
the inflorescence flexuous or recurved, often longer
than the leaves Ocotea laetevirens.

Branches of the inflorescence pubescent, the inflorescence
stiff, shorter than the leaves Ocotea Lundellii.

BEILSCHMIEDEA Nees

Reference: A. J. G. H. Kostermans, Rec. Trav. Bot. Ne'er!. 35:
837-865. 1938.

Trees or shrubs; leaves chartaceous to rigid-coriaceous, glabrous or pubescent,
penninerved, often pruinose beneath; panicles axillary or clustered near the ends
of the branches, usually short and few-flowered; involucre none; flowers perfect,
the perianth tube short, broadly obconic; perianth segments 6, subequal or the
outer ones shorter, deciduous; fertile stamens 9, free, the 6 outer ones with large
ovate anthers, the connective conspicuously produced beyond the large introrse
cells; filaments eglandular; 3 inner stamens with narrower thicker anthers; fila-
ments all with sessile basal glands; staminodia of series 4 large, ovate-acute or
triquetrous, short-stipitate or sessile; ovary subglobose, usually glabrous, the
style short, thick, obtuse; fruit generally ellipsoid, obtuse, often very large.

Fifteen species are known from tropical America, and others
occur in the Old World tropics. Two other species are known from
Costa Rica, and one of them, B. mexicana (Mez) Kosterm., which
occurs also in southern Mexico, is to be expected in the mountains
of Guatemala.

Leaves very densely areolate-reticulate or pitted on the upper surface, tomentose

beneath when young and with persistent pubescence in age B. Anay.

Leaves laxly and openly reticulate-veined on the upper surface, glabrous beneath.

B. hondurensis.

308 FIELDIANA: BOTANY, VOLUME 24

Beilschmiedea Anay (Blake) Kosterm. Rec. Trav. Bot. Ne"erl.
35: 847. 1938. Hufelandia Anay Blake, Journ. Wash. Acad. Sci. 9:
459. /. 1. 1919. Anay.

Wet mixed forest, 350-900 meters; Alta Verapaz; Suchitepe"quez
(type from Finca Compromise, Mazatenango, Wilson Popenoe 754).
Costa Rica; Colombia.

A large tree, as much as 20 meters high, with thick, reddish brown bark, the
young branches thick, densely ferruginous-tomentose or hirsute-tomentose; leaves
chartaceous, on stout petioles 2.5-3.5 cm. long, broadly elliptic to broadly ovate,
12-30 cm. long, 7.5-19 cm. wide, shortly obtuse-acuminate, rounded or short-
cuneate at the base, when young sparsely lanuginous-tomentulose, glabrate above
in age, pruinose beneath, laxly ferruginous-tomentulose or hirsute, the lateral
nerves 10-14 pairs; panicles crowded near the ends of the branches, pyramidal,
densely ferruginous-tomentose, 10-15 cm. long, on peduncles 4-7 cm. long, the
pedicels 2-5 mm. long; flowers pilose, 3-4 mm. long, the tube scarcely 1 mm. long;
perianth segments subequal, erect, densely pilose within, ovate or elliptic, 2.5-3
mm. long; anthers densely pilose, the filaments pilose; basal glands rather large,
subglobose; staminodia large, densely pilose, triangular-ovate; fruit ellipsoid-
pyriform, glossy black, thin-skinned, 10-15 cm. long; seed very large, obovoid.

Through an oversight, we did not investigate the occurrence of
this tree in Guatemala and have no recent material of it. According
to Wilson Popenoe, it grows wild in both the northern and southern
coasts, at low elevations. The fruit is shaped like a pear, the edible
flesh yellow, oily, and of rich flavor.

Beilschmiedea hondurensis Kosterm. Rec. Trav. Bot. Ne"erl.
35: 854. 1938.

Known only from the type, W. A. Schipp 1262, collected at Camp
31, on the boundary between Pete"n and British Honduras.

A small tree, the branchlets glabrous, the branches grayish; leaves alternate
or sometimes crowded at the base of the branchlets, chartaceous, glabrous, some-
what lustrous, lance-elliptic to elliptic or obovate-elliptic, acuminate, acute at
the base, conspicuously and laxly prominulous-reticulate on both surfaces, the
lateral nerves 9-12 pairs, the petioles slender; fruiting panicles 4 cm. long, glabrous;
fruit black, ellipsoid, smooth, 3 cm. long, 1.5 cm. broad, the supporting pedicel
3 mm. long and 2 mm. thick.

CASSYTHA L.

Plants parasitic, scandent, yellowish, herbaceous, perennial, the stems
slender, twining, attached to the host by 1-seriate haustoria; leaves reduced to
small scales or absent; flowers small, greenish or whitish, sessile or pedicellate in
the axil of a scale-like bract and with small bractlets at the base of the perianth,
arranged in mostly pedunculate racemes, spikes, or heads; perianth tube small,
accrescent in fruit and becoming constricted at the apex; perianth segments 6,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 309

the outer ones short, broad, resembling the bracts, the 3 inner ones twice as long;
perfect stamens usually 6 and 2-celled, the 2 outer rows with introrse anthers and
eglandular filaments, the inner ones with extrorse anthers and 2-glandular fila-
ments; staminodia large, subsessile or stipitate; fruit globose, completely included
in the enlarged perianth tube, the perianth segments usually persistent; seed with
a thin testa, the cotyledons carnose, distinct only when young, completely con-
crescent when ripe and having the appearance of carnose endosperm.

Species about 20, one pantropic, the others in tropical Africa,
southern Asia, and Australia.

Cassytha filiformis L. Sp. PI. 35. 1753. C. americana Nees,
Syst. Laur. 644. 1836. Suelda con suelda (Pete'n).

Parasitic on herbs and low shrubs, 300 meters or less, usually
most plentiful near the seashore; Pete'n; Alta Verapaz; Izabal;
Huehuetenango. Southern Florida; southern Mexico; British Hon-
duras, along the Atlantic coast to Panama; West Indies; tropical
South America; widely dispersed in the Old World tropics.

Plants very slender, glabrous or nearly so, scandent, sometimes 3 meters long
but usually much smaller, often forming dense tangles, pale green or yellowish
green; leaves reduced to minute scales; flowers spicate, the spikes lax, usually
solitary in the axil of 3 bracts, slightly or densely tomentulose, 1.5-5 cm. long, the
peduncles 1-3 cm. long; bracts membranous, ovate-lanceolate, 2 mm. long or
shorter, the inner ones ciliate; flowers sessile, glabrous, white or whitish, 2.5 mm.
long; perianth tube almost obsolete, the segments unequal, the outer ones ovate-
orbiculate, ciliate, the inner ovate, obtuse, not ciliate, 2.5 mm. long; stamens
included, glabrous; anthers ovate-triangular; fruit globose, 6 mm. in diameter.

In general appearance the plant is almost like a species of Cuscuta,
this appearance being very deceptive, for the flowers, of course, are
altogether unlike in the two genera.

CINNAMOMUM Burman

Shrubs or large trees, usually with aromatic bark and leaves; leaves coria-
ceous, persistent, opposite or sometimes alternate, triplinerved or penninerved;
panicles axillary or terminal, often congested, the bracts very small or none;
flowers rather small, perfect or by abortion polygamous, the pistillate flowers then
larger; perianth tube funnelform, the segments 6, deciduous at or above the base,
rarely persistent; fertile stamens 9 or fewer; filaments of the 2 outer rows of stamens
eglandular, the anthers introrse, 4-celled; filaments of the third row with stipitate
or subsessile glands, the anthers extrorse, generally 4-celled; anther cells in 2
vertical rows, the upper ones smaller; filaments slender, mostly equaling the ovate
or oblong anthers; staminodia of the fourth row ovate or oblong, cordate or sagit-
tate, stipitate, eglandular; ovary sessile, narrowed into a long slender style, the
stigma obtuse or depressed; fruit usually ellipsoid, the cupule with an entire
margin.

310 FIELDIANA: BOTANY, VOLUME 24

About 130 species, in the tropics of eastern Asia, Australia, and
the Pacific islands. Two of them are often cultivated in tropical
America.

Leaves long-acuminate; fruit less than 1 cm. long C. Camphora.

Leaves acute or obtuse; fruit 1.5 cm. long C. zeylanicum.

Cinnamomum Camphora (L.) Nees & Eberm. Med. Pharm.
Bot. 2: 430. 1831. Laurus Camphora L. Sp. PL 369. 1753. Alcanfor.

Native of eastern Asia, especially of Formosa; planted occasion-
ally for ornament or as a curiosity about Guatemala City, in Alta
Verapaz, and probably elsewhere, but the individuals few.

A small or medium-sized tree, 12 meters high or less, with a dense crown, in
cultivation often only a shrub, glabrous throughout or nearly so; leaves alternate,
coriaceous, on rather long, slender petioles, broadly ovate to lance-oblong, mostly
7-11 cm. long, triplinerved, acuminate or long-acuminate, acute to attenuate at the
base, lustrous on the upper surface; panicles axillary, shorter than the leaves, the
flowers small, yellowish, the branchlets 1-3-flowered; perianth 3 mm. long.

The plant may be recognized readily by the camphor odor of the
crushed leaves. Commercial camphor, which is extracted from the
wood, is produced almost exclusively on the island of Formosa.

Cinnamomum zeylanicum Breyne, Eph. Nat. Cur. Dec.
Ann. 4: 139. 1789. Laurus Cinnamomum L. Sp. PI. 369. 1753.
Canela. Cinnamon.

Native of southeastern Asia, but much planted in other regions
for its bark, source of the cinnamon of commerce; grown occasion-
ally in the mountains of Guatemala for shade or ornament or as a
.curiosity, and planted on a commercial scale in Alta Verapaz, as at
Cubilguitz.

A tree, sometimes 20 meters high, the bark rather thick, reddish inside, pale
outside, the branches glabrous; leaves opposite or subopposite, rarely alternate,
coriaceous, lustrous, ovate to lance-oblong, mostly 6-15 cm. long, acute to very
obtuse, abruptly contracted at the base, conspicuously triplinerved, glabrous, the
young leaves pink, the petioles 1-2.5 cm. long; panicles terminal or subterminal,
pubescent or glabrous, lax, on long slender peduncles, the pedicels 3-4 mm. long,
pubescent; perianth segments 4-7 mm. long, sericeous, oblong or obovate, obtuse;
fertile stamens 9; fruit dark purple, 8-12 mm. long, ellipsoid.

Fairly extensive plantations of cinnamon trees have been made
in the wet lower mountains of Guatemala, apparently are thriving,
and ultimately may afford an important export. Cinnamon is one
of the favorite condiments of Guatemalan (and other Central Ameri-
can) cooks, who consider that almost any dish can be improved by
it. It enters with monotonous persistence into almost every dessert.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 311

LICARIA Aublet

Reference: A. J. G. H. Kostermans, The genus Licaria, with notes
on Phyllostemonodaphne and Dryadodaphne, Rec. Trav. Bot. Ne"erl.
34: 575-605. 1937.

Trees or shrubs, glabrous or pubescent; leaves alternate or opposite, thin-
chartaceous to rigid-coriaceous, penninerved; flowers perfect, in axillary and sub-
terminal panicles, rarely solitary, subumbellate, or capitate; involucre none;
perianth tube usually distinct, rarely shallow; perianth segments 6, biseriate,
equal or unequal, spreading or incurved; stamens of the 2 outer rows modified
into small staminodia or abortive, the stamens of the third series fertile, free,
partly connate into a stamen tube, the filaments distinct or none; anthers 2-celled,
the cells introrse, extrorse, or extrorse-apical, the valves dehiscent from the base
upward; stamens of the fourth series usually abortive, rarely reduced to stami-
nodia, minute, stipe-like; ovary free, ellipsoid or globose-obovoid, glabrous or
pilose; style usually slender, the stigma inconspicuous, truncate or obtuse; fruit
ellipsoid, smooth, mucronulate; cupule hemispheric, with a double or triple margin,
the inner margin erect, thin, the outer one spreading, thick, irregular; cotyledons
flat-convex, large.

Species about 40, in tropical America, mostly in South America.
Three other species have been found in Central America. This
genus may be recognized readily by its fruit, which has two distinct
margins on the cupule, one the normal inner one, usually erect, and,
at a short or longer distance below it, a second one with a spreading,
usually irregular margin. Such a structure is found rarely, if ever,
in other local genera. In the other larger genera such as Nectandra,
Ocotea, and Phoebe, the fruits have no distinctive characters.

Adult leaves velutinous-pilose beneath with dense short spreading hairs.

L. Peckii.
Adult leaves glabrous beneath, or with minute, closely appressed hairs.

Mature leaves covered beneath with minute, closely appressed hairs, the blades

mostly 1.5-2.5 cm. wide L. campechiana.

Mature leaves glabrous beneath, the blades usually much wider.

Flowers capitate at the ends of long slender simple peduncles, the heads

solitary or 2-3 L. capitata.

Flowers scarcely capitate, the peduncles branched.

Stamens connate L. Cervantesii.

Stamens free.

Branchlets hirsute-tomentose L. caudata.

Branchlets glabrous or nearly so L. coriacea.

Licaria campechiana (Standl.) Kosterm. Rec. Trav. Bot.
Ne"erl. 34: 599. 1937. Ocotea campechiana Standl. Carnegie Inst.
Wash. Publ. 461: 56. 1935. Chanekia campechiana Lundell, Phy-
tologia 1: 178. 1935. Misanteca campechiana Lundell, Carnegie

312 FIELDIANA: BOTANY, VOLUME 24

Inst. Wash. Publ. 478: 209. 1937. Phoebe campechiana Standl. ex
Lundell, op. cit. 436: 281. 1934, as syn. Dzol, Ectit (Pet&i, Maya,
fide Lundell); Granadilla (Huehuetenango) ; Copal-chi (Pete"n, fide
Lundell).

Dense or rather thin, moist or wet forest or thickets, often or
usually on limestone, 1,400 meters or less; Pete"n; Alta Verapaz;
Izabal; Huehuetenango. Campeche; British Honduras.

A large shrub or usually a tree of 8-25 meters, the trunk as much as 45 cm. in
diameter, the branches slender, densely and minutely grayish-sericeous, the older
branches gray; leaves coriaceous or chartaceous, on slender petioles 5-8 mm. long,
lance-elliptic to narrowly lanceolate, 4-11 cm. long, 1.5-3 cm. wide, very narrowly
long-acuminate or attenuate, acute at the base, when young laxly sericeous, in
age glabrous above, very lustrous, the nerves and veins not at all elevated, paler
beneath, densely and minutely sericeous or finally glabrate, the costa slender,
elevated, the lateral nerves 11-18 pairs, inconspicuous; panicles axillary, minutely
tomentulose, rather few-flowered, 2-6 cm. long, on slender peduncles 1-2.5 cm.
long; flowers glomerate, the pedicels 1-2 mm. long; perianth subglobose, minutely
tomentulose, 1.5-2 mm. long, the segments carnose, slightly incurved, pilose
within, the outer ones acute, the inner ones smaller; stamens partly connate, the
anthers ovate-elliptic, obtuse, the cells large, extrorse; filaments shorter than the
anthers, slightly pilose, the basal glands small, free; ovary glabrous; fruit about
1 cm. long, oval, the cupule depressed, 8 mm. broad, the margin ciliate, the outer
margin almost regular, very narrow.

Called "laurelillo" in Campeche. The flowers are white or
whitish or sometimes tinged with pink. The wood is lustrous reddish
brown, rather fine-textured, hard, heavy, and strong, apparently
suited for general construction but probably not available in sufficient
quantities for commercial exploitation.

Licaria capitata (Cham. & Schlecht.) Kosterm. Rec. Trav.
Bot. Ne'er!. 34: 592. 1937. Misanteca capitata Cham. & Schlecht.
Linnaea 6: 367. 1831. Acrodiclidium gldbrum Brandeg. Univ. Calif.
Publ. Bot. 6: 497. 1919.

Moist or wet, mixed forest, 1,500 meters or less; reported from
Pete'n; Alta Verapaz; Izabal. Southern Mexico; British Honduras;
Honduras.

A large shrub or a tree, sometimes 12 meters high, the branches grayish, when
young minutely tomentulose or puberulent, the older branches gray; leaves
alternate, usually rigid-coriaceous, on petioles 1-2 cm. long, elliptic to elliptic-
oblong, 12-30 cm. long, 4-10 cm. wide, rather abruptly acute or short-acuminate,
acute at the base, somewhat pulverulent-tomentulose when young but soon gla-
brous, lustrous above, minutely areolate-reticulate or smooth, slightly paler
beneath, the lateral nerves 8-11 pairs, densely areolate-reticulate; inflorescences
clustered at the base of the young branchlets, the peduncles 4-8 cm. long, densely

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 313

and minutely puberulent-tomentulose, bearing a single head of flowers about 1-1.5
cm. in diameter, the flowers sessile, densely tomentulose, 2.5-3 mm. long; perianth
tube urceolate-cylindric, glabrous within; perianth segments erect, ovate-triangu-
lar, glabrous within; stamen tube exserted from the perianth, the filaments pilose;
glands 6; ovary glabrous, the style slender, elongate; fruit ellipsoid, smooth,
mucronulate, 2.5 cm. long and 1.5 cm. broad or smaller; cupule deep red, hemi-
spheric, verruculose, as much as 3 cm. broad and 2 cm. high, the outer margin
thick, irregular, the inner one thin, erect, entire.

Called "aguacatillo" in Honduras.

Licaria caudata (Lundell) Kosterm. Rec. Trav. Bot. Ne"erl. 34:
596. 1937. Chanekia caudata Lundell, Phytologia 1: 178. 1937.

Known only from the type, Pete"n, Camp 32 on the boundary of
British Honduras, 700 meters, W. A. Schipp 1279.

A tree of 7-10 meters, the trunk 7-15 cm. in diameter, the branches slender,
hirsute-tomentose with gray or yellowish hairs, the older branches gray; leaves
alternate, chartaceous, 5-9 cm. long, 1.5-3 cm. wide, on slender petioles 4-7 mm.
long, lanceolate or lance-elliptic, with a long narrow caudate acumination, acute
at the base, the acumen obtuse, glabrous in age except for a few scattered hairs
along the costa beneath, dull, the lateral nerves 6-10 pairs; panicles axillary or
internodal, slender, lax, scarcely branched, few-flowered, 2-3.5 cm. long, on slender
peduncles 1.5-2 cm. long, the slender pedicels 3-5 mm. long, glabrous; flowers
white, subglobose, 1.5-2 mm. long, glabrous, the perianth tube hemispheric,
densely hirsute within; perianth segments erect, subequal, ovate-orbicular, acute,
pilose within; stamens included, hirsute; anthers glabrous, broader than long;
basal glands small, free, orbicular; ovary glabrous, the style usually slightly
exserted beyond the stamens.

Licaria Cervantesii (HBK.) Kosterm. Rec. Trav. Bot. Ne"erl.
34: 587. 1937. Laurus Cervantesii HBK. Nov. Gen. & Sp. 2: 168.
1818. Misanteca Juergensenii Mez, Jahrb. Bot. Gart. Berlin 5: 102.
1889.

Reported from Alta Verapaz ("Matacui," J. D. Smith 1650).
Southern Mexico.

A tree, the branches glabrous, the older ones grayish brown; leaves alternate,
chartaceous, on slender petioles 10-12 mm. long, glabrous, slightly lustrous,
elliptic or rarely broadly elliptic, 11-20 cm. long, 3-8 cm. wide, densely promi-
nulous-areolate on both surfaces, acuminate, narrowed at the base, the lateral
nerves 8-12 pairs; panicles axillary, rather few-flowered, 3-8 cm. long, the pedun-
cles 2-5 cm. long, the branches few, spreading or erect-spreading, sparsely and
minutely tomentulose, the thick pedicels 1 mm. long or shorter; flowers white,
clustered at the ends of the branchlets, 1.5-2 mm. long; perianth tube pulveru-
lent-tomentulose, funnelform, the segments glabrous, erect, ovate-orbicular, acute,
the inner ones narrower; anthers glabrous, triangular, subacute, the filaments
pilose; basal glands free, liguliform; ovary glabrous, the style slender, cylindric;
fruit ellipsoid, as much as 22 mm. long and 15 mm. broad, mucronulate; cupule

314 FIELDIANA: BOTANY, VOLUME 24

hemispheric, verruculose, about 1 cm. high and 2 cm. broad, the outer margin
spreading, thick, irregular, the inner one thin, entire, erect, as much as 5 mm. high.

Licaria coriacea (Lundell) Kosterm. Rec. Trav. Bot. Ne'er!. 34:
604. 1937. Chanekia coriacea Lundell, Phytologia 1: 179. 1937.

Moist or wet, mixed forest, 2,000 meters or less; Pete"n (type from
Camp 31, British Honduras boundary, W. A. Schipp 1282); Alta
Verapaz; Izabal; Zacapa. British Honduras.

A tree of 9-12 meters, the trunk as much as 25 cm. in diameter, glabrous
throughout; leaves on petioles 6-10 mm. long, lanceolate or oblong-lanceolate,
5.5-11 cm. long, 2-4 cm. wide; acuminate with a subobtuse tip, subacute at the
base, coriaceous, the costa prominent, the lateral nerves inconspicuous; inflores-
cences axillary, producing a single fruit, the peduncles 1-3 cm. long, stout; cupule
shallow, verruculose, 1.5-2.5 cm. broad, 1 cm. high; fruit ellipsoid, 17 mm. long,
12 mm. broad, short-apiculate.

The flowers are white, the ripe fruit black, the pedicels red in age.

Licaria Peckii (I. M. Johnston) Kosterm. Rec. Trav. Bot.
Ne'er!. 34: 597. 1937. Misanteca Peckii I. M. Johnston, Contr.
Gray Herb. 70: 70. 1924. Chanekia Peckii Lundell, Phytologia 1:
178. 1937. Senc-cul (Alta Verapaz).

Moist or wet, mixed forest, often or usually on limestone, 400
meters or less; Pete"n; Alta Verapaz; Izabal. British Honduras;
type M. E. Peck 826, without definite locality.

A tree 9-12 meters high or doubtless even taller, the trunk 25-30 cm. or more
in diameter, the branchlets densely hirsute- villous; leaves on petioles 12 mm. long
or shorter, oblanceolate to oblong-obovate, mostly 9-18 cm. long, abruptly
acuminate or long-acuminate, cuneate or subobtuse at the base, coriaceous,
lustrous above, glabrous in age or nearly so, with sunken nerves and thus some-
what bullate, densely and softly pilose or setose-pilose beneath with spreading
hairs, the lateral nerves about 9 pairs; inflorescences racemose or racemiform,
2-3 cm. long or larger, densely brownish- velutinous, few-flowered, lax; flowers
yellowish or greenish white, 2 mm. broad, 1.5 mm. high, the pedicels 1 mm. long
or shorter; perianth segments glabrous, the outer ones very broadly triangular;
fruit ellipsoid, purple-black, about 2 cm. long and 1 cm. broad; cupule reddish or
rose, 1.5 cm. broad.

Called "timber sweet" in British Honduras. The tree has been
reported from British Honduras as Phoebe helicterifolia Mez.

LITSEA Lamarck

Reference: H. H. Bartlett, A synopsis of the American species of
Litsea, Proc. Amer. Acad. 44: 597-602. 1909.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 315

Trees or shrubs, glabrous or pubescent, sometimes glaucous; leaves alternate
or rarely subopposite, penninerved or triplinerved, usually coriaceous; flowers
dioecious, umbellate or capitate, the inflorescences before anthesis included in a
globose involucre, this pedunculate or sessile, the involucres arranged in sessile
or short-pedunculate fascicles or in axillary or lateral racemes; bracts of the
involucre 4-6, decussate-opposite; flowers mostly 4 or 6 in each involucre, the
staminate ones sessile or short-pedicellate; perianth tube ovoid, campanulate, or
almost obsolete; lobes of the limb 6 or 4 or by abortion fewer, rarely minute or
none; stamens in the staminate flower and staminodia in the pistillate flower
usually 9 or 12, those of the first and second series usually eglandular, those of the
third and fourth series, when present, often with a stipitate gland at the base;
filaments usually slender; anthers introrsely 4-locellate; ovary included in the
perianth tube or exserted, attenuate into a short or long style, irregularly somewhat
lobate; fruit surrounded at the base by the unchanged or somewhat accrescent,
cupular base of the perianth tube.

About 100 species, mostly in Asia and Australia, about a dozen
in America. One other Central American one is found in the moun-
tains of Costa Rica.

Leaves pubescent beneath.
Leaves strigose or sericeous beneath with closely appressed hairs.

L. guatemalensis.

Leaves tomentose beneath with lax, more or less spreading, not appressed hairs.

L. Neesiana.

Leaves glabrous beneath L. glaucescens.

Litsea glaucescens HBK. Nov. Gen. & Sp. 2: 168. 1817.
Tetr anther a glaucescens var. subsolitaria Meissn. in DC. Prodr. 15,
pt. 1 : 193. 1864. L. glaucescens var. subsolitaria Hemsl. Biol. Centr.
Amer. Bot. 3: 76. 1882. L. acuminatissima Lundell, Contr. Univ.
Mich. Herb. 4: 3. 1940. L. Matudai Lundell, op. cit. 4 (type from
Volcan de Tacana, Chiapas, E. Matuda 2933). Laurel.

Moist or dry, brushy hillsides, or most often in rather open,
mixed or pine-oak forest, 1,300-3,500 meters; Alta Verapaz; Baja
Verapaz; Zacapa; Chiquimula; Jutiapa; Huehuetenango; Que-
zaltenango; San Marcos. Mexico; Salvador; Honduras.

A shrub or tree, usually 3-12 meters high, rather densely branched, the
branches glabrous or puberulent, slender; leaves coriaceous, on slender petioles
18 mm. long or less, lanceolate or elliptic-lanceolate, 8 cm. long and 2.5 cm. wide
or smaller, glabrous, acute to long-acuminate, acute or subacute at the base, penni-
nerved or obscurely triplinerved, closely and conspicuously reticulate-veined,
lustrous, glaucescent or green beneath; inflorescences axillary, solitary or fascicu-
late, simple or corymbose, the peduncles glabrous, the involucres 5-9-flowered,
the flowers yellow; pedicels glabrous or nearly so; perianth tube none, the lobes
oval, subobtuse or subacute, thin, glabrous; stamens 3-seriate, the filaments
glabrous; fruit globose, black, 9 mm. in diameter.

316 FIELDIANA: BOTANY, VOLUME 24

The leaves of this and other species have an aromatic odor
similar to that of bay leaves (Laurus), and they are much used in
Guatemala for flavoring food of many kinds, especially soup and
meat. Bunches of leafy branches or of dried leaves are on sale in
most of the markets. The trees sometimes are cultivated in the
gardens of Coban on this account. Newly cut branches covered
with leaves are much used for decorations at fiesta times, especially
for making the arches that span streets and roads. The species is a
slightly variable one but we are unable to distinguish the segregates
from it that have been proposed recently, based upon the simple or
corymbose nature of the peduncles and the presence or absence of a
glaucous tinge on the lower leaf surface. Apparently both these
characters are variable and are unsuitable as a basis for specific
segregation.

Litsea guatemalensis Mez, Jahrb. Bot. Gart. Berlin 5: 479.
1889. Laurel; Aguarel (Jalapa).

Dense, moist or wet, mixed forest or often in open pine forest or
in thickets, 1,500-3,150 meters; endemic; Jalapa; Guatemala;
Sacatepe"quez; Chimaltenango; Solola (type from Godinez, Hartweg
613).

A shrub or small tree, seldom more than 6 meters high, the branches slender,
brown or brownish, when young velutinous-pubescent; leaves coriaceous, on
petioles 1.5 cm. long or shorter, lanceolate or elliptic-lanceolate, about 8 cm. long
and 2.5 cm. wide, acuminate or long-acuminate, acute or subacute at the base,
lustrous above and glabrous, penninerved, much paler beneath, sparsely or densely
strigose or in age often glabrate; peduncles simple, axillary, solitary or fasciculate,
tomentulose, 15 mm. long or less, 5-11-flowered; bracts of the involucre deciduous,
tomentulose, the pedicels strigillose, slightly longer than the flowers; perianth tube
none, the segments oval, subobtuse; filaments glabrous, half as long as the anthers.

Litsea Neesiana (Schauer) Hemsl. Biol. Centr. Amer. Bot. 3:
76. 1882. Tetranthera Neesiana Schauer, Linnaea 19: 712. 1847.
Laurel; Spac-tze (Huehuetenango).

Moist or dry, often rocky, brushy hillsides, often in oak forest,
1,900-3,000 meters; Solola; reported from Quiche"; Huehuetenango;
Quezaltenango. Southern Mexico.

A shrub or tree 3-9 meters high, the branches slender, terete, the young ones
densely reddish- or brownish-tomentose; leaves coriaceous, on rather slender
petioles 2 cm. long or shorter, lustrous, ovate to ovate-lanceolate or narrowly
lanceolate, about 6.5 cm. long and 2.5 cm. wide, acuminate or acute, acute or
obtuse at the base, at first densely ochraceous-tomentose, in age green and
glabrate above, the pubescence persistent on the lower surface; inflorescences

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 317

3-7-flowered, mostly axillary and simple, the peduncles 1.5 cm. long or less,
tomentulose; bracts of the involucre deciduous, more or less tomentose outside;
perianth tube none, the segments more or less lanceolate, subobtuse, thin; fila-
ments glabrous, longer than the anthers; ovary glabrous; fruit black, globose,
8-9 mm. in diameter.

NECTANDRA Rolander

Large or small trees, rarely shrubs, glabrous or pubescent; leaves alternate
or rarely opposite, coriaceous or chartaceous; panicles pyramidal or subcorymbose,
rarely racemose, mostly axillary; flowers without an involucre, generally rather
large, perfect or dioecious; perianth tube conspicuous or almost none, the segments
6, usually spreading, equal or nearly so, deciduous; fertile stamens 9, those of the
fourth series reduced to staminodia, small, or wanting; anthers 4-celled, usually
papillose, the cells in a horizontal, straight or slightly arcuate row, those of the
outer 6 stamens mostly introrse, those of the third row extrorse; filaments of the
2 outer rows of stamens usually short or none, those of the third row with 2 sessile
glands; ovary globose or ellipsoid, commonly glabrous; style usually short, rarely
longer than the ovary; fruit globose or ellipsoid; cupule with a simple entire margin,
saucer-shaped to hemispheric.

About 90 species, in tropical America, most numerous in South
America. A few besides those listed here are known from southern
Central America. Some trees of this genus furnish valuable lumber,
especially in case of certain South American species. In general the
heartwood is greenish yellow to dark olive-brown; luster usually
silky or silvery; odor spicy or resinous, the taste mild to pronounced;
rather light and soft to moderately hard and heavy, the specific
gravity usually 0.60-0.75; texture medium to somewhat coarse, the
grain straight to roey; seasons readily without splitting.

Leaves densely and softly pilose beneath with conspicuous spreading hairs, some-
times densely tomentose.

Margins of the leaves conspicuously recurved at the base; leaves long-acumi-
nate N. reticulata.

Margins of the leaves not recurved; leaves rounded or very obtuse at the apex

and abruptly apiculate or apiculate-acuminate N. sinuata.

Leaves glabrous beneath or the pubescence minute and appressed.

Leaf blades finely sericeous, puberulent, or strigillose beneath, the pubescence
persistent in age, usually inconspicuous to the naked eye but evident under
a lens.
Leaves acute or subobtuse, mostly elliptic-oblong AT. surinamensis.

Leaves narrowly long-acuminate, lanceolate or oblong-lanceolate.

N. membranacea.

Leaf blades glabrous beneath or practically so, at least at maturity.

Flowers glabrous or essentially so 2V. Heydeana.

Flowers conspicuously and usually densely pubescent.

318 FIELDIANA: BOTANY, VOLUME 24

Anthers of the outermost stamens on conspicuous filaments; leaves barbate
beneath in the axils of the nerves; branches of the inflorescence gla-
brous, puberulent, or hirtellous N. sanguined.

Anthers of the outer stamens sessile.

Branches of the inflorescence usually minutely sericeous, the hairs all
or mostly appressed; leaves usually barbate beneath in the axils of

the nerves N. globosa.

Branches of the inflorescence not sericeous, sometimes almost glabrous,
but often puberulent or short-hirtellous; leaves not barbate beneath.

N. glabrescens.

Nectandra glabrescens Benth. Bot. Voy. Sulph. 161. 1844.
Aguacatillo (reported from Izabal); Pubabac (Alta Verapaz; deter-
mination uncertain).

Moist or wet, mixed forest, 1,400 meters or less; Alta Verapaz;
Izabal; Retalhuleu; Quezaltenango. Southern Mexico; British
Honduras to Panama; Colombia.

A tree, sometimes 18 meters high with a trunk 45 cm. in diameter, the branch-
lets at first minutely tomentulose, soon glabrate; leaves chartaceous, dull or some-
what lustrous, elliptic to ovate-lanceolate, mostly 12-20 cm. long and 4-7 cm.
wide, acuminate or long-acuminate, acute at the base, glabrous or practically so,
at least at maturity; inflorescences many-flowered, laxly corymbose-paniculate,
puberulent or almost glabrous, shorter than the leaves; flowers white or whitish,
tomentulose, 9-10 mm. broad or sometimes smaller, perfect; perianth tube very
short or obsolete, the segments oval, obtuse; anthers sessile, papillose, depressed-
triangular; staminodia small, stipe-like, glabrous; ovary glabrous.

Known in British Honduras by the names "laurel" and "sweet-
wood"; "pimiento" (Salvador).

Nectandra globosa (Aubl.) Mez, Jahrb. Bot. Gart. Berlin 5:
415. 1889. Laurus globosa Aubl. PI. Guian. 364. 1775. Canoj;
Zunonte, Sacalante (Pete"n, fide Lundell) ; Coyokiche (reported as the
Quecchi name).

Moist or wet, mixed forest, sometimes in pastures or along road-
sides, often on limestone, 1,500 meters or less; Peteri; Alta Verapaz;
Santa Rosa; Escuintla; Guatemala; Sacatepe"quez; Suchitepe"quez;
Retalhuleu. British Honduras to Salvador and Panama; northern
South America.

A small to large tree, commonly 9-15 meters high or sometimes much larger,
with a thick trunk, the young branchlets mostly sericeous with grayish or brownish
hairs, soon glabrate; leaves coriaceous or chartaceous, on petioles 1-1.5 cm. long,
subopposite or alternate, lanceolate or ovate-lanceolate, mostly 14-22 cm. long
and 4-8.5 cm. wide, acuminate or long-acuminate, acute at the base, glabrous and
lustrous above, the venation little if at all elevated, paler beneath, glabrous or
practically so except when very young, the venation prominulous, laxly reticulate;

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 319

inflorescence corymbose-paniculate, minutely sericeous or glabrate, many-flowered,
much shorter than the leaves, the pedicels 1-5 mm. long; flowers perfect, white,
fragrant, 1 cm. broad or somewhat smaller, larger than in most other local species,
tomentulose; perianth tube conspicuous, suburceolate, the segments broadly
ovate or oval, obtuse or rounded at the apex, papillose-tomentulose within; anthers
sessile, papillose, those of the outer series subacute or obtuse, those of the third
series attenuate, with globose sessile glands; ovary glabrous; fruit ellipsoid or
ovoid, about 1 cm. long, black or purple-black, the cupule short, with a simple
entire margin.

Sometimes called "wild pear" and "timber sweet" in British
Honduras; "aguacatillo," "sangre blanca" (Honduras); "aguacate
de monte" (Salvador). One of the common trees of the lowlands of
Guatemala and other countries of Central America, sometimes
plentiful along stream banks. It is not altogether certain that the
name used here is the correct one, but it is employed in the sense in
which it was applied by Mez. The genus has not been monographed
recently, and the proper terminology for some of the commonest
species of the genus is decidedly uncertain. The specific names here
used are those of Mez's monograph of the American Lauraceae, and
are mostly associable with material determined at one time or
another by him.

Nectandra Heydeana Mez & Bonn. Smith, Bot. Gaz. 19: 262.
pi. 25. 1894. Phoebe platyphylla Lundell, Contr. Univ. Mich. Herb.
6: 23. 1941 (type from Chiapas).

Known in Guatemala only from the original locality, city of
Santa Rosa, Santa Rosa, 900 meters, the type being Heyde & Lux
4260. Chiapas.

A large shrub or a tree 4-12 meters high, the trunk sometimes 60 cm. in
diameter, the branchlets glabrous, slender; leaves on slender petioles 2 cm. long
or shorter, membranaceous or thick-membranaceous, elliptic to lance-oblong,
mostly 10-20 cm. long and 4-11 cm. wide, obtuse to short-acuminate with an
obtuse tip, usually olivaceous when dry, somewhat lustrous, glabrous or nearly
so in age but usually barbate in the axils of the nerves, rounded to broadly cuneate
at the base, when young somewhat strigillose, laxly prominulous-reticulate on both
surfaces; inflorescence subcorymbose or subpyramidal, shorter than the leaves,
lax, with rather numerous flowers, glabrous; flowers perfect, glabrous, about 7
mm. in diameter, the perianth segments elliptic, rounded or very obtuse at the
apex, spreading; anthers of the outer series of stamens sessile, those of the inner
series on very thick filaments, these with 2 minute sessile globose basal glands;
outer anthers suborbicular, rounded at the apex, papillose; ovary globose, glabrous,
about equaling the stout style.

Nectandra membranacea (Swartz) Griseb. Fl. Brit. W. Ind.
282. 1862. Laurus membranacea Swartz, Prodr. Veg. Ind. Occ. 65.

320 FIELDIANA: BOTANY, VOLUME 24

1788. N. Gentlei Lundell, Contr. Univ. Mich. Herb. 6: 13. 1941 (type
from Stann Creek, Mullins River, British Honduras). N. perdubia
Lundell, Lloydia 4: 47. 1941 (based in part on Pete"n material).
Coajche (fide Aguilar); Zunonte (Maya?), Laurel, Laurel bianco
(Pet&i, fide Lundell).

Moist or wet, mixed forest or in second-growth thickets, often
on limestone, 1,200 meters or less; Pete"n; Alta Verapaz; Huehue-
tenango. British Honduras to Honduras and Panama; West Indies.

Usually a tree of 9-20 meters, the young branches thinly tomentulose or
sericeous or glabrate; leaves on petioles 1.5 cm. long or less, coriaceous or sub-
coriaceous, lanceolate or narrowly lanceolate to ovate-lanceolate, mostly 12-22
cm. long and 3-6 cm. wide, narrowly long-acuminate, acute or obtuse at the base,
glabrous or glabrate above, little if at all lustrous, usually densely and minutely
sericeous beneath or appearing glabrous to the naked eye, the venation not elevated
on the upper surface, sometimes impressed, little elevated on the lower surface,
laxly reticulate; inflorescence pyramidal-paniculate, tomentulose or almost wholly
glabrous; flowers white, perfect, tomentulose or almost glabrous, 4-5 mm. broad;
perianth tube conspicuous, the segments ovate or elliptic, subacute or obtuse;
filaments about equaling the anthers, glabrous, those of the third series with large
globose sessile glands at the base; anthers suborbicular, truncate or subemarginate
at the apex; staminodia stipe-like; ovary glabrous; fruit black, globose, about

1 cm. in diameter, the cupule small and shallow.

Nectandra reticulata (Ruiz & Pavon) Mez, Jahrb. Bot. Gart.
Berlin 5: 404. 1889. Laurus reticulata Ruiz & Pavon, Fl. Peruv. 4:
pi 348. 1802. Chuala (Alta Verapaz) ; Canoj.

Moist or wet, mixed forest, or sometimes in rather dry thickets
or second growth, 900 meters or less; Retalhuleu; Quiche"; Quezal-
tenango; San Marcos. Southern Mexico; Costa Rica; Panama;
tropical South America.

A tree of 6-25 meters, the trunk sometimes 75 cm. in diameter, with smooth
gray bark, the branches ferruginous- villous or tomentose; leaves on stout petioles

2 cm. long or mostly shorter, coriaceous, lanceolate to elliptic or lance-ovate,
mostly 17-35 cm. long and 4-10 cm. wide, acuminate or long-acuminate, some-
what narrowed to the base but the base itself usually rounded or subcordate and
with incurved margins often forming a short pocket, tomentulose or pilose above
or in age glabrate, densely and softly pilose beneath or brownish-tomentose, the
venation usually impressed on the upper surface, very prominently and laxly
reticulate beneath; inflorescence pyramidal-paniculate or corymbose-paniculate,
villous or tomentose, many-flowered, shorter than the leaves, the pedicels 3-8
mm. long; flowers white, perfect, 10-14 mm. broad, villous or tomentose; perianth
tube almost obsolete, the segments equal, broadly elliptic or suborbicular, obtuse;
outer anthers sessile, acute, papillose, those of the third series on short filaments,
laterally dehiscent; staminodia small, pilose, liguliform; ovary glabrous or very
sparsely pilose, the style elongate; fruit ellipsoid, 13 mm. long, 8 mm. broad, the
cupule short and spreading.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 321

Nectandra sanguinea Rottb. Act. Litt. Univ. Hafn. 1: 279.
1778. Aguacatillo, Laurel bianco (Pete"n).

Moist or wet, mixed forest, often on limestone, 800 meters or less;
Pete"n; Alta Verapaz; Baja Verapaz; Izabal; El Progreso; Que-
zaltenango. Southern Mexico; British Honduras to Panama; West
Indies; northwestern South America.

A tree of 9-12 meters or sometimes larger, the branchlets tomentulose or
glabrate, dark brown or grayish; leaves on petioles 1.5 cm. long or usually shorter,
mostly elliptic to obovate-lanceolate or obovate, averaging about 13 cm. long and
4 cm. wide, usually acute or acuminate, acute at the base, glabrous or practically
so but usually barbate beneath in the axils of the nerves, generally very lustrous
on the upper surface, somewhat paler beneath, laxly prominulous-reticulate on
both surfaces; inflorescences paniculate or corymbose-paniculate, equaling or
shorter than the leaves, puberulent or glabrate, the branches slender, usually
reddish, the pedicels 2-4 mm. long; flowers white or pinkish, 10 mm. broad or
smaller, fragrant, puberulent; perianth tube obsolete, the segments lanceolate to
elliptic, obtuse or rounded at the apex; filaments pilose, equaling or shorter than
the anthers, those of the third series with 2 large globose glands at the base; anthers
depressed-orbicular, truncate or emarginate at the apex; staminodia conspicuous,
glabrous, capitulate-thickened at the apex; ovary glabrous, shorter than the style;
fruit broadly ellipsoid, purple-black, about 12 mm. long and 10 mm. broad, the
cupule saucer-shaped.

Called "laurel" and "timber sweet" in British Honduras;
"piecito de paloma" (Tabasco). This is a very common tree of the
Yucatan Peninsula. It sometimes flowers and fruits when only a
large shrub.

Nectandra sinuata Mez, Jahrb. Bot. Gart. Berlin 5: 402. 1889.
Persea Matudai Lundell, Lloydia 4: 49. 1941 (type from Chiapas).
Tepeaguacate rojo; Aguacatillo; Canoj negro; Canoj bianco.

Mostly in damp or wet, often dense, mixed forest, frequently
on open, moist or rather dry, brushy hillsides, 200-2,300 meters;
Alta Verapaz; Zacapa; Santa Rosa; Guatemala; Sacatepe*quez;
Solola; Suchitepe'quez; Retalhuleu (type collected by Bernoulli &
Cario, no. 2581); Quezaltenango; San Marcos; Huehuetenango.

A small or large tree, sometimes 35 meters high with a trunk 1.2 meters in
diameter, the trunk tall and slender in the large trees, the crown narrow, the young
branches densely pilose or tomentose with fulvous or grayish hairs; leaves mem-
branaceous, on stout or slender petioles 1-3 cm. long, usually very broadly obovate,
mostly 20-30 cm. long and 10-15 cm. wide, rounded or very obtuse at the apex
and abruptly apiculate-acute or short-acuminate, obtuse or rounded at the base,
thinly pilose above, beneath usually densely velutinous-pilose or often hirsute;
inflorescence many-flowered, axillary, laxly corymbose-paniculate, densely
tomentose or pilose, long-pedunculate, often longer than the leaves, the pedicels
5-20 mm. long; flowers perfect, 17-19 mm. broad, densely pilose or villous, greenish

322 FIELDIANA: BOTANY, VOLUME 24

white or usually dull red or often pink outside; perianth tube obsolete, the seg-
ments broadly ovate, subacute; anthers sessile, papillose, those of the outer series
subfoliaceous, obtuse; staminodia none; ovary densely villous, much shorter than
the style; fruit broadly ellipsoid, almost 2 cm. long, the cupule 1.5 cm. broad.

Known in Salvador by the names "trompillo," "chipinahuaca,"
"trompito," "aguacate de mico," and "aguacate amarillo." The
bark and wood are said to yield a yellow dye. The Indian name
"canoj" is given to this and most other Lauraceae in Guatemala,
chiefly in the mountains of the Occidente. In the departments of
San Marcos and Huehuetenango there are caserios with the name
Canoj.

Nectandra surinamensis Mez, Jahrb. Bot. Gart. Berlin 5:
454. 1889.

Moist or wet, mixed forest, 300-600 meters; Alta Verapaz;
Escuintla. Perhaps also farther south in Central America; Guianas.

A tree, the branchlets yellowish-tomentulose or pilose; leaves on petioles 1.5
cm. long or shorter, chartaceous, oblong or subelliptic, mostly 9-17 cm. long and
3.5-5 cm. wide, acute or short-acuminate, acute or obtuse at the base, rather
densely appressed-pilose or strigillose beneath, glabrate on the upper surface, the
venation prominulous and laxly reticulate on both surfaces; inflorescence panicu-
late or pyramidal-paniculate, thinly tomentulose, many-flowered, sometimes
slightly exceeding the leaves, the pedicels 2-5 mm. long; flowers perfect, white,
strigose, 5-6 mm. broad; perianth tube obsolete or nearly so, the segments ovate-
lanceolate, pilose within, subobtuse; filaments of the outermost stamens very
short, the anthers suborbicular, truncate at the apex; filaments of the third series
of stamens bearing 2 sessile, rather large, basal glands; staminodia stipe-like,
capitulate at the apex; ovary glabrous, the style short.

The determination of the Guatemalan material is questionable,
but one of the collections was determined by Mez.

OCOTEA Aublet

Trees or shrubs, glabrous or pubescent, often blackening when dried; leaves
alternate, petiolate, membranaceous to rigid-coriaceous, penninerved; panicles
axillary or pseudoterminal, few-many-flowered, dichotomously branched; flowers
perfect or dioecious; perianth tube none or conspicuous, the segments equal,
usually deciduous; stamens of the 3 outer series fertile, those of the fourth series
reduced to staminodia or wanting; stamens of the third series with usually sessile
basal glands; anthers 4-celled, the cells in 2 vertical rows; cells of the 6 outer
anthers introrse, of the third series extrorse or lateral; ovary ovoid or ellipsoid,
glabrous or pilose, the style usually elongate; fruit globose or ellipsoid, the cupule
with a simple or double margin, hemispheric or saucer-shaped.

About 200 species in tropical America, with a few scattered
species in Africa and the Mascarene Islands. Other species are

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 323

known from southern Central America, especially in Costa Rica.
Most of the Guatemalan Lauraceae whose leaves become blackish
in drying belong to this genus.

Branches narrowly winged 0. subalata.

Branches not winged.
Flowers glabrous.

Perianth with a conspicuous tube, the segments short.

Flowers perfect; inflorescence few-many-flowered O. Bernoulliana.

Flowers dioecious; inflorescences mostly many-flowered O. cernua.

Perianth cleft nearly or quite to the base, the tube none or very short.
Perianth segments widely spreading; venation of the upper leaf surface

not elevated, very inconspicuous 0. Dendrodaphne.

Perianth segments erect or nearly so; venation of the upper leaf surface

elevated and conspicuously reticulate, lax O. verapazensis.

Flowers pubescent, sometimes sparsely so.

Leaves rounded to obtuse at the apex, sometimes acute but with a short,

very obtuse tip.

Petioles broadly winged almost or quite to the base, the leaves practically
sessile, the blades usually more or less pubescent beneath, often bar-
bate in the axils of the nerves 0. chiapensis.

Petioles not winged, conspicuous, the blades glabrous, not barbate beneath.

Leaf blades mostly 19-29 cm. long O. Standleyi.

Leaf blades mostly 9-15 cm. long O. veraguensis.

Leaves very acute to long-acuminate at the apex.

Leaf blades broadest above the middle, long-attenuate to the base.

0. eucuneata.

Leaf blades broadest at or below the middle, not long-attenuate to the base.

Leaves barbate beneath in all or most of the axils of the nerves, the blades

usually blackish when dried, mostly 1.5-3 cm. wide 0. effusa.

Leaves not barbate beneath, not blackish when dried, the blades usually
wider.

Branches of the inflorescence glabrous or essentially so, the inflores-
cence flexuous or curved, often longer than the leaves.

O. laetevirens.

Branches of the inflorescence usually pubescent, the inflorescence stiff,
straight, shorter than the leaves 0. Lundellii.

Ocotea Bernoulliana Mez, Jahrb. Bot. Gart. Berlin 5: 275.
1889. Canoj.

Dense, moist or wet, usually mixed forest, 300-1,650 meters;
Alta Verapaz; Escuintla; Retalhuleu (type from Mujulia, Bernoulli
& Cario 2590); San Marcos; Huehuetenango. Southern Mexico;
probably extending south to Panama.

A large shrub or usually a slender, sparsely branched tree about 6 meters high,
glabrous throughout; leaves chartaceous, usually lustrous, on slender petioles 1.5

324 FIELDIANA: BOTANY, VOLUME 24

cm. long or usually shorter, elliptic, mostly 10-15 cm. long and 3.5-7 cm. wide,
abruptly acuminate or usually caudate-acuminate, acute at the base, penninerved,
with usually 5-6 pairs of lateral nerves, prominulous-reticulate on both surfaces;
inflorescence laxly paniculate, longer or usually shorter than the leaves, the
branches spreading or somewhat reflexed, the slender pedicels 3-5 mm. long;
flowers perfect, glabrous, 3 mm. long, green; perianth tube conspicuous, not con-
stricted at the apex, the segments short, ovate, subacute; filaments sparsely pilose,
longer than the anthers; filaments of the third series of stamens bearing 2 small
sessile subglobose glands; anthers ovate, subacute; staminodia none; ovary gla-
brous, globose, the style short; fruit depressed-globose or ellipsoid, as much as
1.5 cm. long; cupule truncate, semiglobose, 1 cm. broad.

Called "laurel" and "timber sweet" in British Honduras;
"laurel de bajo" (Campeche); "aguacatillo" (Honduras); "laurel
amarillo" (Veracruz). In some parts of its range the tree reaches
a height of 12 meters and a trunk diameter of 30 cm.; the crown is
dense and spreading or narrow and irregular; bark light to dark
brown, the inner bark pinkish. The wood is white or yellowish,
turning brown on exposure to air, the heartwood sometimes dark
brown. Material of this species usually has been referred to 0. cernua
(Nees) Mez, but all or most of the continental collections so named
are referable rather to 0. Bernoulliana, which is a common species
in many regions of the Central American lowlands, chiefly in dense
wet rain forest.

Ocotea cernua (Nees) Mez, Jahrb. Bot. Gart. Berlin 5: 377.
1889. Oreodaphne cernua Nees, Syst. Laur. 424. 1836.

Wet mixed forest, at or little above sea level; Izabal (probably
in this department; S. Watson 450). Southern Mexico; British
Honduras; Honduras; Costa Rica; Panama; reported from South
America.

A large shrub or small tree, the branchlets pubescent at first but soon glabrous;
leaves on slender petioles 12 mm. long or less, chartaceous or subcoriaceous, gla-
brous, usually oblong-elliptic, 16 cm. long and 6.5 cm. wide or smaller, gradually
or abruptly acuminate, rounded or obtuse at the base, the lateral nerves 4-6
pairs; panicles numerous, glabrous, many-flowered, branched, axillary; flowers
glabrous, yellowish, not more than 2 mm. long, dioecious, the pedicels filiform,
4 mm. long or shorter; staminate and pistillate flowers much alike except in their
stamens and pistils; fruit black, ellipsoid, apiculate, 14 mm. long and 9 mm.
broad or smaller, the woody cupule about 11 mm. broad and 7 mm. long.

Called "aguacatillo" in British Honduras; "laurel" (Tabasco).
In general appearance this is exactly like 0. Bernoulliana, and Miss
Allen suggests that the latter may be only "a different manifestation"
of 0. cernua.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 325

Ocotea chiapensis (Lundell) Standl. & Steyerm. Field Mus.
Bot. 23: 114. 1944. Nectandra chiapensis Lundell, Contr. Univ.
Mich. Herb. 6: 12. 1941. Canoj.

Moist or wet, mixed forest, 1,400-2,800 meters; Huehuetenango;
San Marcos. Chiapas, the type from Rodeo, Siltepec, at 2,800
meters.

A tree 12 meters high or perhaps sometimes larger, the branchlets thick,
often conspicuously angulate, minutely and densely sericeous when young with
brownish or grayish hairs; petioles thick and broad, 2 cm. long or usually shorter,
winged almost or quite to the base, the leaves thus essentially sessile; leaf blades
chartaceous or coriaceous, oblong-elliptic to oblanceolate-oblong or obovate-
oblong, 10-17 cm. long, 4-6.5 cm. wide, subacute to rounded at the apex, gradually
narrowed to the acute base, the lower part of the margin strongly recurved and
forming a narrow pocket, glabrous above or nearly so, very closely brownish-
sericeous beneath at first, in age glabrate, usually densely barbate in the axils of
the nerves, the lateral nerves 9-12 pairs, the veins slightly prominulous-reticulate,
especially beneath; panicles axillary, thinly brownish-sericeous, many-flowered,
usually broad and long-pedunculate, the pedicels 1-2.5 mm. long; flowers 6-7 mm.
broad, densely brownish-sericeous; perianth tube very short, the segments ovate
or oblong-ovate, subacute or very obtuse, spreading or ascending-spreading;
filaments of the outer stamens shorter than the anthers, sparsely pilose; anthers
truncate at the apex; ovary glabrous, about as long as the style; fruit oblong-
ellipsoid, 3-3.5 cm. long, 1.5 cm. broad; cupule rose-red, 1.5 cm. broad, the sup-
porting pedicel much thickened.

This species is very close to 0. nicaraguensis Mez, and perhaps
identical with it. That was based on fruiting material from San
Juan, Nicaragua.

Ocotea Dendrodaphne Mez, Jahrb. Bot. Gart. Berlin 5: 238.
1889. 0. ovandensis Lundell, Contr. Univ. Mich. Herb. 6: 16. 1941.
Aguacate de mico.

Moist or wet, mixed forest, 900-1,500 meters or higher; El
Progreso(?); Quezaltenango; Huehuetenango (?). Chiapas; Atlantic
coast of Honduras; Costa Rica.

A tree 12-30 meters high, the trunk as much as 50 cm. in diameter, the bark
gray, slightly roughened, the branchlets stout, minutely strigillose or puberulent
at first, soon glabrate, the older ones gray, subterete, striate; leaves coriaceous or
subcoriaceous, on naked petioles 2 cm. long or shorter, oblong or elliptic-oblong,
glabrous, mostly 16-25 cm. long and 6-10 cm. wide, abruptly short-acuminate,
cuneate at the base, grayish or fuscous when dried, slightly lustrous on the upper
surface, the venation not or scarcely elevated, finely and closely prominulous-
reticulate beneath, penninerved, the lateral nerves 9-11 pairs; inflorescences
broadly paniculate, many-flowered, shorter than the leaves, on long or short
peduncles, very minutely puberulent or almost glabrous, the pedicels 5 mm. long
or less; flowers white, fragrant, perfect, very minutely puberulent or strigillose or

326 FIELDIANA: BOTANY, VOLUME 24

almost glabrous; perianth tube very short, the segments oblong or elliptic-oblong,
3-4 mm. long, obtuse, pubescent within at the base; filaments very short, pilose;
anthers oblong, obtuse; staminodia minute, stipe-like, pilose; ovary glabrous,
equaling or longer than the style; fruit ellipsoid, 18 mm. long, 11 mm. broad, the
cupule 7-13 mm. broad.

Ocotea effusa (Meissn.) Hemsl. Biol. Centr. Amer. Bot. 3: 73.
1882. Oreodaphne effusa Meissn. in DC. Prodr. 15, pt. 1: 120. 1864.
Canoj bianco (Quezaltenango) .

Moist or wet, mixed forest, sometimes in thickets, 2,500 meters
or less, mostly at 1,200 meters or lower; Pete"n; Alta Verapaz;
Izabal(?); Escuintla; Sacatepe"quez; Solola; Quezaltenango; San
Marcos; Huehuetenango. Southern Mexico.

A large shrub or a small tree, mostly 6-12 meters high, the branches very
slender, sparsely pubescent or glabrate; leaves on slender petioles 1 cm. long or
less, lanceolate or oblong-lanceolate, mostly 7-11 cm. long and 1.5-2.5 cm. wide,
narrowly long-acuminate, acute or subobtuse at the base, membranaceous or sub-
chartaceous, deep green above, lustrous, glabrous or nearly so, paler beneath,
usually barbellate in the axils of the nerves, somewhat pubescent on the nerves,
elsewhere glabrous or nearly so, penninerved, the venation not elevated on the
upper surface, beneath barely prominulous, laxly reticulate; inflorescence few-
many-flowered, laxly paniculate, almost or quite glabrous, often much longer than
the leaves, the slender pedicels 3-10 mm. long; flowers perfect, sparsely pubescent,
2 mm. long; perianth tube conspicuous, the segments ovate, acute; filaments
glabrous, very short, those of the third series of stamens with 2 rather large, sessile,
subglobose glands at the base; anthers rounded at the apex; staminodia stipe-like,
densely pilose; ovary glabrous, attenuate into a short style; fruit purple-black,
1-2 cm. long, 7 mm. broad, the cupule rose-red, the pedicel much thickened.

Ocotea eucuneata Lundell, Contr. Univ. Mich. Herb. 6: 16.
1941.

Dense wet mixed forest, 1,500 meters or less; Alta Verapaz;
Izabal. British Honduras, the type from Middlesex, Stann Creek
District, P. H. Gentle 3068.

A medium-sized or large tree, 9 meters high or more, the trunk said to be as
much as a meter in diameter, the branchlets appressed-pubescent, stout or rather
slender, terete or slightly angulate; leaves on petioles 5-12 mm. long, chartaceous
or thick-membranaceous, oblanceolate to obovate-oblong, mostly 8-23 cm. long
and 3-7.5 cm. wide, usually drying blackish, abruptly acuminate, attenuate to the
narrow base, glabrous above or nearly so, the venation not elevated, somewhat
paler beneath, puberulent or glabrate, usually barbate in the axils of the nerves,
laxly prominulous-reticulate, the lateral nerves 5-8 pairs; inflorescence slender-
pedunculate, paniculate, 9 cm. long or shorter, grayish-puberulent ; flowers perfect,
short-pedicellate, densely grayish-puberulent; perianth tube conspicuous, the
segments ovate, 2 mm. long; filaments pilosulous, about as long as the anthers,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 327

these truncate or subemarginate at the apex; ovary glabrous, shorter than the
style.

Ocotea laetevirens Standl. & Steyerm. Field Mus. Bot. 23:
114. 1944.

In forest, 800-2,000 meters; endemic; Huehuetenango (type
from Cerro Chiblac, between Finca San Rafael and Ixcan, Steyer-
mark 49189).

A tree of 9 meters, the branches very slender, almost glabrous or when young
sparsely and minutely puberulent, terete; leaves thick-membranaceous, yellowish
green or olivaceous when dry, on slender petioles 7-10 mm. long, oblong-elliptic
or lance-oblong, 12-18 cm. long, 4-6.5 cm. wide, gradually or abruptly acuminate,
with a narrow obtuse tip, acute at the base, glabrous, dull on the upper surface,
the venation scarcely prominulous, somewhat paler beneath, penninerved, the
lateral nerves about 8 pairs, the veins laxly prominulous-reticulate; inflorescence
lax, many-flowered, paniculate, often longer than the leaves, about 6 cm. long, the
slender, flexuous, perhaps recurved peduncle 4-9 cm. long, the pedicels subumbel-
late, 2-3 mm. long, glabrous or sparsely and minutely strigillose; flowers globose,
scarcely 2 mm. long, minutely and sparsely puberulent; perianth tube very short,
the segments equal, broadly elliptic, very obtuse, suberect; filaments of the outer-
most stamens broad and thick, slightly longer than the anthers, glabrous, the
anthers broadly ovate, very obtuse at the apex; basal glands of the third series of
stamens large, globose, sessile; staminodia stipe-like, very slender and short, or
none; cupule of the fruit hemispheric, 1 cm. broad, the margin simple, rose-red.

Ocotea Lundellii Standl. Carnegie Inst. Wash. Publ. 461: 56.
1935. Yaaxhochoc (Pete'n, Maya, fide Lundell) ; Laurel.

Moist or wet, mixed forest, often or perhaps usually on lime-
stone, 1,500 meters or less, mostly at 300 meters or lower; Pete'n
(type from ruins of Ixlu, Lago de Pete'n, C. L. Lundell 4359) ; Alta
Verapaz; Izabal; Huehuetenango (?). Campeche.

A tree, sometimes 15 meters high, often much lower, with a trunk 30 cm. or
probably more in diameter, the branches glabrous; leaves lustrous, coriaceous, on
slender petioles 7-14 mm. long, lance-oblong to ovate or ovate-elliptic, mostly
9-12 cm. long and 3.5-4.5 cm. wide, rather abruptly short-acuminate or long-
acuminate, with an obtuse tip, acute at the base, glabrous, penninerved, the veins
prominulous and laxly reticulate on both surfaces, paler beneath, the lateral nerves
about 6 pairs; inflorescences axillary, cymose-paniculate, 3-6 cm. long, lax, few-
many-flowered, shorter than the leaves, stiff, the branches glabrous or puberulent,
often reddish, the pedicels 7 mm. long or less, glabrous or puberulent; flowers
4.5-5 mm. long, white or greenish white, sparsely and minutely sericeous outside
or glabrate, the perianth segments papillose-villosulous within, ascending or some-
what spreading, the tube very short; fruit purple-black, oval or ellipsoid, as much
as 2.5 cm. long and 1.5 cm. broad or usually smaller, the cupule short, about 6 mm.
broad, the pedicel much thickened.

328 FIELDIANA: BOTANY, VOLUME 24

Ocotea Standleyi Allen, Journ. Arnold Arb. 26: 343. 1945.
Phoebe macrophylla Standl. & Steyerm. Field Mus. Bot. 23: 116.
1944, not Mez (type collected southeast of Tactic, Alta Verapaz,
Standley 70009).

Dense, moist or wet, mixed, mountain forest, 1,200-1,700 meters;
endemic; Alta Verapaz; Quezaltenango; San Marcos.

A large shrub or a tree of 6 meters or more, glabrous outside the inflorescence,
the branches slender, terete or obtusely angulate, the older ones grayish brown;
leaves on stout naked petioles 1.5-2 cm. long, chartaceous, oval to obovate-elliptic,
19-29 cm. long, 9-12 cm. wide, rounded or very obtuse at the apex, sometimes
acute, rounded or very obtuse at the base or subcordate, penninerved, the lateral
nerves about 13 pairs, the veins prominulous and laxly reticulate on both surfaces,
often lustrous, when dry usually brownish; panicles axillary, long-pedunculate,
lax, many-flowered, sometimes equaling the leaves, the lower branches glabrous,
the upper ones sparsely puberulent or pilosulous, the pedicels 1.5-3 mm. long;
flowers greenish, 2.5-3 mm. long, sparsely strigillose; perianth tube well developed,
broadly turbinate, the segments ascending or almost erect, suborbicular, rounded
at the apex; filaments almost equaling the anthers, thick, sparsely pilosulous or
almost glabrous, the anthers oblong-quadrate, obtuse; staminodia small, short-
stipitate, oblong or ovate; ovary glabrous, ovoid, about equaling the thick style;
fruit oval or ellipsoid, 2 cm. long, 1.5 cm. broad, rounded at the apex, the cupule
turbinate-campanulate, 1 cm. broad, the margin simple.

Ocotea subalata Lundell, Lloydia 4: 48. 1941.

Known only from the type, north side of Volcan de Tacana,
Chiapas, 2,100 meters, E. Matuda 2957; doubtless occurring in San
Marcos; material from Volcan de Tajumulco, at 1,300-2,000 meters,
probably belongs here.

A tree, the branchlets angulate and narrowly winged, at first hirsute-tomentose
with brown hairs, usually drying blackish; leaves on petioles 13-20 mm. long,
chartaceous, oblong-lanceolate or oblong, 8-21 cm. long, 4-8 cm. wide, abruptly
acuminate with an obtuse tip, acute at the base, glabrous on the upper surface,
the costa and nerves subimpressed, persistently pubescent beneath with sub-
appressed brownish hairs, reticulate- veined, the lateral nerves 6-9 pairs; inflores-
cences axillary, corymbose-paniculate, long-pedunculate, in anthesis about
equaling the leaves, in fruit as much as 40 cm. long, subappressed-pilosulous, the
branches subangulate, sometimes slightly winged, the pedicels 4 mm. long or less,
much elongate in fruit; flowers perfect, appressed-pilosulous, 2.5 mm. long; peri-
anth tube short, the segments ovate, obtuse; filaments half as long as the anthers,
sparsely pubescent; glands at the base of the third series of stamens sessile, con-
spicuous; anthers ovate, obtuse; ovary glabrous, longer than the style; fruit
ellipsoid, black, lustrous, as much as 2.5 cm. long and 17 mm. broad, the cupule
very shallow, 11 mm. broad.

Ocotea veraguensis (Meissn.) Mez, Jahrb. Bot. Gart. Berlin 5:
240. 1889. Sassafridium veraguense Meissn. in DC. Prodr. 15, pt. 1:

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 329

171. 1864. Pimiento; Pimienton; Pububuc (reported as the Quecchi
name).

Moist or dry forest or thickets, most common along stream banks,
often on dry rocky hillsides, 1,400 meters or less, chiefly at 700
meters or lower and probably most common on the Pacific plains;
El Progreso; Zacapa; Santa Rosa; Escuintla; Guatemala; Chimalte-
nango; Suchitepe"quez ; Retalhuleu; Quezaltenango ; Huehuetenango.
Chiapas; Salvador to Panama.

A large shrub or usually a tree of 6-12 meters, often larger, the trunk generally
dense, broad, and spreading, the bark almost smooth, grayish, the young branch-
lets ferruginous-tomentulose but in age glabrous, subangulate or terete; leaves
coriaceous, lustrous, on slender naked petioles mostly 1 cm. long or shorter,
narrowly elliptic to oblong-lanceolate, mostly 9-15 cm. long and 3.5-4.5 cm. wide,
chiefly obtuse but frequently acute, glabrous, acute at the base, penninerved, the
venation not elevated on the upper surface, prominulous-reticulate beneath;
panicles many-flowered, pyramidal or subcorymbose, sparsely pilosulous, equaling
or shorter than the leaves, the pedicels 4-10 mm. long; flowers creamy white,
perfect, pilosulous, 3 mm. long; perianth tube short, the segments broadly elliptic,
acute, somewhat spreading; anthers sessile, densely papillose, the connective long-
produced beyond the cells; staminodia abortive; ovary glabrous, the style short;
fruit black, ellipsoid, 17-20 mm. long, 10 mm. broad, the cupule shallow, with a
double margin.

Sometimes called "canelo" in Salvador; "aguacatillo" (Hon-
duras). One of the commonest trees along stream banks on the
Pacific plains, often growing at the very edge of the water, the
branches extending far out over the stream.

Ocotea verapazensis Standl. & Steyerm. Field Mus. Bot. 23:
114. 1944.

Dense wet mixed mountain forest, 1,650 meters or less; endemic;
Alta Verapaz (type from Tactic, Standley 71421); Izabal; San
Marcos.

A tree of 6-12 meters, the branchlets slender, obtusely angulate, glabrous or
nearly so; leaves chartaceous, blackish when dried, on narrowly marginate petioles
1 cm. long or shorter, oblanceolate-oblong, mostly 14-27 cm. long and 4.5-8 cm.
wide, gradually or abruptly and shortly obtuse-acuminate, gradually narrowed
to the acute or subobtuse base, glabrous, penninerved, the veins closely prominu-
lous-reticulate on both surfaces, the lateral nerves about 8 pairs, not barbate in
the axils; panicles lax, many-flowered, on long slender peduncles, 15 cm. long or
shorter, shorter than the leaves, the branches glabrous, the slender pedicels 3.5
mm. long or less; flowers green, glabrous, 2.5 mm. long; perianth tube very
short, the segments broadly elliptic, obtuse; anthers almost sessile, ovate-quadrate,
very obtuse or subtruncate at the apex, the filaments thick, glabrous; staminodia
abortive; ovary globose, glabrous, the style very short; fruit ellipsoid, lustrous,

330 FIELDIANA: BOTANY, VOLUME 24

2.5 cm. long, 1.5 cm. broad, the cupule red, turbinate-campanulate, 12-15 mm.
broad, the rather long pedicel greatly thickened.

PERSEA Miller

Trees or shrubs; leaves alternate, petiolate, chartaceous or coriaceous, usually
somewhat pubescent; panicles axillary or subterminal; flowers large, cymose or
subumbellate, not involucrate, perfect; perianth tube very short or none, the 6
segments equal or the outer ones usually smaller, mostly persistent; stamens
9, generally all fertile, those of the fourth row reduced to staminodia; filaments
filiform, commonly longer than the anthers, pilose or glabrous; third row of stamens
with stipitate, usually large glands, the stipes of the glands united with the fila-
ments; anthers commonly 4-celled, ovate, the cells large; outer anthers introrse,
the inner 6 anthers extrorse or extrorse-lateral ; staminodia large, distinctly
stipitate, cordate or sagittate, often pubescent at the apex; ovary subglobose,
glabrous or pilose, the style usually longer, glabrous or pilose, the stigma large,
dilated; fruit globose or ellipsoid, small or often very large; perianth not enlarged
in fruit.

Species about 60, in tropical or subtropical America. A very
few additional species are found in Central America.

Leaves sessile or nearly so P. sessilis.

Leaves long-petiolate.

Ovary pubescent; fruit usually large, commonly 3.5-10 cm. long or even larger.

Pedicels 8-15 mm. long; head of the staminodia elliptic, about as broad as
the stipe; branchlets densely ferruginous-tomentose; leaves usually very
densely and softly pubescent beneath P. Schiedeana.

Pedicels 1-10 mm. long; head of the staminodia triangular, much broader
than the stipe; leaves almost or quite glabrous beneath except along the
nerves, at least in age.

Leaf blades mostly 6-10 cm. long P. Steyermarkii.

Leaf blades mostly 12-20 cm. long.

Leaves not anise-scented; perianth deciduous P. americana.

Leaves with the odor of anise or sassafras; perianth usually persistent.

P. americana var. drymifolia.

Ovary glabrous; fruit small, often only 1 cm. long but sometimes larger.
Leaves densely tomentose beneath with lax, more or less spreading hairs.

P. Donnell-Smithii.

Leaves glabrous beneath or minutely sericeous, or with a minute and very
closely appressed tomentum.

Leaves glabrous beneath P. Standleyi.

Leaves covered beneath with a very dense but minute and closely appressed
tomentum, appearing glabrous to the naked eye P. vesticula.

Persea americana Mill. Gard. Diet. ed. 8. 1768. Laurus Persea
L. Sp. PI. 370. 1753. P. gratissima Gaertn. Fruct. 3: 222. pi. 221.
1807. Aguacate; 0, Oj, Ju, Un, Um, On (various Indian dialects of

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 331

Guatemala); Tsumon (soft-skinned fruit), Tc'om (hard-shelled
fruit), both names used at Jacaltenango.

Cultivated at all elevations in Guatemala, in its various forms
and varieties; in many localities more or less naturalized and in
some regions perhaps native, or possibly only a relic of former
cultivation; such apparently wild trees have been collected or noted
in the mountains of Zacapa, Chiquimula, Huehuetenango, Que-
zaltenango, and elsewhere. Native in tropical America, doubtless
in many regions of Mexico and Central America, and common in
cultivation in many other parts of tropical America, also in the Old
World tropics.

A large or medium-sized tree, often 20 meters high, with a very dense, rounded
or elongate crown, the young branches glabrous to puberulent or pilosulous, often
glaucous; leaves on slender petioles 2-6 cm. long, oval to elliptic or obovate-oval
or sometimes ovate, mostly 10-30 cm. long, acute or acuminate, unequal at the
base and acute to rounded, chartaceous, penninerved, deep green above, glabrous
or nearly so, often lustrous, pale and glaucescent beneath, glabrous or pilosulous
with short spreading hairs, especially along the nerves; panicles densely grayish-
puberulent or sericeous, few or many near the ends of the branches, 6-20 cm. long,
pedunculate, the slender pedicels 3-6 mm. long; perianth pale greenish, 5-7 mm.
long, densely grayish-tomentulose, the segments elliptic or lance-elliptic to oval-
ovate, obtuse, the outer ones shorter; filaments pilose; staminodia 2-2.8 mm. long,
the head triangular, acute, truncate or sagittate-cordate at the base, slightly
shorter than the stipe; fruit highly variable in size, shape, color, and quality of the
flesh.

Called "pear" and "butter-pear" in British Honduras. The
usual names in the United States are "avocado" and "alligator
pear." The avocado is one of the most abundant and popular
fruits of Guatemala, and this country produces some of the finest,
if not the finest, avocados of America. The trees are planted in
every inhabited region, from sea level to the summits of the moun-
tain ranges, or at least to 3,000 meters and more. The varieties are
innumerable, based upon shape, size, and color of the fruit and on the
thickness of its skin. Some of the best of these varieties have been
introduced into other parts of the earth, but chiefly into the United
States, in Florida and southern California, where the trees produce
well and have become in recent years the basis of a substantial
industry. The fruit grown to maturity in Florida and California is
of good quality, but as it reaches the markets of the northern and
eastern United States it usually is very inferior, principally because
of faulty harvesting and shipping. The fruits still are somewhat of a
luxury in northern markets, being retailed at about twenty-five
cents each, a sum that in Guatemala would buy a large number of

332 FIELDIANA: BOTANY, VOLUME 24

much better avocados. The person chiefly responsible for intro-
duction of this fruit tree into the United States is Dr. Wilson
Popenoe, formerly of the United States Department of Agriculture
and later the United Fruit Company, who, with his wife, is insepa-
rably associated with "The House" of Antigua. Many years ago
he explored on muleback the remotest mountains of Guatemala
in search of the best varieties for introduction elsewhere, and he
proposed the horticultural classification of the fruit most used for
practical purposes, which is as follows: (1) Mexican type, the leaves
anise-scented, the skin of the fruit thin and soft, Per sea americana
var. drymifolia (see below); (2) West Indian type, the leaves not
anise-scented, the surface of the fruit usually smooth, the skin
leathery but thin; (3) Guatemalan type, the surface of the fruit
usually rough or warty, the skin brittle, granular, relatively thick
and hard. Of these three races only two are common in Guatemala.
The Guatemalan type is grown at 900 meters and upward to the
limit of cultivation; at 750 meters and lower is planted the West
Indian type, which ripens chiefly in July and September. The
Mexican race is almost unknown in Guatemala, but there are a few
trees in Sacatepe"quez, Chimaltenango, and elsewhere. Avocados,
because of the wide range of elevation at which they are planted,
may be obtained in Guatemala at all seasons of the year, and they
are produced in vast quantities. It may be said that all of them are
good, although some are better than others, and the hard-skinned
fruits usually are preferred to the West Indian type. Most people
are very fond of avocados, which are eaten rather as a salad vegetable
than as a fruit, although people often pluck them from the tree and
eat them like an apple. The fruit is rich in oil and highly nutritious,
and with bread affords a good meal. In Guatemala it is much served
on the table as a salad or appetizer, and it often appears in the form
of guacamol, the pulp separated from the skin, mashed, and flavored
with oil, vinegar, onion, garlic, chile, and other substances. The
fruit is eaten by all domestic animals; even dogs are fond of it, and
many or most wild animals relish it.

Besides the three chief horticultural forms of this fruit, there are
many minor varieties distinguishable by size, shape, and color. The
fruit is mostly obovoid and green, but it is often tinged with red and
yellow, and the shape varies greatly in some of the rare varieties.
One with sausage-shaped fruit is of rare occurrence. There is said
to be sold occasionally in the Quezaltenango market a delicious
small avocado, about 2.5 cm. in diameter, with a large seed and scant
thin flesh, perhaps from wild trees.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 333

The name "aguacate" given commonly in all parts of Central
America to this fruit is of Nahuatl origin, derived from the term
ahuacatl or ahuacuahuitl. The former word is also the Aztec term
for testicle, but this is probably a derived application. There is,
however, a belief popular in Mexico and extending also into Central
America that the avocado has aphrodisiac properties. The name
Aguacate is much used in local geographic names, being applied
to settlements in at least fourteen of the departments of Guatemala.
Most important is the well-known pueblo, Aguacatan, in Huehue-
tenango.

The sap of the avocado seed makes an indelible stain on cloth
and is sometimes used for marking clothing. The pulverized seeds
mixed with cheese, tallow, or other substances are used, strangely
enough, for poisoning mice and other destructive animals. The
Indian women often boil the bark with dyes for textiles, to set them.
The rind of the fruit is employed as a vermifuge. The fruit contains
about 14 per cent of fat or oil, and in recent years it is being extracted
on a rather large scale in some parts of tropical America. It is a
common commercial article in Guatemala, being extracted locally
and used principally as a substitute for olive oil on the table and
elsewhere. It often is applied to the hair to improve its appearance.

Persea americana var. drymifolia (Schlecht. & Cham.) Blake,
Journ. Wash. Acad. Sci. 10: 15. 1920. Persea drymifolia Schlecht. &
Cham. Linnaea 6: 365. 1831. Aguacate de anis.

Cultivated in Guatemala, but infrequently, as mentioned above.
Differing in its leaves, which have the odor of anise or sassafras;
fruit thin-skinned.

This is the common avocado of Mexico, but it is of rare occur-
rence in Guatemala. So far as we know, the two forms can not be
distinguished by herbarium specimens.

Persea Donnell-Smithii Mez ex Bonn. Smith, Enum. PI.
Guat. 2: 67. 1891; Mez, Arb. Bot. Gart. Breslau 1: 113. 1892.
Aguacate; Sacsi (Coban, Quecchi).

Chiefly in open pine forest, sometimes in dense wet mixed forest,
often in pastures or sometimes in open swamps, 1,200-2,000 meters;
Alta Verapaz (type from Chicoyonito, J. D. Smith 1718); Baja
Verapaz; Chiquimula. Southern Mexico.

A tree of 5-12 meters or often larger, with a rather thick trunk and a low dense
rounded crown, the young branches very densely tomentose with lax spreading

334 FIELDIANA: BOTANY, VOLUME 24

brownish hairs; leaves coriaceous or thick-coriaceous, on thick, densely brownish-
tomentose petioles 2-4 cm. long, oblong-oval to almost orbicular or oval-ovate,
mostly 10-20 cm. long, rounded or obtuse at the apex, rounded to subacute and
often conspicuously unequal at the base, glabrate in age on the upper surface,
beneath brownish and very densely and laxly tomentose, the nerves elevated and
very conspicuous beneath; inflorescences usually numerous in the upper leaf axils,
long-pedunculate, shorter or usually longer than the petioles, few-many-flowered,
very dense, sparsely branched, the floriferous portion usually shorter than the
stout peduncle, densely ferruginous-tomentose, the flowers sessile or nearly so;
sepals very unequal, the outer ones short, the inner ones broadly ovate to sub-
orbicular, densely tomentose, rounded or very obtuse at the apex, persistent in
fruit; ovary glabrous; young fruit globose, probably about 1 cm. in diameter at
maturity.

This is a very common tree in the Coban region, abundant in
many of the pastures, where it often is left for no apparent reason.
The fruits, so far as we know, are not edible.

Persea Schiedeana Nees, Syst. Laur. 130. 1836. P. gratissima
var. Schiedeana Meissn. in DC. Prodr. 15, pt. 1: 53. 1864. P. Pit-
tieri Mez, Bot. Jahrb. 30: Beibl. 67: 15. 1901. Coy 6, Coyocte, Kivo,
Kiyau, Cotoyo (Alta Verapaz); Chucte, Chaucte (El Progreso);
Xucte (Zacapa); Aguacate de monte (Huehuetenango) ; Chalte
(Zacapa).

Moist or wet, mixed forest, often in open, pine or oak forest,
frequently in open fields or pastures, 900-2,700 meters; Alta Vera-
paz; El Progreso; Izabal; Zacapa; Chiquimula; Huehuetenango; San
Marcos. Southern Mexico; Honduras; Costa Rica; Panama.

Usually a tree of 15-20 meters but sometimes as much as 50 meters high,
with a large crown, the branchlets stout, densely tomentose with mostly ferrugi-
nous, sometimes grayish pubescence; leaves on slender petioles 1.5-4.5 cm. long,
thick-membranaceous or chartaceous, obovate to elliptic-obovate or oval, 12-30
cm. long, 7-15 cm. wide, broadly rounded and apiculate to subacute at the apex,
broadly rounded or obtuse at the base, penninerved, green above, glabrous or
nearly so in age, when young often tomentose, paler or glaucous beneath, densely
pilose with short spreading velutinous hairs; panicles long or short, densely grayish-
tomentulose, mostly 10-12 cm. long, long-pedunculate, the slender pedicels 8-15
mm. long; perianth greenish yellow, 6-8 mm. long, densely grayish-tomentulose,
the segments subequal, lance-elliptic, subacute; filaments pilose; staminodia
pilose, the stipe subulate, 2-3 times as long and about as thick as the elliptic
obtuse head; ovary densely pilose; fruit similar to that of P. americana, variable
in size and shape, the skin thick blit leathery and pliable, the flesh brownish white,
of fine oily texture, permeated by numerous coarse tough fibers; cotyledons rose-
pink (whitish in P. americana).

Called "yas" hi Costa Rica, "chuti" in Honduras, and "chinini"
in southern Mexico. The tree is common in the mountain forests of
various parts of Guatemala, but especially in the mountains of Alta

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 335

Verapaz. The trees lose their leaves there during the dry season.
They usually are left when the forest is cleared and often are plentiful
in pastures. The fruit varies greatly in quality, that of most wild
trees being unpleasantly fibrous and having scant flesh. However,
the flavor is so good that the fruit is much appreciated, and it is
sold commonly in the markets during its relatively brief season.
Some trees have large fruits in which the fiber is not conspicuous.
Occasionally the trees are planted in fincas but most of the fruit is
harvested from wild trees.

Persea sessilis Standl. & Steyerm. Field Mus. Bot. 23: 115. 1944.

Moist mixed mountain forest, 2,100-2,400 meters; known only
from the type, Zacapa, Sierra de las Minas, along Rio Repollal to
summit of mountain, Steyermark 42487.

A shrub of 1.5 meters, the branchlets stout, terete, densely leafy, fuscous-
ferruginous, glabrous or glabrate; leaves on petioles 4 mm. long, rather rigidly
coriaceous, lustrous, narrowly lance-oblong, about 20 cm. long and 5.5-7 cm. wide,
acute or acuminate with an obtuse tip, slightly attenuate to the base, the base
shallowly cordate, glabrous, the lateral nerves about 15 on each side; panicles
much shorter than the leaves, cymose, few-flowered, minutely and not densely
pilosulous-tomentulose, the flowers short-pedicellate, the branches ascending;
perianth segments subequal or the outer ones somewhat shorter, very broadly
ovate or almost rounded, very obtuse or rounded at the apex, sericeous on both
surfaces, in fruit persistent and spreading; immature fruit globose, 1 cm. in
diameter.

Persea Standleyi Allen, Journ. Arnold Arb. 26: 301. 1945.

Moist mixed mountain forest, 1,500-2,100 meters; Chiquimula
(Volcan de Quezaltepeque) ; Solola (type collected along trail,
slopes of Volcan de Santa Clara toward San Pedro, Steyermark
47130). Veracruz.

A tree of 7-12 meters, the branchlets glabrate, densely leafy; leaves alternate
or subverticillate, the petioles 3.5 cm. long or shorter, slightly pubescent, reddish;
leaf blades glabrous, coriaceous, greenish brown when dried, lanceolate or oblance-
olate, as much as 20 cm. long and 4.5 cm. wide, obtuse or acute, obtuse at the base,
with 10-12 nerves on each side; inflorescences axillary, subcapitate, shorter than
the leaves, as much as 5 cm. long, fulvous-sericeous, few-flowered; flowers short-
pedicellate, the perianth fulvous-tomentose, the lobes distinctly 5-nerved, ovate,
pubescent on both surfaces; gynoecium glabrous; fruit (immature?) globose,
apiculate, 9 mm. in diameter, subtended by the persistent perianth lobes.

Persea Steyermarkii Allen, Journ. Arnold Arb. 26: 286. 1945.

Known only from the type, San Marcos, trail between Finca El
Porvenir and San Sebastian, upper slopes of Volcan de Tajumulco,
1,300-1,400 meters, Steyermark 37061.

336 FIELDIANA: BOTANY, VOLUME 24

A small tree about 10 meters high, the branchlets densely leafy at the apex,
glabrous, dark reddish, becoming gray and rugulose; petioles 2 cm. long or shorter,
dark reddish, glabrous; leaf blades glabrous, coriaceous, pale beneath, lance-ellip-
tic or oblong-elliptic, 6-10.5 cm. long, 2.5-4 cm. wide, rounded or acute at the apex,
obtuse at the base, penninerved, with 6-7 pairs of nerves; inflorescences axillary,
shorter than the leaves, subverticillate, paniculate, 3-5 cm. long, few-flowered, the
peduncle glabrate, 3 cm. long or less; flowers 5.5-8 mm. long, the pedicels 7-10
mm. long, appressed-pubescent; perianth campanulate, yellow-green, the lobes
reflexed, the outer ones 4.5 mm. long, the inner 6 mm. long; gynoecium pubescent.

Persea vesticula Standl. & Steyerm. Field Mus. Bot. 23: 116.
1944.

Moist or wet, mixed, mountain forest, 1,500-3,000 meters; El
Progreso (Sierra de las Minas, hills north of Finca Piamonte) ; San
Marcos (type from Volcan de Tacana, between La Vega ridge and
northeast slopes of the volcano, near the Mexican boundary,
Steyermark 36207); Huehuetenango (Cerro Huitz, Sierra de los
Cuchumatanes). Doubtless extending into Chiapas.

A tree of 15-30 meters, the branchlets very thick, rugose, fuscous-ferruginous
or cinnamoa-brown, covered with a minute appressed tomentum; leaves rigid-
coriaceous, on petioles 1-2 cm. long, oblong or elliptic-oblong, 10-17 cm. long,
3.5-6.5 cm. wide, obtuse or subacute, obtuse or rounded at the base and some-
times more or less unequal, lustrous and glabrous above, brownish beneath,
covered everywhere with a very dense, minute, closely appressed, ochraceous or
brownish tomentum, penninerved, with about 9 pairs of nerves; inflorescences
numerous, borne in the upper leaf axils or densely clustered at the ends of the
branchlets, about 14 cm. long, densely tomentulose, the very thick pedicels
scarcely more than 2 mm. long; perianth 5-6 mm. long, densely tomentulose, the
segments broadly ovate or elliptic, obtuse, the outer ones slightly shorter; fila-
ments pilosulous, about equaling the anthers, these 4-celled, broadly oblong,
obtuse at the apex; ovary glabrous; fruit globose, rounded at the apex, about 3.5
cm. long.

PHOEBE Nees

Large or small trees, rarely shrubs; leaves chartaceous or coriaceous, alternate,
often 3-nerved; panicles axillary, mostly rather few-flowered and small but often
large and lax, the flowers generally cymose, perfect, not involu crate; perianth
tube very short or none, the segments 6, equal or nearly so, usually persistent;
fertile stamens 9, free; filaments equaling or shorter than the anthers, pilose or
glabrous, those of the 2 outer series eglandular, those of the third series with 2
sessile basal glands; anthers usually 4-celled, in the outer series introrse, in the
third series extrorse or lateral; staminodia conspicuous, cordate-sagittate, borne
on a pilose stipe; ovary usually glabrous, globose or ellipsoid, the style equaling
or shorter than the ovary, the stigma obtuse or discoid; fruit ellipsoid or sub-
globose, the perianth lobes usually persistent at its base, the pedicel thickened,
the cupule rarely broad and saucer-shaped or deciduous.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 337

Probably about 50 species, or more, all American and most
numerous in tropical North America. Other species occur in
southern Central America, especially in Costa Rica and Panama.
The genus has not been studied critically during the past 50 years,
and is very much in need of attention from a competent taxonomist.
There are no general characters by which the genus may be recog-
nized easily without dissection of the flowers, but Guatemalan
Lauraceae with triplinerved leaves are referable to Phoebe. Many
of the species have penninerved leaves.

Leaves densely tomentose beneath or densely pilose with spreading hairs.

Leaves covered beneath with a very dense and close, rufous tomentum, this

persistent in age, the leaves bicolored P. Salvini.

Leaves not densely and closely tomentose beneath, the leaves not bicolored.
Margins of the leaves conspicuously recurved at the base and often forming a

conspicuous pocket P. amplifolia.

Margins of the leaves not recurved at the base.
Flowers pubescent.

Leaf blades very obtuse or rounded at the base, mostly 3.5-6 cm. wide.

P. mollis.

Leaf blades acute or narrowly obtuse at the base, mostly 2-3.5 cm. wide.

P. Bourgeauviana.
Flowers glabrous.

Leaves acute or subacute at the base, small, mostly 2-2.5 cm. wide.

P. Bourgeauviana.

Leaves rounded or very obtuse at the base, sometimes subcordate, usually

much wider P. helicterifolia.

Leaves glabrous beneath or with inconspicuous appressed pubescence, sometimes
puberulent or sparsely pilose along the nerves or barbate in the axils of the
nerves, or the nerves tomentose or with abundant spreading hairs.
Leaves evidently triplinerved.
Flowers densely pubescent.

Leaves small, mostly 7-8 cm. long; inflorescences usually about 3 cm. long,

corymbif orm, few-flowered P. savannarum.

Leaves large, mostly 12-18 cm. long; inflorescences large, paniculate, many-
flowered P. mexicana.

Flowers glabrous or essentially so P. areolata.

Leaves penninerved.

Flowers glabrous or practically so.

Leaves narrowly lanceolate, mostly 1.5-2.5 cm. wide. . .P. acuminatissima.
Leaves elliptic or lance-elliptic, mostly 3.5-5.5 cm. wide. . .P. padiformis.
Flowers conspicuously and densely pubescent.

Leaves rounded or obtuse at the apex, sometimes abruptly contracted into
a short, very obtuse tip.

Leaves with large perforations or pits beneath in the axils of the nerves.

P. mayana.

Leaves not pitted beneath in the axils of the nerves P. ambigens.

Leaves very acute to long-acuminate at the apex.

Leaves mostly 2.5-4.5 cm. wide P. longicaudata.

Leaves mostly 7-12 cm. wide P. Gentlei.

338 FIELDIANA: BOTANY, VOLUME 24

Phoebe acuminatissima Lundell, Contr. Univ. Mich. Herb.
6: 19. 1941 (type from Mount Ovando, Chiapas). P. saxchanalensis
Lundell, op. cit. 7: 14. 1942 (type from Saxchanal, Chiapas).

Moist or wet, mixed, mountain forest, sometimes in pine forest,
1,300-2,600 meters; Santa Rosa, Sacatepe"quez; Suchitepe"quez;
Huehuetenango; Quezaltenango. Chiapas.

A tree of 5-18 meters, the trunk sometimes 45 cm. in diameter, the bark
smooth, grayish, the branchlets sericeous at first, soon glabrate; leaves on slender
petioles 6-12 mm. long, chartaceous, narrowly lanceolate, mostly 5-10 cm. long
and 1.5-2.5 cm. wide, narrowly long-acuminate or attenuate, acute at the base,
green and glabrate above, paler beneath, sericeous at first, glabrate in age, pen-
ninerved, the lateral nerves 7-11 pairs, the veins closely prominulous-reticulate
beneath; panicles axillary, narrow, usually racemiform, generally half as long as
the leaves or shorter, rather densely appressed-pilose, mostly many-flowered;
flowers slender-pedicellate, yellowish green, glabrous or with a few appressed
hairs; perianth tube short, the segments oval, obtuse or rounded at the apex,
spreading, deciduous; fruit ellipsoid, about 2 cm. long and 1 cm. broad or smaller;
cupule short, 6 mm. broad, the pedicel much thickened.

Phoebe ambigens Blake, Contr. U. S. Nat. Herb. 24: 3. pi. 2.
1922. Aguacatillo.

Known definitely in Guatemala only from Las Playitas, Izabal,
at 120 meters. Honduras, the type from Rodezno, Copan.

A tree as much as 35 meters high with a trunk a meter in diameter, the branch-
lets angulate, strigillose at first, soon glabrate; leaves on naked petioles 1.5-3
cm. long, elliptic to elliptic-oblong, 10-26 cm. long, 3.5-10 cm. wide, obtuse at the
apex or short-pointed with an obtuse tip, attenuate to the very acute base, char-
taceous, glabrous or nearly so in age, often barbate beneath in the axils of the
nerves, prominulous-reticulate on both surfaces, the lateral nerves 5-7 pairs;
panicles axillary, long-pedunculate, pyramidal, lax, many-flowered, shorter than
the leaves or almost equaling them, grayish-puberulent; flowers umbellate in
3's or 4's, 7 mm. long, 15 mm. broad, on pedicels 4-9 mm. long; perianth tube
very short, the segments oval, rounded at the apex, grayish-puberulent; anthers
short-stipitate, truncate at the apex; style equaling the ovary.

Called "guambo" in Honduras. The trunk often has low
buttresses.

Phoebe amplifolia Mez & Donn. Smith ex Donn. Smith, Enum.
PI. Guat. 3: 71. 1893, nomen; Bot. Gaz. 19: 261. pi. 24. 1894.

Moist or wet, mixed, mountain forest, 2,300-3,000 meters; El
Progreso; Quich£ (type from El Jute, Heyde&Lux 3033); Huehue-
tenango. Costa Rica.

A tree of 9-18 meters, the trunk as much as 45 cm. in diameter, the branchlets
stout, densely ferruginous-tomentose; leaves on stout petioles 3 cm. long or shorter,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 339

chartaceous, broadly elliptic to oblong-ovate, 14-30 cm. long, 4.5-16 cm. wide,
acute or subacuminate, obtuse or almost rounded at the base, the margins often
recurved at the base, brown-tomentulose above when young but in age glabrous,
densely and closely brown-tomentose beneath, penninerved, laxly and prominently
reticulate-veined; inflorescence paniculate, densely ferruginous-tomentose, many-
flowered, much shorter than the leaves, long-pedunculate, the branches stout, the
pedicels stout, 3 mm. long or less; flowers greenish white or greenish yellow, densely
ferruginous-tomentulose; perianth segments equal, obtuse; filaments pilose, much
shorter than the anthers, those of the third series of stamens with 2 large sessile
glands at the base; anthers subquadrate, obtuse; staminodia conspicuous, on an
evident stipe; ovary glabrous, the style stout, of about the same length; fruit
about 33 mm. long and 22 mm. broad, ellipsoid, the cupule red, shallow, obscurely
double-marginate, the pedicel much thickened.

Phoebe areolata Lundell, Contr. Univ. Mich. Herb. 7: 13. 1942.

Wet, mixed forest, 300-400 meters; Alta Verapaz (south of
Cubilgiiitz, Steyermark 44494). Chiapas, the type from Saxchanal;
Cockscomb Mountains of British Honduras.

A tree of 18-24 meters, the branchlets glabrous or sparsely sericeous; leaves on
stout petioles 13 mm. long or less, thick-coriaceous, broadly elliptic-ovate to
lance-oblong, 6-11 cm. long, 2.5-4.5 cm. wide, acute or acuminate, rounded or
very obtuse at the base, often abruptly contracted, conspicuously or rather
obscurely triplinerved, with 5-6 pairs of lateral nerves, glabrous or nearly so, the
venation very finely and closely prominulous-reticulate on both surfaces, the leaves
appearing pitted, often barbate beneath in the axils of the nerves; inflorescences
corymbose-paniculate, equaling or shorter than the leaves, long-pedunculate,
few-many-flowered, laxly branched, sparsely sericeous or glabrate, the slender
pedicels 4-7 mm. long; flowers glabrous or practically so, 3-3.5 mm. long, yellowish
green; perianth tube very short, the segments oblong-ovate, obtuse, densely
sericeous within; filaments appressed-pilose, slightly shorter than the anthers;
staminodia large, the stipe thick, appressed-pilose; ovary glabrous, about as long
as the style.

Phoebe Bourgeauviana Mez, Jahrb. Bot. Gart. Berlin 5: 194.
1889. P. purpurea Mez, op. cit. 196 (type from Laraxquica, Alta
Verapaz, Tuerckheim 371).

Moist or wet, mixed or pine, mountain forest, frequently in
wooded swamps, 1,200-2,850 meters; Alta Verapaz; Baja Verapaz;
Zacapa; Chiquimula; El Progreso; Suchitepe'quez ; Solola; Quiche1;
Huehuetenango; Quezaltenango; San Marcos. Southern Mexico;
Honduras.

A slender shrub or tree of 3-6 meters, the branchlets fulvous-villous, glabrate
and brown in age; leaves on naked petioles 8 mm. long or shorter, chartaceous,
mostly lanceolate or lance-oblong, about 10 cm. long and 3 cm. wide or mostly
smaller, acuminate, acute or subobtuse at the base, penninerved, glabrate above,
densely and softly pubescent beneath, the veins prominulous and reticulate

340 FIELDIANA: BOTANY, VOLUME 24

beneath; inflorescences axillary, corymbose-paniculate, pilose or villous, few-
flowered, slender-pedunculate, shorter than the leaves, the pedicels 1-4 mm. long;
flowers usually glabrous, 3 mm. long; perianth tube obsolete, the segments equal,
ovate, subacute; filaments glabrous, very short, those of the third series of stamens
with 2 large acute sessile glands at the base; anthers ovate, acute; staminodia
conspicuous, cordate, sessile; ovary glabrous, the style very short; fruit black,
subglobose, 10-12 mm. long, the cupule small, shallow.

Phoebe Gentlei (Lundell) Standl. & Steyerm. Field Mus. Bot.
23: 117. 1944. Per sea Gentlei Lundell, Contr. Univ. Mich. Herb. 6:
18. 1941.

Moist, mixed forest, at or little above sea level; British Honduras;
endemic; type from Mountain Cow Ridge, Stann Creek Valley,
P. H. Gentle 3288.

A tree, the trunk 25-30 cm. in diameter, the branchlets stout or slender,
grayish-sericeous at first, becoming gray; leaves on stout petioles 2.5 cm. long or
less, chartaceous or subcoriaceous, elliptic-oblong to oval, 12-32 cm. long, 6-14
cm. wide, acute or usually abruptly short-acuminate, acute at the base, pen-
ninerved, with 8-11 pairs of nerves, bright green and very lustrous above, glabrous
in age, densely sericeous or tomentulose beneath at first, glabrate in age; inflores-
cences large, many-flowered, pedunculate, shorter than the leaves, sericeous-
tomentulose, 18 cm. long or less; flowers white, fragrant, sericeous, 4 mm. long,
on pedicels 4 mm. long or shorter; perianth tube very short, the segments equal,
rounded-obovate, rounded at the apex; filaments half as long as the anthers;
staminodia conspicuous, the stipe sparsely pubescent; fruit ellipsoid, 18 mm. long,
11 mm. broad, the cupule 1 cm. long.

Called "timber sweet" and "wild pear."

Phoebe helicterifolia (Meissn.) Mez, Jahrb. Bot. Gart. Berlin
5: 193. 1889. Oreodaphne helicterifolia Meissn. in DC. Prodr. 15,
pt. 1: 123. 1864. Ocotea helicterifolia Hemsl. Biol. Centr. Amer. Bot.
3:73.1882. P. betazensis Mez, op. cit. 192. 1889. P. nectandroides
Mez, op. cit. 194. 1889. Aguacate de monte; Aguacate de mico;
Ismard (Alta Verapaz); Sacsi (Coban, Quecchi) ; Ojche (fide Aguilar).

Moist or wet, mixed forest or often in rather dry forest or thickets,
common in some regions in pine forest, 2,500 meters or less; Alta
Verapaz; El Progreso; Izabal; Santa Rosa; Escuintla; Sacatepe"quez;
Chimaltenango; Suchitepe"quez ; Retalhuleu; Quiche"; Huehuete-
nango. Southern Mexico; Nicaragua.

A large shrub or a tree of 9-15 meters or more, the branchlets stout, densely
villous or villous-tomentose with usually fulvous or grayish hairs; leaves on slender
naked petioles 3 cm. long or shorter, chartaceous or almost membranaceous,
obovate to oblong-obovate or broadly elliptic, sometimes rounded-obovate, mostly
15-25 cm. long and 8-15 cm. wide, abruptly acuminate or short-acuminate,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 341

subcordate to rounded at the base or rarely cuneate, penninerved, usually pilose
or hirsute on both surfaces with long spreading hairs, often glabrate on the upper
surface, the venation sometimes impressed above, very prominent and laxly
reticulate beneath; inflorescences corymbose-paniculate, large and lax, many-
flowered, shorter than the leaves, pedunculate, very sparsely white-hirsute or
almost wholly glabrous; flowers dull yellow or yellowish white, glabrous, on pedi-
cels 3-6 mm. long; perianth tube very short, the segments spreading, equal, ovate,
subacute, 3.5 mm. long; filaments glabrous, equaling or shorter than the anthers,
the glands of the third series small, sessile; anthers elliptic, obtuse; staminodia
small, cordate, glabrous; ovary glabrous, little longer than the style; fruit ellipsoid,
black, about 2 cm. long and 1.5 cm. broad, the cupule saucer-shaped, simple-
margined, the pedicel much thickened.

The three species names listed above have been treated as repre-
senting distinct species by all or most authors, but in spite of the
characters used in Mez's key, it is not apparent how the material
now at hand may be separated into so many distinct groups.

Phoebe longicaudata Lundell, Bull. Torrey Club 64: 548. 1937.

Moist or wet, mixed, lowland forest or thickets, 1,100 meters or
less; Alta Verapaz; Izabal; Chiquimula. British Honduras (type
collected near San Agustin, El Cayo District, C. L. Lundell 6833);
Chiapas; Honduras.

A shrub of 3 meters or usually a tree of 9-12 meters, the branchlets slender,
densely brownish-pilose, soon glabrate; leaves chartaceous, on petioles 4-10 mm.
long, oblong-lanceolate to lance-elliptic, mostly 5-11 cm. long and 2-4.5 cm. wide,
rather abruptly acuminate or caudate-acuminate, acute or obtuse at the base,
usually very lustrous, in age glabrous or nearly so but usually pilose beneath along
the costa with spreading brownish hairs, the veins little if at all elevated above,
laxly prominulous-reticulate beneath; panicles axillary, few-flowered, 4.5 cm. long
or less, much shorter than the leaves, slender-pedunculate, sparsely short-pilose,
the pedicels 2-3 mm. long; perianth white, the tube very short, the segments
subequal, 3 mm. long, sparsely pubescent, tomentose within, elliptic-obovate or
oblong-spatulate, obtuse; filaments short, pubescent, the anthers subquadrate;
staminodia conspicuous, sagittate, the stipe pilose; ovary glabrous; fruit ellipsoid,
black, 1 cm. long, the cupule small, shallow, red.

Called "aguacatillo" and "white laurel" in British Honduras.

Phoebe mayana Lundell, Amer. Midi. Nat. 29: 473. 1943.
Granadillo.

At edge of forest in pasture, 350-450 meters; Alta Verapaz (near
Cubilgiiitz, Steyermark 44649). British Honduras, the type from
Baboon Ridge, Stann Creek Valley, P. H. Gentle 3187.

A tree of 9-15 meters, the trunk 30 cm. or more in diameter, the bark whitish,
the branchlets stout, appressed-puberulent, soon glabrate; leaves chartaceous, on

342 FIELDIANA: BOTANY, VOLUME 24

stout petioles 6-15 mm. long, oblanceolate or oblong-oblanceolate, 7-14 cm. long,
2-4.5 cm. wide, narrowed to an obtuse apex, long-attenuate to the base, blackish
when dried, glabrous or nearly so and lustrous on the upper surface, minutely
sericeous beneath or glabrate, usually barbellate in the axils of the nerves, pen-
ninerved, the lateral nerves 8-10 pairs, the veins not or scarcely elevated on the
upper surface, prominulous and rather laxly reticulate beneath; panicles axillary,
13 cm. long or less, minutely appressed-pubescent at first, glabrate in age; fruiting
pedicels thick, 8 mm. long; fruit oblong, 1.5 cm. long, 5-7 mm. broad, rounded at
the apex, the cupule very shallow, 5 mm. broad.

The species is known only from fruiting material, and its generic
position is therefore problematical.

Phoebe mexicana Meissn. in DC. Prodr. 15, pt. 1: 31. 1864.
Persea mexicana Hemsl. Biol. Centr. Amer. Bot. 3: 72. 1882. Agua-
catillo (Pete"n).

Moist or wet, mixed forest or in thickets, 2,400 meters or less,
chiefly at very low elevations; Pete"n(?); Izabal; El Progreso;
Retalhuleu; San Marcos. Southern Mexico; British Honduras;
Honduras; Costa Rica.

A tree of 9-12 meters, or sometimes lower, the trunk as much as 25 cm. in
diameter, the branchlets fulvous-tomentose at first, soon glabrate; leaves on
petioles 1-3 cm. long, coriaceous or chartaceous, narrowly ovate to ovate-lance-
olate or elliptic-lanceolate, mostly 11-20 cm. long and 4-8 cm. wide, rather
abruptly acuminate or long-acuminate, acute at the base, usually conspicuously
triplinerved, barbate beneath in the axils of the nerves, elsewhere glabrous or
obscurely and very sparsely sericeous, the veins prominulous-reticulate beneath;
inflorescence densely whitish-pilose or sericeous, many-flowered, pyramidal-
paniculate, pedunculate, equaling or shorter than the leaves, the pedicels 2-3 mm.
long; flowers white or whitish, densely pilose, 3 mm. long; perianth tube obsolete,
the segments equal, ovate, acute, suberect; filaments about equaling the anthers,
sparsely pubescent, those of the third series with 2 rather large, subglobose, sessile,
basal glands; anthers glabrous, elongate-ovate; staminodia large, subcordate-
sagittate, acuminate, very sparsely pubescent dorsally, the stipe shorter, pilose;
ovary glabrous, globose, the style 2-3 times as long, slender; fruit ellipsoid, 12 mm.
long, 7 mm. broad.

Called "aguacate negro" in Honduras.

Phoebe mollis Mez, Jahrb. Bot. Gart. Berlin 5: 192. 1889.
(?)P. belizensis Lundell, Contr. Univ. Mich. Herb. 6: 20. 1941 (type
from Stann Creek Valley, Mountain Cow Ridge, British Honduras,
P. H. Gentle 3304).

Dense wet mixed forest, 2,000 meters or lower; Alta Verapaz;
Guatemala; Suchitepe"quez ; Quezaltenango ; San Marcos. Southern
Mexico; British Honduras.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 343

A tree, the branchlets fulvous-tomentose; leaf blades coriaceous or subcoria-
ceous, oblong-lanceolate, mostly 10-15 cm. long and 5-6 cm. wide, long-acuminate,
rounded or cordate at the base, on petioles 1 cm. long or shorter, the lateral nerves
10 or fewer pairs; inflorescences slender, paniculate, few-flowered, fulvous-tomen-
tose at first, glabrescent in age, the peduncles 8 cm. long or less; flowers pubescent,
3 mm. long, on pedicels about 1.7 mm. long; perianth segments rounded-ovate,
subacute, 1.7 mm. long; ovary glabrous, with a short style.

Phoebe padiformis Standl. & Steyerm. Field Mus. Bot. 23: 117.
1944.

Dense wet mixed forest, 550-2,000 meters; endemic; Huehue-
tenango; Quezaltenango (type from Colomba, A. F. Skutch 1367).

A tree of 6-15 meters, the trunk as much as 30 cm. in diameter, the branchlets
slender, at first rather sparsely appressed-pilose, soon glabrate, striate-angulate;
leaves on petioles 6-10 mm. long, oblong-elliptic or oblanceolate-oblong, 8-11 cm.
long, 3-5.5 cm. wide, abruptly acute or short-acuminate with an obtuse tip, acute
at the base, when young grayish-sericeous but soon glabrate and at maturity almost
wholly glabrous, densely white-barbate beneath in the axils of the nerves, pen-
ninerved, the lateral nerves about 6 pairs, the veins not elevated on the upper
surface, prominulous and laxly reticulate beneath; panicles axillary, racemiform
or racemose, half as long as the leaves or shorter, with very short lower branches,
laxly few-many-flowered, glabrous, the pedicels 3-4 mm. long, straight; flowers
greenish white, glabrous, 3 mm. long; perianth tube almost none, the segments
broadly elliptic, obtuse, suberect; filaments slender, twice as long as the anthers or
longer, glabrous or pilosulous near the base; anthers small, oblong, obtuse; glands
of the filaments of the third series of stamens large, thick, cordate, sessile; stami-
nodia conspicuous, ovate, acute, short-stipitate; ovary globose, glabrous, about
equaling the thick style.

Phoebe Salvini (Mez) Lundell, Contr. Univ. Mich. Herb. 6:
23. 1941. Ocotea Salvini Mez, Jahrb. Bot. Gart. Berlin 5: 264. 1889.

Dense, moist or wet, mixed, mountain forest, 1,800-3,200 meters;
endemic; El Progreso; Guatemala; Chimaltenango (type from Las
Calderas, Volcan de Acatenango [not Fuego as labeled], Salviri);
Solola; San Marcos.

A tree of 9-15 meters or more, the branches stout, terete, densely and closely
ferruginous-tomentose; leaves on stout petioles 2.5 cm. long or shorter, coriaceous,
elliptic to ovate-elliptic, 9-16 cm. long, 3.5-8.5 cm. wide, acute or acuminate,
acute to rounded at the base, the margins recurved at the base and forming a
pocket, bright green and lustrous on the upper surface, glabrous or nearly so, the
venation prominent and closely reticulate, covered beneath with a very dense,
close, ferruginous tomentum; panicles subpyramidal, shorter than the leaves,
densely ferruginous-tomentose, many-flowered, the stout pedicels 1-3 mm. long;
flowers densely ferruginous-tomentulose; perianth tube obsolete, the segments
broadly ovate, acute; filaments sparsely pubescent, short, those of the third series
with 2 small sessile globose basal glands; anthers suborbicular, rounded at the apex;

344 FIELDIANA: BOTANY, VOLUME 24

staminodia conspicuous, cordate-sagittate, the stipe sparsely pilose; ovary gla-
brous, the style slightly shorter; mature fruit oval-globose, 3 cm. long, 2 cm. broad,
broadly rounded at the apex; cupule saucer-shaped, 11 mm. broad, double-
marginate, the short pedicel very thick.

Among all Lauraceae of Guatemala this is recognized readily
by the very dense, close, ferruginous tomentum of the lower leaf
surface.

Phoebe savannarum Standl. & Steyerm. Field Mus. Bot. 23:
118. 1944.

Known only from the type, Alta Verapaz, along stream bordering
forest, savanna between base of Cerro Chinaja at Sachaj and
Sacacao, 150-180 meters, Steyermark 45712.

A tree of 9 meters, the branches very slender, terete, densely sordid-pilosulous
with ascending hairs, soon glabrate and blackish brown; leaves on naked or nar-
rowly marginate petioles 3-5 mm. long, chartaceous, blackish brown when dried,
elliptic to ovate-elliptic or oblong-elliptic, 6-8.5 cm. long, 2-3.5 cm. wide, abruptly
long-caudate-acuminate with an obtuse tip, obtuse at the base, lustrous and
glabrous above, the veins not elevated, beneath minutely and inconspicuously
pilosulous on the nerves, elsewhere glabrous, triplinerved, the veins densely
prominulous-reticulate; panicles axillary, 3-4 cm. long, few-flowered, on long
slender peduncles, cymiform, minutely pilosulous or puberulent, the pedicels
puberulent, scarcely more than 2 mm. long; flowers white, densely and minutely
puberulent or strigillose, 2.5 mm. long; perianth tube very short, the segments
broadly elliptic, obtuse, spreading, densely tomentulose within; outer anthers
large, suborbicular, rounded at the apex, on very short filaments; glands of the
third series of stamens globose, sessile; staminodia short-stipitate, conspicuous,
broadly ovate, obtuse; ovary glabrous, globose-ovoid, the style short, thick.

HERNANDIAGEAE

Trees or shrubs, sometimes woody vines; leaves alternate, without stipules,
simple, entire or lobate, penninerved or palmate-nerved, oil cells and cystoliths
often present in the foliage; flowers perfect or unisexual, small, in axillary or
pseudo-terminal long-pedunculate corymbose panicles; perianth segments usually
in 2 valvate 3-5-parted series, or in 1 imbricate 4-10-parted series; stamens 3-5,
in a single series, opposite the outer perianth segments; anthers 2-celled, introrse,
dehiscent by valves, the filaments often with basal glands; ovary inferior, 1-celled,
the ovule 1, pendulous, anatropous; fruit dry, large and winged, or included in an
enlarged cupule; seed without endosperm, the embryo straight, the cotyledons
large.

Four genera and about 35 species are known, in the tropics of
both hemispheres. Only the following genera occur in America.

Leaves lobate; fruit bearing 2 long, narrowly spatulate wings Gyrocarpus.

Leaves entire; fruit not winged.

Leaves peltate; flowers surrounded by bracts Hernandia.

Leaves not peltate; flowers not bracteate Sparattanthelium .

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 345

GYROCARPUS Jacquin

Deciduous trees with thick branches; leaves mostly clustered at the ends of
the branches, petiolate, broad, palmate-nerved, usually trilobate; inflorescences
terminal, umbel-like or lax and many-flowered, not bracteate; flowers small,
perfect or unisexual, the staminate numerous, the pistillate and perfect ones few;
sepals of the staminate flowers 4-7, concave, pubescent; stamens 4-7, some of them
often reduced to staminodia, the filaments thick, pubescent; sepals 8 in the pistil-
late flower, tomentose, 2 of them large and quadrangular, 4 of them small and
united with the large ones as lateral appendages, the other 2 free and caducous;
ovary tomentose, the style very short; fruit subglobose, subtended by the 2 large
sepals which are greatly accrescent, elongate, and linear-spatulate; cotyledons
foliaceous, spirally twisted.

The genus consists of a single species.

Gyrocarpus americanus Jacq. Stirp. Amer. 282. pi. 178, f. 80.
1763. Volantin; Palo hediondo; Campdn (Zacapa) ; Tregador (Chiqui-
mula) ; Titirillo (Gualan, fide Record) ; Felipdn.

Dry hillsides or plains, ascending from near sea level to about
1,400 meters; Zacapa; Chiquimula; El Progreso; Jutiapa; Santa
Rosa; Escuintla; Guatemala; Retalhuleu; Quiche"; Huehuetenango.
Southern Mexico; Salvador; Nicaragua; Costa Rica; northern South
America; tropical Asia, Africa, and Australia.

Often only a shrub but usually a small or medium-sized tree, reported to
attain a height of 20 meters but usually lower, the trunk and branches thick, with
rather smooth, whitish bark; leaves usually on very long petioles, the blades large
and thin, broad, often 30 cm. wide or larger, entire or usually palmately 3-5-lobate,
the lobes entire, acuminate, truncate or broadly cordate at the base, green and
glabrate above, paler beneath, at first often white-tomentose, finally glabrate;
flowers small and greenish, the 2 largest calyx segments in age 10-12 cm. long
and about 1 cm. wide, rounded at the apex, suberect or spreading, tomentulose
or glabrate; nut ellipsoid, about 2 cm. long, densely tomentulose; seed broadly
oblong, terete, the testa coriaceous.

Called in Yucatan "ciis" or "xkis" (Maya), "volador," "palo
hediondo"; in Salvador "tambor," "lagarto," and "corroncha de
lagarto." The tree is abundant in the dry hilly parts of Guatemala,
about Amatitlan, Zacapa, and Sacapulas, and also extends down
upon the Pacific plains. It is unattractive in appearance, leafless
during the dry season, at which time it produces flowers and fruits.
The latter are of distinctive form, of a "parachute" type, so that
when they fall from the tree they spin in the air, to come to the
ground usually at some distance from the tree. The foliage has a
disagreeable odor. The wood, apparently, is not utilized unless for
firewood. It is soft, white, and of light weight.

346 FIELDIANA: BOTANY, VOLUME 24

HERNANDIA L.

Trees; leaves alternate, usually palmate-nerved, entire, long-petiolate; flowers
monoecious, in lax corymbiform panicles, each branch terminating in an involucrate
cluster of 2-3 flowers, the central flower pistillate and sessile, the lateral staminate
and short-pedicellate; staminate flower with 6-8 perianth segments; stamens 3,
opposite the outer segments; pistillate flower subtended at the base by a cupule,
the perianth segments 8, 4 glands present opposite the outer segments; ovary
inferior, the style short, the stigma dilated, irregular, peltate-discoid; fruit a
globose black hard nut, more or less 8-costate, included in the greatly enlarged
and inflated cupule; cotyledons flattened, somewhat rugose.

About 14 species, in the tropics of both hemispheres. Another
Central American species, H. didymantha Bonn. Smith, with oblong
leaves, is found in Costa Rica and Panama, and H. stenura Standl.,
with linear-caudate leaves, has been described from Costa Rica.

Hernandia sonora L. Sp. PI. 981. 1753. H. guianensis Aubl.
PL Guian. 849. pi. 329. 1775. H. peltata Sesse" & Mocino, Fl. Mex.
ed. 2. 213. 1894. Tanajita (Tinajita?).

Lowland, wet or dry forest, at 750 meters or less; Alta Verapaz;
Izabal; Escuintla. Veracruz; Honduras; Costa Rica; Colombia to
the Guianas; Old World tropics.

A large shrub or a small tree said to attain sometimes a height of 20 meters, the
branchlets thick, glabrous or nearly so; leaves large, membranaceous or charta-
ceous, on very long petioles, broadly ovate, usually peltate and attached near the
base, mostly 14-20 cm. long and 7-12 cm. wide, short-acuminate, rounded or
truncate at the base, glabrous or nearly so, 5-nerved; panicles borne in the upper
leaf axils, long-pedunculate; bracts oblong or spatulate, 1 cm. long or less; seg-
ments of the staminate flower fleshy, elliptic, obtuse, 6.5 mm. long or less, densely
tomentulose outside, densely pilose within; stamens 3, the filaments glabrous;
segments of the pistillate perianth elliptic, 6 mm. long or less; cupule surrounding
the fruit in age inflated and globose, with a small opening at the apex, about 6 cm.
in diameter; fruit ellipsoid-ovoid, longitudinally 6-8-costate, sessile or short-
stipitate, 2.5 cm. long, umbonate.

Said to be called "palo de chicalpexte" in Veracruz; "hoja de
tamal," "mano de leon," "tambor" (Honduras).

SPARATTANTHELIUM Martius

Shrubs, usually scandent; leaves trinerved or triplinerved, entire; flowers
small, polygamo-dioecious, in axillary or subterminal, panicled cymes, without
bracts; perianth of 4-7 subequal segements subimbricate in bud; perianth tube
in the perfect flowers united with the ovary, in the staminate flowers almost
obsolete; fertile stamens 4-5, opposite the perianth segments, the filaments fili-
form, glandless; anthers oblong-linear, the cells introrsely dehiscent; staminodia

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 347

none; style cylindric, the stigma subcapitate, small; fruit dry, ovoid or ovoid-
ellipsoid, smooth, the endocarp coriaceous or ligneous.

About 12 species in tropical America, ranging from Guatemala
to Bolivia and Brazil. Only one species is known from Central
America.

Sparattanthelium guatemalense Standl. Proc. Biol. Soc.
Wash. 37: 51. 1924.

Type collected in wet thicket near Puerto Barrios, Izabal, at
sea level, Standley 25066 in 1922. Also in the Atlantic coast of
Honduras.

A shrub or small tree 3-6 meters high, perhaps sometimes scandent, the slender
branches glabrous; leaves on slender petioles 1.5-3.5 cm. long, oblong-lanceolate,
11-13 cm. long, 3-4.5 cm. wide, abruptly long-acuminate, obtuse at the base,
glabrous, 3-nerved, the 2 lateral nerves extending two-thirds the distance to the
apex; panicles slender-pedunculate, about 7 cm. long, many-flowered, the very
slender branches minutely gray-puberulent, the pedicels puberulent, often twice
as long as the calyx but sometimes shorter than the segments; calyx 4-parted,
0.5 mm. long, minutely gray-puberulent.

PAPAVERACEAE. Poppy Family
Reference: Friedrich Fedde, Pflanzenreich IV. 104: 1-430. 1909.

Herbs or rarely shrubs or trees, the sap usually colored; leaves alternate,
entire to lobate or dissected, without stipules; flowers perfect, regular, often large
and showy; sepals 2-3, free, caducous; petals usually 4-6, hypogynous, free,
deciduous, imbricate; stamens hypogynous, usually numerous, free, the filaments
filiform; anthers erect, 2-celled, the cells longitudinally dehiscent; ovary free,
1-many-celled, the placentae parietal; style short or obsolete; stigmas as many
as the placentae, distinct or confluent, often adnate to the apex of the ovary and
radiately spreading; ovules usually numerous, sometimes few, anatropous, ascend-
ing or horizontal; fruit capsular, dehiscent by pores or valves, rarely indehiscent;
seeds globose or subreniform, smooth or scrobiculate, the raphe cristate or naked;
embryo minute, the endosperm oily-flesby.

About 25 genera, widely distributed but chiefly in temperate
regions. No other genera are found in Central America.

Plants trees or shrubs; petals none; fruit usually 1-seeded Bocconia.

Plants herbaceous; petals present; fruit many-seeded.

Capsule linear; flowers bright yellow; leaves divided into linear lobes.

Eschscholtzia.

Capsule globose, obovoid, or oblong.

Leaves prickly-margined Argemone.

Leaves not prickly Papaver.

348 FIELDIANA: BOTANY, VOLUME 24

ARGEMONE L.

Herbs or rarely shrubs, glaucous, with yellow sap; leaves incised-pinnatifid,
generally spinose-dentate and rigid-setose; flowers large, white or yellow, rarely
red or purple, the buds erect; sepals 2-3; petals 4-6; stamens numerous; ovary
with 4-6 placentae, the style very short or almost obsolete, the stigma depressed-
dilated, the lobes radiating from the center; capsule oblong, dehiscent by short
valves; seeds scrobiculate.

About 10 species, in temperate and tropical regions of America,
one of the species naturalized in the Old World.

Argemone mexicana L. Sp. PL 508. 1753. A. ochroleuca Sweet,
Brit. Fl. Gard. 3: pi. 242. 1828. A. mexicana var. ochroleuca Lindl.
Bot. Reg. pi. 1343. 1830. Chicalote; Cardosanto; Cajhuoc, Ixmucur
(Quich^ ; fide Tejada); Kixatucan (Totonicapan fide Tejada);
Sajquix (Huehuetenango fide Tejada) ; another name reported, with-
out locality, is Cahhouc.

Dry or moist fields or thickets, often along roadsides or in sandy
stream beds, ascending from sea level to about 2,500 meters, or
perhaps even higher; Pete"n; Zacapa; Jalapa; Jutiapa; Santa Rosa;
Escuintla; Guatemala; Sacatepe*quez; Chimaltenango; Quiche";
Totonicapan; Huehuetenango; Quezaltenango; probably in all the
other departments. Mexico and British Honduras to Panama;
West Indies; South America; naturalized in the Old World.

A coarse stout annual a meter high or less, sometimes perhaps enduring for
more than one year, glabrous but armed throughout with numerous sharp, rather
stiff prickles, very glaucous, the foliage somewhat mottled; leaves sinuate-pin-
natifid, 8-20 cm. long, the lobes short and broad, with prickly margins; flowers
solitary at the ends of the branches, each subtended by 2-3 leaf -like bracts; sepals
3, prickly, tipped with a stout terete spinose horn; petals 6, white, creamy white,
or yellow, commonly 2-3 cm. long; capsule 4-6-valvate, 4-5 cm. long, armed with
few stiff spines; seeds globose, numerous, reticulate, about 2.5 mm. in diameter.

The Maya names of Yucatan are reported variously as "kix-
zaclol," "kixcanlol," "canlal," "ixcanlol." The typical form of the
species has white petals. In var. ochroleuca the petals are bright or
pale yellow. Both forms occur commonly in Guatemala, the white-
flowered plants apparently the more common, although in some
regions, as about Escuintla and Amatitlan, yellow flowers are more
plentiful. The two forms often grow in the same region or even in
the same spot. In the highlands of the Occidente the prickly poppy
is conspicuous during the dry months, since it is one of the few plants
that continue to grow during the cold season. Apparently the sheep,
which destroy most vegetation at this time, do not touch it. It

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 349

produces large quantities of seeds and springs up abundantly in
cornfields and other cultivated ground. The flowers vary greatly
in size of petals. The seeds may well be poisonous. They are
sometimes administered as an emetic or purgative but their use is
perhaps somewhat dangerous. In Guatemala the latex is sometimes
placed in the eyes to relieve eye affections. It is recorded that the
Indians of San Miguel Acatan (Huehuetenango) employ the plant
to cure drunkenness, but the manner of administering it is not
described. The seeds are reported to contain about 36 per cent of
oil. This has been used in Mexico in soap-making but is said not
to be very satisfactory. The oil has purgative or vomitive-purgative
properties.

BOCCONIA L.

Reference: J. Hutchinson, Bocconia and Macleaya, Kew Bull.
275-282. 1920.

Shrubs, small trees, or large herbs, often glaucous, glabrous or pubescent, the
sap yellow or orange; leaves large, lobate, dentate, or entire; flowers small, in
large terminal panicles; sepals 2; petals none; stamens numerous or sometimes of
definite number; ovary with 2 placentae, these sterile or bearing only a few ovules,
only a basal ovule fertile; style short or somewhat elongate, the stigma lobes
oblong or linear, erect or recurved; capsule more or less stipitate, ellipsoid, dehis-
cent to the base, the valves recurved; seed usually 1, surrounded at the base by a
pulpy aril.

Ten species are known, in tropical America. Only the following
occur in Central America.

Leaves sessile, the blade attenuate to the point of insertion B. vulcanica.

Leaves distinctly petiolate, the petiole often long.

Leaves not lobate but merely serrate, crenate, or entire.

Leaves glaucous and glabrous beneath B. glaucifolia.

Leaves not glaucous beneath, puberulent or villosulous B. gracilis.

Leaves pinnate-lobate.

Leaves cuneate at the base, the lobes attenuate or acuminate; lower leaves

deeply pinnate-lobate B. arborea.

Leaves mostly truncate or rounded at the base, the lobes obtuse or subacute;
lower leaves lobed less than halfway to the costa B. frutescens.

Bocconia arborea Wats. Proc. Amer. Acad. 25: 141. 1890.

Quiebra-muelas ; Palo de matates; Llora-sangre ; Sangre de chucho;
Saupe de chucho (fide Aguilar).

Damp or wet thickets or forest, frequently in oak forest, some-
times in second growth, 500-2,630 meters; Chiquimula; Jalapa;
Jutiapa; Santa Rosa; Escuintla; Guatemala; Sacatepe*quez ; Chi-

350 FIELDIANA: BOTANY, VOLUME 24

maltenango; Suchitepe'quez; Quiche"; Quezaltenango; San Marcos.
Central and southern Mexico; Salvador; Costa Rica; Panama.

A shrub or tree 2.5-6 meters high with few thick branches, the young branches
tomentose; leaves as much as 45 cm. long and 30 cm. wide but usually smaller,
deeply pinnate-lobate, glabrous above, grayish or brownish beneath and more or
less tomentose, in age sometimes glabrate, the lobes narrow, serrate, mostly long-
acuminate or attenuate; panicles large, often 20 cm. long or more, usually recurved,
at least in age, the flowers pedicellate, the pedicels 1 cm. long or less; sepals acumi-
nate, usually 10-12 mm. long, glabrous; stamens about 12; fruit about 7 mm. long,
stipitate, recurved, ellipsoid, 1 cm. long or less, crowned by the persistent and
elongate style.

This is an abundant and showy plant at many places in the
Occidente and in the Pacific bocacosta. It is sometimes planted
for ornament in the parks, as at Huehuetenango. It is used in
Guatemala as a dye plant, the bark giving a yellow color that was
said to have been used by the aboriginal inhabitants of Mexico for
dyeing feathers and other objects. The orange sap is a common
remedy for toothache in Guatemala. The plant, studied by Mexi-
can pharmacists, is said to contain several alkaloids similar to those
of Papaver, and these, injected beneath the skin, cause local anes-
thesia. They have been used by surgeons of Mexico City while
performing operations. The wood is reported to be used sometimes
in Mexico for tanning. In Salvador the tree is called "tine-canasta"
and "brasil."

Bocconia frutescens L. Sp. PL 505. 1753. Sangre de toro;
Camotillo (Pete'n).

Moist thickets or forest, ascending from little above sea level to
about 2,800 meters; Pete'n; Alta Verapaz; El Progreso; Jalapa;
Guatemala; Quezaltenango; San Marcos. Southern Mexico to
British Honduras; Costa Rica; West Indies.

Usually a shrub of 1.5-3 meters, simple or branched, the young branches
somewhat lanate-tomentose; lower leaves petiolate, usually truncate or rounded
at the base, 15-35 cm. long, 10-20 cm. wide, glabrous or nearly so above, somewhat
tomentose beneath or glabrate, often glaucous but sometimes green, the lobes
short, very obtuse to rounded, repand-denticulate; panicles 40 cm. long or less,
lax and many-flowered, the pedicels 1 cm. long or less; sepals abruptly acuminate,
pale, about 1 cm. long or often somewhat shorter; stamens about 16; fruit narrowly
or broadly ellipsoid, 6-8 mm. long, usually acute at each end, long-stipitate; seed
6 mm. long, somewhat muricate.

Collections from Pete'n and British Honduras are noteworthy for
their small sepals, although not apparently unique. It appears that
the sepals in this genus often enlarge considerably during and after

STANDLE.Y AND STEYERMARK: FLORA OF GUATEMALA 351

anthesis. The yellow or orange sap is bitter, acrid, and has a dis-
agreeable odor. Fedde recognizes two forms of the species, both
found in Guatemala, but neither appears to be of much significance
from a taxonomic standpoint. They are f. glaucescens (Kuntze)
Fedde (Pflanzenreich IV. 104: 218. 1909), with leaves glaucous and
glabrate beneath; and f. subtomentosa (L'He"r.) Fedde (loc. cit.),
with leaves pale green or green beneath, often copiously tomentose.

Bocconia glaucifolia Hutchinson, Kew Bull. 281. 1920.
B. integrifolia var. mexicana DC. Prodr. 1: 121. 1824 (type collected
by Sesse" and Mocifio). B. integrifolia f. mexicana subforma glau-
cescens Fedde, Pflanzenreich IV. 104: 220. 1909. Saupe (fide
Aguilar).

Moist or wet forest, 1,600-2,600 meters; Quich£ (type from San
Miguel Uspantan, Heyde & Lux 2899); Huehuetenango; endemic.

A shrub 3.5 meters tall, glabrous thr6ughout or nearly so, the branches
glaucous; leaves long-petiolate, oblong-oblanceolate, 10-30 cm. long, 3-8 cm.
wide, acute or subobtuse, attenuate to the base or sometimes rounded, subentire
or undulate-serrate, very glaucous beneath; panicles 35 cm. long or less, peduncu-
late, pendent, lax and many-flowered; pedicels 10 mm. long or less, glaucous;
sepals acuminate, about 1 cm. long, glaucous; stamens about 12.

It is of interest to record that a specimen of this species is in the
Sesse" and Mocifio Herbarium (No. 1807), and since the plant is
unknown from Mexico — although it may occur there — it seems
probable that the collection was made in Guatemala. This is one
of the most distinct species of the genus, and it is curious that Fedde
considered it merely a subform of a form; but his treatment of this
genus, as well as of some other groups of Papaveraceae, is notoriously
inadequate.

Bocconia gracilis Hutchinson, Kew Bull. 280. 1920. (?)£.
integrifolia var. Seleri Fedde, Pflanzenreich IV. 104: 220. 1909 (type
from Yalambohoch, Huehuetenango, Seler 2700). Achote de monte.

Dense wet forest, 1,100-1,650 meters; Alta Verapaz (type from
Pansamala, Tuerckheim 1236; collected also in the regions of Tactic
and Coban); Huehuetenango; endemic.

A shrub of 2-3 meters, the young branches brownish-tomentose; leaves slender-
petiolate, elliptic-oblong or oblanceolate-oblong, 8-25 cm. long, 3-9 cm. wide,
acute or acuminate, short-cuneate at the base, coarsely or rather finely and
remotely serrate, glabrous above, brownish-tomentulose or glabrate beneath,
green; panicles lax, many-flowered, 20 cm. long or less, pedunculate, the slender

352 FIELDIANA: BOTANY, VOLUME 24

pedicels 1-1.5 cm. long; sepals abruptly acuminate, glabrous, about 1 cm. long;
stamens 12 ; ovary stipitate.

This has been reported from Guatemala under the name B.
frutescens var. cernua DC.

Bocconia vulcanica Bonn. Smith, Bot. Gaz. 16: 1. 1891.
B. oUanceolala Lundell, Contr. Univ. Mich. Herb. 4: 5. 1940 (type
from Volcan de Tacana, Chiapas, Matuda 2916). Cerbatana;
Quiebra-muelas.

Moist or wet, usually dense forest, sometimes in Cupressus
forest, 2,000-3,800 meters; Sacatepe"quez (type from Volcan de
Agua, 3,200 meters, J. D. Smith 2172); Jalapa; Chimaltenango;
Quezaltenango; San Marcos. Adjacent Chiapas (Volcan de Tacana).

A shrub or tree 3-8 meters tall, the thick branches glabrous or nearly so;
leaves oblanceolate to obovate-oblong, mostly 10-35 cm. long and 5-11 cm. wide,
acute or short-acuminate, long-attenuate to the sessile base, closely and rather
finely serrate, glabrous, green beneath; panicles recurved, usually narrower than
in other species, pedunculate, many-flowered, often dense; pedicels 5 mm. long,
or in fruit elongate and recurved; sepals caudate-acuminate, 1 cm. long or less,
glabrous; stamens 10-15; fruit about 1 cm. long, ellipsoid, long-beaked by the
persistent style.

This species, like the others, is used as a remedy for toothache,
the seeds or fruit being placed in cavities. The plant is abundant
in many parts of Quezaltenango and San Marcos. It was stated on
the Volcan de Agua that the plant is poisonous but it is hard to
imagine how one could obtain a fatal dose of it.

ESCHSCHOLTZIA Chamisso

Glabrous, more or less glaucous annuals or perennials; leaves much cleft,
with linear segments; flowers yellow, long-pedunculate, often large and showy;
torus more or less cupular-dilated at the apex, the petals and stamens thus per-
igynous; sepals coherent, dehiscent as a cap; petals 4; stamens numerous; placentae
of the ovary 2; style short, the stigma divided into 4-6 linear divergent lobes;
capsule linear, 10-sulcate, dehiscent to the base, the valves rigid, recurved; seeds
not cristate.

A group of 10 or more species, in western United States and
northern Mexico. One species has been introduced into cultivation
in many parts of the earth.

Eschscholtzia californica Cham, in Nees, Horae Phys. Berol.
73. pi. 15. 1820. Chorchitas; Popa de oro; Popi (a corruption of the
English word "poppy").

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 353

Planted commonly in gardens at low, middle, and rather high
elevations; noted as more or less naturalized as a weed in a corn-
field at Chichicastenango. Native of California.

Plants annual or perennial, diffusely branched, 30-60 cm. tall; leaves tripin-
natifid, glaucous, the segments linear or nearly so; flowers 3-5 cm. broad, variable
in size, the petals bright yellow, flabellif orm ; capsules 5-6 cm. long; seeds globose,
reticulate.

The California poppy is a favorite garden flower and often is
grown in the parks. Large bunches of flowers often are on sale in
the markets but this is not a good cut flower for vases, since the
blossoms do not last long in water. The petals are open in sunshine
but the flowers close in the evening and during cloudy weather.
Called "adormidera" in Salvador.

PAPAVER L. Poppy

Herbs, sometimes hispid, often glaucous, with milky sap; leaves usually lobate
or dissected; peduncles elongate, the buds nutant, the flowers often large and
showy, red, purple, white, or yellow; sepals commonly 2; petals 4 or rarely 6;
stamens numerous; placentae of the ovary 4 to many, intruded, ovuliferous on all
sides, the ovary more or less septate; stigma at the apex of the ovary disk-like,
convex or pyramidal, adnate to the ovary, the lobes radiating from the center;
capsule globose, ovoid, or oblong, dehiscent below the apex by transverse pores
between the placentae; seeds scrobiculate.

About 100 species are recognized by Fedde, while Bentham and
Hooker in 1862 give the number as 14! They are mostly natives of
the Old World, but two species are indigenous in California and
Baja California.

Stems hispid; plants not glaucous P. Rhoeas.

Stems glabrous; plants glaucous. . P. somniferum.

Papaver Rhoeas L. Sp. PI. 507. 1753. Adormidera.

Grown commonly for ornament in parks and gardens of the
central uplands, and more or less throughout the higher regions, as
well as in Alta Verapaz. Native of Europe; occasionally naturalized
in North America.

Plants erect, branched, mostly 50-80 cm. tall, hispid with long spreading
hairs; lower leaves petiolate, the upper smaller and sessile, pinnatifid, with lanceo-
late acute serrate lobes; flowers 5-10 cm. broad, usually scarlet with a dark center;
capsule subglobose or turbinate, glabrous. *

This is the corn or field poppy of Europe, whose cultivated forms
are known in the United States by the name "Shirley poppy."

354 FIELDIANA: BOTANY, VOLUME 24

Papaver somniferum L. Sp. PL 508. 1753. Adormidera;
Azumbador; Amapola.

Grown commonly for ornament in gardens and parks, through-
out the cooler and cold regions, also about Coban; often running wild
in old fields in Quezaltenango and San Marcos and some other
regions, but not persisting long. Native of Europe and Asia.

Plants tall and stout, often a meter high, glabrous or nearly so, very glaucous,
sparsely branched; leaves sessile and clasping by a broad base, undulate, lobate,
or dentate; flowers 7-10 cm. wide or larger, bluish white with darker center or
often pink or red; capsule large, globose, glabrous.

This is the well-known opium poppy, from whose pods the drug
opium and its derivatives are obtained, this product having its
origin in eastern Asia. From the seeds is obtained poppy oil. The
seeds are much used in the United States for sprinkling upon rolls,
to which they impart a distinctive flavor. Poppies are grown in
large quantities in the highlands of Guatemala for sale in the markets.
In these regions they are somewhat persistent in cornfields but
probably would not persist long unless the supply of seeds was
renewed from cultivated plants.

CRUCIFERAE. Mustard Family

Annual or perennial herbs, rarely suffrutescent, the sap watery, often acrid,
the pubescence of simple or often branched hairs; leaves alternate, simple or dis-
sected, the basal ones often forming a rosette; stipules none; flowers perfect,
regular, racemose, the racemes terminal or axillary, usually ebracteate; corolla
white, purple, pink, or dark red; sepals 4, free, the inner ones sometimes saccate
at the base, usually imbricate; petals 4, rarely none, cruciately spreading, entire
or bilobate, convolute or imbricate; glands usually present at or above the base
of the torus, usually 4 and opposite the sepals; stamens 6 and of 2 lengths, or often
more or fewer, the filaments subulate, the longer ones often 1-dentate; anthers
2-celled or rarely 1-celled, longitudinally dehiscent, basifixed, oblong-cordate or
sagittate, sometimes linear and twisted; ovary sessile or rarely stipitate, 2-carpel-
late, 1-celled or usually 2-celled; style simple, the stigmas 2, or sometimes connate;
ovules generally numerous, horizontal or pendulous, campylotropous or amphi-
tropous; fruit usually a silique or silicle, i.e. elongate and narrow, but very variable
in form, 2-celled or 1-celled, usually 2-valvate, the valves separating from the
septum, sometimes indehiscent; seeds small, often mucilaginous when wet, fre-
quently winged or marginate; endosperm usually none, sometimes present and
oily; cotyledons mostly plano-convex.

Genera about 225, widely distributed, best represented in tem-
perate regions. In the tropics most of the species, except a few
weedy ones, are found only in the mountains. No other genera are
known in Central America.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 355

Fruit transversely 2-articulate, the terminal joint beak-like; plants of seashores.

Cakile.
Fruit not transversely articulate; plants found rarely if ever on seashores.

Fruit indehiscent; cultivated plants or rare weeds Raphanus.

Fruit dehiscent.

Pods orbicular to oblong, usually little more than twice as long as broad,
often about as broad as long.

Fruit not compressed; cultivated plants, the leaves linear or nearly so.

Lobularia.
Fruits strongly compressed; native plants or introduced weeds.

Fruit compressed parallel with the partition, twice as long as broad or
more; flowers yellow Draba.

Fruit compressed contrary to the partition ; flowers white or pale yellow.

Pods obtriangular, not at all winged Capsella.

Pods rounded or oval, often winged at the apex Lepidium.

Pods linear, often much elongate, several or many times as long as broad.

Petals 1.5-2 cm. long or larger, usually deep red or purple, never yellow.

Pubescence of stellate hairs; cultivated plants Matthiola.

Pubescence of appressed hairs, each with 2 branches; native plants.

Erysimum.
Petals usually much less than 1 cm. long, various in color.

Plants densely and finely stellate-pubescent throughout; leaves mostly
2-pinnatifid Descurainia.

Plants glabrous, or the pubescence of simple hairs, a few branched hairs

sometimes present.
Flowers yellow, or rarely white in one cultivated species.

Flowers 1.5-2 mm. long; pods terminated by a very short style,

never long-rostrate Rorippa.

Flowers much larger; pods often long-rostrate Brassica.

Flowers white or purple, never yellow.

Leaves pinnately divided, with 3-many leaflets or segments.

Pods compressed; plants terrestrial or often growing in wet soil.

Cardamine.

Pods not compressed; plants aquatic Nasturtium.

Leaves simple.

Cauline leaves auriculate-clasping Romanschulzia.

Cauline leaves narrowed to the base Lamprophragma.

Armoracia lapathifolia Gilibert (A. rusticana Gaertn. Mey. &
Scherb.), native of Europe, is planted rarely in Guatemala, and has
been noted at Quezaltenango and Coban. It seems to thrive in
gardens, but is little used on the table in Guatemala, except perhaps
by foreigners. The English name is "horse-radish," the Spanish
"rabano picante." The large thick roots are exceedingly acrid, and
when grated are much used in the United States as a condiment,
to flavor meat and pickles.

356 FIELDIANA: BOTANY, VOLUME 24

BRASSICA L.

Reference: O. E. Schulz, Pflanzenreich IV. 105: 21-84. 1919.
L. H. Bailey, The cultivated Brassicas, Gentes Herb. 1: 53-108.
1922.

Annual or biennial herbs, sometimes of longer duration, glabrous or with
pubescence of simple hairs; leaves alternate, the lowest often rosulate, petiolate,
sessile, or amplexicaul, simple or pinnately parted; outer sepals oblong, obtuse at
the apex and more or less cucullate, the inner ones usually ovate, subacute, sub-
saccate at the base; petals obovate, unguiculate, usually yellow, rarely white;
stamens 6, the anthers obtuse or pointed, yellow; ovary cylindric, few-many-ovu-
late, the ovules generally 1-seriate, the style usually long; stigma capitate or
somewhat bilobate, usually slightly wider than the style; silique narrowly or
broadly linear or oblong, straight or sometimes flexuous, the valves convex, usually
terminated by a conic beak; valves 1-nerved; seeds globose or rarely ovoid, pendu-
lous, not marginate, brown; cotyledons longitudinally conduplicate, sessile, deeply
emarginate.

Species about 30, most of them native in the Mediterranean
region. None are native in America but some have become natural-
ized widely as weeds. Because of the fact that most of the species
have been in cultivation for many centuries as food plants, the Latin
nomenclature is highly complicated, and scarcely two authors agree
as to what the various elements should be called. The nomencla-
ture used here is that employed by Dr. L. H. Bailey in his horticul-
tural publications. The following key to species includes only the
forms likely to be found in Guatemala in a more or less wild state.

Cauline leaves dilated and amplexicaul at the base B. campestris.

Cauline leaves neither dilated nor amplexicaul at the base, usually petiolate.
Pedicels about 4 mm. long; pods appressed to the rachis, 12-25 mm. long.

B. nigra.

Pedicels mostly 6-10 mm. long or longer; pods erect or ascending, not appressed

to the rachis, mostly 25-50 mm. long.
Principal leaves pinnately parted to the costa and with 2-3 pairs of leaflets;

pods 2-3.5 mm. thick, the beak 6-10 mm. long B. juncea.

Principal leaves simple, coarsely dentate; pods 1-2 mm. thick, the beak
usually 3-7 mm. long B. integrifolia.

Brassica alboglabra L. H. Bailey, Gentes Herb. 1: 79. 1922.

Plants apparently referable to this species were collected along
a roadside between Finca Pirineos and Calahuache", Quezaltenango,
Steyermark 35201. The species is cultivated in China for its edible
foliage, and has been introduced into cultivation in the United
States. We have not seen it in Guatemalan markets. It is an annual
with very glaucous, glabrous foliage, the leaves oval, petiolate, not

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 357

clasping; the inflorescence is much elongate, the flowers large and
white, rather than of the usual yellow.

Brassica campestris L. Sp. PI. 666. 1753. Moztaza. Field
mustard.

A common weed in cultivated or abandoned fields, waste ground,
roadsides, and various other situations, abundant in many parts of
Guatemala, 1,200-3,300 meters; Alta Verapaz; Zacapa; Jalapa;
Sacatepe"quez; Chimaltenango; Quiche"; Huehuetenango; Quezalte-
nango; San Marcos; doubtless to be found in other departments.
Native probably of Europe, but widely naturalized in other regions,
and found in many parts of Central America.

Plants annual, glabrous, glaucous, with a slender root, the stems erect, a
meter high or less, usually branched; basal and lowest cauline leaves lyrate-pin-
natifid, the upper cauline leaves narrowly or broadly oblong, obtuse or acute, often
entire, dilated and clasping at the base; flowers 7-10 mm. long, bright yellow;
pedicels spreading or ascending, 5-15 mm. long or more; pods erect-spreading,
3-6 cm. long or even longer, the beak 1-2 cm. long, conical at the base; seeds 1 mm.
in diameter, dark brown.

This and B. Rapa, the turnip, are closely related, and some of the
forms passing as B. campestris are probably seedlings from neglected
turnip patches. The country people of Guatemala recognize this
fact, and are quite as likely to call the plant "nabo" as "mostaza."
In some regions of the earth this species is grown for its oil-yielding
seeds. In Central America the young plants are cooked and eaten.

Brassica caulorapa (DC.) Pasq. Cat. Ort. Bot. Napoli 17:
1867. B. oleracea var. caulorapa DC. Syst. Nat. 2: 586. 1821.
Colinabo. Kohlrabi.

Kohlrabi has been in cultivation for many centuries and is
unknown in the wild state, although presumably of European origin.
Like cabbage, it is a biennial, and its distinguishing character is the
thick turnip-like swelling of the stem, just above the ground. Very
fine kohlrabi is grown in Quezaltenango, at Almolonga and Zunil,
and it is fairly common in the large market of Quezaltenango. It is
seen also in the Coban market, but the Indians do not care for it,
or for cabbage either. It is not a popular vegetable in the United
States.

Brassica integrifolia (Willd.) Rupr. Fl. Ingr. 1: 96. 1860.
Sinapis integrifolia Willd. Hort. Berol. 14. pi. 14- 1806.

358 FIELDIANA: BOTANY, VOLUME 24 -

A weed in waste ground, at or little above sea level; British
Honduras. Doubtless of Old World origin, and widely dispersed
there; occasional in tropical America, but in Central America
infrequent.

An erect annual a meter high or less, glabrous, usually very glaucous; lower
leaves long-petiolate, elliptic or oval, large, coarsely and irregularly dentate, acute
to rounded at the apex, cuneate at the base; upper cauline leaves sessile or short-
petiolate, lanceolate or oblanceolate, often almost entire; flowers yellow, long-
pedicellate.

Brassica juncea (L.) Coss. Bull. Soc. Bot. France 6: 609. 1859.
Sinapis juncea L. Sp. PI. 668. 1753. Mostaza.

Often planted for food, and found occasionally as an escape or
weed in waste ground; Pete"n; Alta Verapaz; Guatemala; not com-
mon in Guatemala or elsewhere in Central America. Native
probably of Asia.

A glabrous, more or less glaucous annual, a meter high or less, usually branched,
with a slender taproot; lower leaves large, broadly oblong or obovate in outline,
lyrately lobate or divided, the upper cauline leaves simple, narrower, lobate,
dentate, or entire; flowers bright yellow, often fragrant, 7-10 mm. long; pods
4-7 cm. long, erect or ascending on stout pedicels, the beak 3-10 mm. long; seeds
1 mm. in diameter, mostly very dark brown.

Var. japonica (Thunb.) L. H. Bailey, in which the upper cauline
leaves are incised-pinnatifid, with narrow lobes, has been found in
cultivation in Guatemala (San Marcos) . B. juncea is often planted
in Guatemala for its leaves, which are cooked and eaten, and sold
frequently in the markets. The seeds are used locally as a condi-
ment, and medicinally. They are sometimes boiled with meat and
other foods to flavor them.

Brassica nigra (L.) Koch in Roehling, Deutschl. Fl. ed. 2. 4:
713. 1833. Sinapis nigra L. Sp. PL 668. 1753. Mostaza.

Cultivated occasionally for food, sometimes escaping in waste or
cultivated ground, as in Jalapa and San Marcos, but an infrequent
weed in Central America. Native of Eurasia, but widely planted
in other regions, and often naturalized as a weed.

Plants annual, erect, usually glabrous, sometimes hispid, with a slender
taproot, usually a meter high or less; leaves mostly petiolate, the lower ones pin-
nately parted, the terminal segment much larger than the others, finely and closely
serrate; upper cauline leaves simple, narrowly ovate to oblong or linear; flowers
bright yellow, 5-8 mm. long; pods erect and appressed to the rachis, 1-2 cm. long,
somewhat 4-sided, the beak only 1-3 mm. long; seeds 1-1.5 mm. in diameter, dark
brown.

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 359

Known in the United States as "black mustard." This plant is
there the chief source of the mustard used on the table. The leaves
may be cooked and eaten like those of all or most other species.

Brassica oleracea L. Sp. PL 667. 1753.

The wild plant, presumably the ancestor of what Dr. Bailey
justly calls "a marvellous progeny," is a native of the coasts of
western and southern Europe, often growing in calcareous soil or on
chalk or limestone cliffs. In general appearance the wild plant looks
much like the collards cultivated so commonly in the southern
United States. It has a somewhat elongate stalk, with the large,
broad, very glaucous leaves spreading from the lower thickened
portion of the stem.

Brassica oleracea var. acephala DC. Syst. Nat. 2: 583. 1821.

This variety includes various plants known in the United States
as "kale." The common kale has not been seen by us in Guatemala,
but probably it has been planted or at least tested there. Another
form referred to this variety by Bailey is collards, which we have
noted a few times, as at San Lucas, Sacatepe'quez. Apparently it is
grown as a curiosity. Rather frequent in cultivation for ornament
in Guatemala is what is presumably the tree kale, B. oleracea var.
acephala sub var. palmifolia DC. This is a tall plant with a thick
simple stem a meter high or often taller, bearing near the top many
crowded leaves, which usually are purplish and much curled or
fringed. It is grown chiefly for ornament, but the leaves are some-
times sold in the markets for food. At Totonicapan the market
women gave them the name of "colinabo" but they were not leaves
of kohlrabi, to which that Spanish name properly applies. This
purple-leafed kale is seen mostly in the highlands, in Chimaltenango
and westward through Los Altos, usually only one or two plants in
gardens or parks. There are many plants in the cemetery at Tactic
(Alta Verapaz).

Brassica oleracea var. botrytis L. Sp. PL 667. 1753. Coliflor.
Cauliflower.

Distinguished from cabbage by its dense whitish head of fasciated
flower clusters, surrounded by whorls of large leaves. The plants
bloom the second year from seed, producing panicles of whitish
flowers. Cauliflower thrives in the cooler parts of Guatemala,
especially at elevations of 2,000 meters or more. It would be hard

360 FIELDIANA: BOTANY, VOLUME 24

to find finer cauliflower than that produced in the gardens of Almo-
longa and Zunil, which reaches the Quezaltenango market still wet
with dew. It is sold commonly with all the leaves attached, these
being cooked and eaten like cabbage or collards. Dr. F. Webster
McBryde, at the suggestion of the senior author, questioned a large
number of Guatemalan people, rich and poor, as to what fruits and
vegetables they like best. The results were not very satisfactory,
except that when asked what was their favorite vegetable, the
majority named cauliflower. This may well be because it is scarcer
and more expensive than most other vegetables, or partly because it
usually appears on the table properly cooked, in contrast with cab-
bage, which, in Guatemala as well as in the United States, usually
is boiled for an hour or more, until it is indigestible and has lost all
its original flavor.

Brassica oleracea var. capitata L. Sp. PI. 667. 1753. Repollo;
Col; Culic (Jacaltenango, Huehuetenango). Cabbage.

A plant of European origin, presumably derived from the wild
B. oleracea, cultivated for many centuries and now represented by
innumerable varieties. It is a biennial, blooming the second year
from seed. Cabbage is one of the common vegetables of Guatemala,
either raw or cooked, and is grown almost everywhere at middle and
high elevations, not or rarely in the lowlands. Much is cultivated
through the verano under irrigation. On the slopes of Volcan de
Zunil there are large fields planted on the very steep slopes of white
sand. Here the plants grow luxuriantly through the dry season
because the slopes are covered every night with dense fog and clouds
that provide abundant moisture, in spite of the fact that there is no
rain. The sand is so loose and the mountain side so steep that the
plants maintain a rather precarious foothold. The senior author has
seen a large boulder rolling down the slopes, large cabbage heads
hurtling down behind it like so many cannon balls. Savoy cabbage
(sometimes called B. oleracea var. Sabauda L., treated by Bailey as
a mere form of var. capitata) is cultivated rarely in Guatemala.
Red cabbage (B. oleracea var. rubra L.) has not been noted in Guate-
mala, but probably is planted occasionally, at least in the German
fincas. Its leaves are deep purple-red.

Brassica oleracea var. gemmifera (DC.) Zenker, Fl. Thuering.
15: 2. 1836(7). B. oleracea var. bullata DC. subvar. gemmifera DC.
Syst. Nat. 2: 583. 1821. Repollitos; Colitos. Brussels sprouts.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 361

This is distinguished by its tall thick stems, bearing large soft
buds, 2-3 cm. in diameter and resembling small cabbages, along
almost the whole length of the stem. The flowers are produced
the second year from seed. Brussels sprouts is not a very common
vegetable of Guatemala but it is grown in the gardens of Almolonga
and Zunil and doubtless elsewhere, and is sold rather commonly in
the Quezaltenango and Guatemala markets.

Brassica oleracea var. italica Plenck, Icon. PI. Med. 6: 29.
pi 534- 1794. Broccoli.

This is somewhat similar to cauliflower, but the head is composed
of loose, green and purplish, thick and somewhat fasciated branches,
which, unlike the fasciated branches of cauliflower, bear normal
flowers. The plants are eaten before the flowers open. This plant
was not seen by the writers in Guatemala but doubtless it has been
planted there. It was introduced rather recently into the United
States and has become popular and common only during the last
ten years or so.

Brassica pekinensis (Lour.) Rupr. Fl. Ingr. 96. 1860. Sinapis
pekinensis Lour. Fl. Cochin. 400. 1790. B. Pe-Tsai L. H. Bailey,
Cornell Univ. Agr. Exp. Sta. Bull. 67: 178, 190. 1894. Pe-tsai;
Chinese cabbage.

Grown occasionally in Guatemala for food, but infrequent.
Introduced from China not many years ago, it has become rather
common in United States markets. When well grown, the plants
form elongate narrow heads of soft, bright light-green leaves, that
look more like giant lettuce than cabbage. The leaves are eaten
either raw in salads or cooked.

Brassica Rapa L. Sp. PI. 666. 1753. Nabo. Turnip.

Turnips are grown commonly in Guatemala at middle and high
elevations, and many of those seen are of excellent quality and some
of great size. The plant is normally a biennial but in Central
America it probably blooms the first year from seed. The turnips
of Guatemala are rather uniform in appearance, and doubtless are
grown, like most other vegetables, from seed imported from the
United States. Some in the Totonicapan market were somewhat
elongate and almost oblong and very large. It is possible that they
were rutabagas (B. Napobrassica Mill.), but they looked more like
common turnips. In the Huehuetenango market, and probably

362 FIELDIANA: BOTANY, VOLUME 24

elsewhere, there are sold what are called "nabitos," young turnip
plants with ample foliage and slender taproots, to be cooked and
eaten like mustard.

CAKILE Miller
Reference: 0. E. Schulz, Pflanzenreich IV. 105, pt. 2: 18-28. 1923.

Succulent, glabrous, annual or biennial herbs, usually growing along seashores,
the stout stems branched, often decumbent; leaves pinnatifid to entire; flowers
purple, pink, or white, in ebracteate racemes, the pedicels short, thickened in
fruit; sepals erect, the outer ones linear, obtuse and subcucullate at the apex, the
2 inner ones broadly oblong, subacute; petals unguiculate, obovate, rounded or
subemarginate at the base, closely veined; stamens 6, the anthers oblong, obtuse;
pistil broadly cylindric, sessile, biarticulate, the lower joint short, 1-ovulate, the
upper joint thick, usually 1-ovulate; stigma depressed-capitate, narrower than the
style; fruit a silique, composed of 2 joints, more or less tetragonous, 3-nerved on
each side; lower joint turbinate, often 2-corniculate, 1-seeded; upper joint easily
separating from the lower, usually broader, gradually attenuate to the beak;
seeds rather large, oblong, somewhat rugulose; cotyledons oblong.

Different authors have varied greatly in their treatment of this
genus of seaside plants, but Schulz recognizes 4 species, in Europe,
northern Africa, western Asia, and North and Middle America.
Only one is found in Central America, where it is confined, appar-
ently, to the Atlantic coast, and is not common.

Cakile lanceolata (Willd.) 0. E. Schulz in Urban, Symb.
Antill. 3: 504. 1903. Raphanus lanceolatus Willd. Sp. PI. 3: 562. 1800.

Reported (as C. maritima Scop.) from Livingston, Izabal,
Tuerckheim 8835. British Honduras (keys off the coast; specimens
in flower, the specific determination uncertain). Southern Florida;
Yucatan Peninsula of Mexico; Honduras; West Indies; Colombia
and Venezuela.

Plants stout, erect or decumbent, the stems 20-50 cm. long; leaves petiolate,
oblong-elliptic to linear-oblanceolate, obtuse, attenuate at the base, undulate-
dentate; fruiting racemes rather lax; petals 6-8 mm. long, mostly white, obovate;
ovary 2-4-ovulate; fruit elongate, 18-30 mm. long, 4 mm. thick, subterete and
somewhat sulcate; lower joint cylindric, the upper joint 2-4 times as long, dagger-
like, obtuse or acutish.

The single British Honduras collection is in flower and can not
be determined with certainty, if certainty is possible in this genus.
The specific characters are found in the fruits. The British Hon-
duras plant may be rather C. edentula (Bigel.) Hook. var. alacranensis
(Millsp.) 0. E. Schulz, the common species of Yucatan, in which
the terminal joint of the fruit is ovoid.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 363

CAPSELLA Medicus. Shepherd's-purse

Slender and usually low annuals, glabrous or with pubescence of branched
hairs, the stems simple or branched; radical leaves forming a rosette, entire or
lobate; flowers very small, white, slender-pedicellate, in elongate racemes; sepals
spreading, not saccate; stamens free; silique usually obcuneate, laterally com-
pressed and flattened, the valves strongly compressed, carinate, the septum very
narrow, membranaceous; style short, the stigma sessile; seeds numerous, not
winged.

Half a dozen species, as treated by most authors, in temperate
regions of both hemispheres. No species are native in Central
America, but several have been described from Mexico.

Gapsella Bursa-pastoris (L.) Medic. Pflanzengatt. 1: 85. 1792.
Thlaspi Bursa-pastoris L. Sp. PI. 647. 1753. Bolsa de pastor.

Waste or cultivated ground, often a weed in gardens or corn
fields, or in dooryards, sandy fields or on sandbars along streams,
pastures, coffee plantations, 1,300-3,900 meters; Alta Verapaz;
El Progreso; Jalapa; Guatemala; Sacatepe"quez; Chimaltenango;
Huehuetenango; Quezaltenango; San Marcos. Native of Europe
but now naturalized in many temperate regions of the earth; a
common weed of the United States; mountains of Costa Rica.

Plants slender, erect, from a long slender root, usually 40 cm. high or less,
simple or branched, stellate-pubescent below, glabrous above; basal leaves lobate
or pinnatifid, forming a large dense rosette, 4-10 cm. long; cauline leaves few,
lanceolate, auriculate at the base, dentate or entire; flowers white, about 2 mm.
long, the slender pedicels spreading or ascending; pods triangular, cuneate at the
base, 4-8 mm. long, truncate or emarginate at the broad apex; seeds 10-12 in
each cell.

This is a rather frequent weed in the mountains of Guatemala,
but it is apparently rare in other parts of tropical America and in
Mexico. The leaves have a flavor similar to that of Lepidium.
They may be cooked and eaten as a pot herb, but so far as we know
are not used thus in Guatemala.

CARDAMINE L.

Reference: 0. E. Schulz, Monographic der Gattung Cardamine,
Bot. Jahrb. 32: 280-623. 1903.

Annual, biennial, or perennial herbs, glabrous or with pubescence of simple
hairs, sometimes with rhizomes, usually low, simple or branched; leaves mostly
petiolate, simple or variously pinnatisect; flowers small, generally racemose, the
racemes often corymbiform in an thesis, usually ebracteate; sepals oblong or ovate,
erect-spreading; petals unguiculate, rarely none, usually obovate, white, pink, or

364 FIELDI AN A: BOTANY, VOLUME 24

purple; stamens 6, the anthers oblong, sagittate at the base; ovary cylindric,
4-10-ovulate, the ovules 1-seriate, the ovary attenuate to the usually filiform
style; stigma somewhat 2-lobate, minute; silique narrowly or broadly linear,
straight, compressed, the valves plane, acuminate, not thickened on the margins,
not or obscurely nerved; seeds 1-seriate, elliptic or quadrate-oblong, more or less
compressed, not marginate or rarely narrowly winged.

Species more than 100, in almost all cold and temperate regions,
in the tropics found in the mountains. One or two additional
species occur in southern Central America.

Leaflets 3-5 or sometimes 9-13, the principal ones 2-4.5 cm. long; flowers 6-10
mm. long.

Leaflets 9-13 C. balnearia.

Leaflets usually 3, rarely 5.

Racemes leafy-bracteate at the base C. fulcrata.

Racemes naked C. innovans.

Leaflets 3 or usually more numerous, the largest 2 cm. long and most of them

much smaller; flowers 5 mm. long or shorter.
Plants cespitose from a lignescent root, erect; leaflets all alike, oblanceolate,

entire, scarcely more than 2 mm. wide C. eremita.

Plants annual, or perennial with very slender, soft stolons, erect or procumbent;
leaflets often dissimilar, the terminal one orbicular or nearly so, most of the
leaflets much more than 2 mm. wide.
Plants perennial, with slender stolons, procumbent; stems with several or

numerous leaves C. flaccida.

Plants annual, erect; stems naked or with a single leaf C. jejuna.

Cardamine balnearia Standl. & Steyerm. Field Mus. Bot. 23:
157. 1944.

Known only from the type, Quezaltenango, wet mossy bank,
Aguas Amargas, western slope of Volcan de Zunil, 2,450 meters,
Standley 83332.

An erect perennial herb about 35 cm. high, glabrous throughout, the root
perpendicular, emitting very numerous slender roots, apparently not stoloniferous;
stems simple, naked near the base, very densely leafy about the base for a short
distance; leaves very numerous, long-petiolate, about 14-15 cm. long, 9-13-folio-
late, the slender petiole naked, at the base somewhat dilated and almost clasping;
leaflets alternate or the upper ones opposite, thin, often remote, on petiolules
4-6 mm. long, broadly ovate to ovate-oblong or lance-oblong, 1-3 cm. long, 4-12
mm. wide, subacute to very obtuse, rounded to subacute and often oblique at the
base, with a few remote subulate-tipped teeth or very shallowly and remotely
lobulate, the terminal leaflet generally larger than the lateral ones; racemes
terminal, simple or sparsely branched from the base, leafy-bracteate only at the
base or naked, about 14 cm. long, lax, many-flowered, the slender pedicels 7-12
mm. long, ascending; flowers 6 mm. long, the sepals purplish, almost 3 mm. long;
petals white, tinged with purple; immature siliques 3.5 cm. long, 0.8 mm. broad,
the style 3 mm. long, scarcely narrowed upward.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 365

The plant probably is rare, for the senior author has collected
several times at the type locality and found it but once.

Cardamine eremita Standl. & Steyerm. Field Mus. Bot. 23:
53. 1944.

On rocks in alpine situations in pine forest, 3,300-3,700 meters;
endemic; Huehuetenango (Sierra de los Cuchumatanes; type col-
lected between Tojquia and Caxin Bluff, Steyermark 50143; collected
also at Tunima).

A glabrous perennial, erect or ascending, more or less cespitose, the slender
caudex laxly branched, the few stems 8-20 cm. long, sparsely leafy; radical leaves
2-4 cm. long, about 7-foliolate, the segments sessile, small, thick, linear-oblance-
olate or oblanceolate, 3-7 mm. long, 2.5 mm. wide or narrower, obtuse or very
obtuse, gradually attenuate to the base, entire; cauline leaves similar to the basal
ones, petiolate, the lowest flower usually leafy-bracted at the base; racemes with
few or rather numerous flowers, in fruit as much as 7 cm. long, usually shorter,
the flowers sometimes somewhat secund, the slender fruiting pedicels ascending,
5-9 mm. long; sepals oblong, 2.5 mm. long, rounded at the apex, white-margined,
tinged with purple; petals white, 5-6 mm. long; pods linear, 20-27 mm. long,
scarcely 1 mm. wide, gradually long-attenuate at the apex, the style 1-1.8 mm.
long; seeds few, brownish, marginate.

Cardamine flaccida Cham. & Schlecht. Linnaea 1: 21. 1826.

Wet fields or hillsides, often on wet shaded stream banks or along
irrigating ditches, sometimes in rocky stream beds, 1,200-3,500
meters; Zacapa; Guatemala; Sacatepe"quez; Chimaltenango; Hue-
huetenango; Quezaltenango; San Marcos. Mexico; Costa Rica;
widely distributed in South America.

A slender weak succulent perennial, with very slender stolons, the stems com-
monly 10-40 cm. long, sometimes forming large masses of foliage, procumbent and
rooting near the base, often much branched from the base, abundantly leafy;
lower leaves with 3-4 pairs of leaflets; terminal leaflet orbicular or reniform, 1-1.5
cm. long and as wide, or sometimes larger, obscurely and coarsely crenate or sub-
entire, petiolulate; lateral leaflets somewhat smaller, obliquely ovate, 1-2-crenate
on each side or almost entire, petiolulate, glabrous; racemes very lax, often
elongate, few-many-flowered, the fruiting pedicels 8-15 mm. long; flowers white,
3.5-4 mm. long; sepals ovate, 2 mm. long; pods about 22 mm. long and 1.2 mm.
wide, attenuate to a slender style 0.5-1 mm. long; seeds 1 mm. long, fulvous,
marginate.

This plant is confined to very wet soil and may even grow in
shallow water. The species is a highly variable one, and Schulz
recognizes numerous subspecies and varieties which seem to be
vaguely limited.

366 FIELDIANA: BOTANY, VOLUME 24

Cardamine fulcrata Greene, Pittonia 3: 155. 1897. Quilete
(Jalapa) ; Jazmin (Quezaltenango) ; Yacan-chamel (Huehuetenango) ;
Berro amargo (fide Aguilar).

Moist or wet, usually mixed forest, sometimes in Alnus forest
and growing in white sand, rarely somewhat epiphytic, 2,000-2,900
meters; El Progreso; Jalapa; Chimaltenango; Huehuetenango;
Quezaltenango; San Marcos. Southern Mexico.

Plants perennial, erect or ascending, sometimes almost suffrutescent below,
30-60 cm. high, often much branched, the stems sparsely or densely puberulent;
leaves all or mostly 3-foliolate, large, long-petiolate, mostly cauline; terminal leaflet
ovate or ovate-oblong, petiolulate, 2.5-9.5 cm. long, acute or acuminate, crenate-
serrate, the lateral leaflets similar but smaller, sparsely pilose with short white
hairs; racemes leafy-bracteate at the base or higher, the bracts 3-foliolate or 3-
lobate; fruiting pedicels 1.5 cm. long or shorter; flowers 6-10 mm. long, the sepals
3.5-4 mm. long; petals white, rounded at the apex; pedicels erect-spreading in
fruit; pods 3.5-4 cm. long, 2 mm. wide, the style 1.5-5 mm. long; seeds 2.5 mm.
long, greenish brown.

The name "quilete," if properly given to this plant, would
indicate that it was used as a pot herb, which may well be the case.
Almost all plants of this family, if young and tender, may be eaten
either raw or cooked.

Cardamine innovans 0. E. Schulz, Bot. Jahrb. 32: 417. 1903.
Napscul (Huehuetenango); Chilillo de agua (fide Aguilar).

Moist or wet, usually dense, mixed forest, 1,300-3,000 meters;
Alta Verapaz; Chimaltenango (type from mountains above Tecpam,
F. C. Lehmann 1475); Solola; Quiche"; Huehuetenango; Quezalte-
nango; San Marcos; endemic.

Perennial, erect or decumbent, the stems 20-50 cm. long, bearing few or rather
numerous leaves, simple or branched above, glabrous or nearly so; leaves mostly
3-foliolate, sometimes 5-foliolate, the leaflets petiolulate, large, the terminal one
ovate or oblong-ovate, 3-4.5 cm. long, acute or obtuse, undulate-dentate or repand-
dentate, the lateral leaflets smaller, all glabrous or nearly so; racemes lax, 5-12-
flowered, the flowers white or purplish, 7-8 mm. long; pods on pedicels 1.5-2 cm.
long, suberect, 4-5 cm. long, attenuate to the style, this about 6 mm. long.

This species is very closely related to C. fulcrata, and the two
probably should be combined.

Cardamine jejuna Standl. & Steyerm. Field Mus. Bot. 23: 54.
1944.

Known only from the type, Huehuetenango, forested summit,
Cerro Pixpix, above San Ildefonso Ixtahuacan, 2,800 meters, Sierra
de los Cuchumatanes, Steyermark 50569.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 367

A dwarf annual, erect from a long slender root, the stems very slender, simple,
naked or bearing a single leaf, 3-4-flowered, glabrous; radical leaves 1.5-2.5 cm.
long, usually 3-foliolate, sometimes 5-foliolate or simple, the lateral leaflets petiolu-
late, the terminal one long-petiolulate, all the leaflets 3-6 mm. long and as wide,
obtuse or rounded at the apex, truncate or rounded at the base, entire or usually
shallowly 3-lobulate, the lobes mucronate, the leaflets glabrous beneath, some-
times hispidulous above; racemes short and lax, the pedicels very unequal, as
much as 13 mm. long, filiform; sepals pale green, obovate-oval, 2 mm. long, rounded
at the apex, pale-margined, glabrous; petals white, 3 mm. long; pods glabrous,
narrowly linear, 2.5 cm. long, 0.7 mm. wide, short-attenuate at the base, gradually
narrowed at the apex into a beak almost 5 mm. long, the style 1.5 mm. long.

DESCURAINIA Webb & Berthelot
Reference: O. E. Schulz, Pflanzenreich IV. 105: 305-346. 1924.

Chiefly annuals, erect or ascending, often much branched, the pubescence
mostly of stellate hairs, often grayish or tomentose, sometimes with gland-tipped
hairs; leaves pinnatisect, often much divided, the lower ones petiolate, the upper
sessile or nearly so; flowers minute, white or yellowish, the racemes almost always
ebracteate, the fruiting pedicels filiform; sepals erect-spreading, the outer ones
narrowly oblong, the inner ones broader, obtuse at the apex and not cucullate, not
saccate at the base; petals spatulate, generally equaling or shorter than the sepals;
stamens 6, often longer than the petals; ovary sessile, 6-many-ovulate, the style
very short, the stigma depressed-capitate; siliques short, 4.5 cm. long or shorter,
2-celled, 2-valvate, the valves nerved; seeds 1-2-seriate, oblong or ellipsoid,
mucilaginous when wet; cotyledons oblong, as long as the radicle.

About 40 closely related species, chiefly in temperate regions.
Only the following is found in Central America.

Descurainia streptocarpa (Fourn.) 0. E. Schulz, Pflanzen-
reich IV. 105: 317. 1924. Sisymbrium streptocarpum Fourn. Recherch.
Crucif. 58. 1865.

Usually a weed in gardens or old grain fields, sometimes on sand-
bars along streams, 1,500-2,550 meters; Guatemala; Quiche*; Totoni-
capan; Quezaltenango. Central and southern Mexico.

An erect annual, generally a meter high or less, usually much branched, green
or grayish, the stems minutely stellate-pubescent, eglandular; lower leaves ovate
in outline, with about 4 pairs of segments, these divided into small narrow obtuse
lobes, the upper leaves with narrower segments, finely stellate-pubescent, often
very densely so or more or less stellate-tomentose; racemes, at least below, with
small, pinnatifid or entire bracts, dense at first, usually greatly elongate in age and
many-flowered, the fruiting pedicels 12 mm. long or shorter; sepals 2 mm. long,
glabrous; petals yellowish or greenish yellow, sometimes white, equaling the sepals;
ovary 18-30-ovulate; pods erect or ascending on the spreading pedicels, 8-15 mm.
long, often somewhat curved, 1 mm. thick or less, glabrous, acute, the style very
short, the valves 1-nerved; seeds 1-seriate, oblong or oval, brown.

368 FIELDIANA: BOTANY, VOLUME 24

This is one of the commonest weeds of gardens and old fields
in the valley of Quezaltenango. The seeds are much sought by
small birds. The species has been reported from Guatemala as
Sisymbrium Galeottianum Fourn. It is by no means certain that D.
streptocarpa is distinct from the Mexican D. impatiens (Cham. &
Schlecht.) 0. E. Schulz. If they should be united, the latter is the
older name.

DRABA L.

Reference: O. E. Schulz, Pflanzenreich IV. 105: 16-343. 1927.

Plants annual or perennial, usually herbaceous, the stems scapose or leafy,
usually pubescent; leaves simple, the basal ones often forming rosettes, petiolate,
the cauline leaves sessile; flowers small, white or yellow, naked or bracteate;
sepals erect-spreading, the outer oblong or elliptic, the inner ones broader, rounded
or obtuse at the apex; petals unguiculate, obovate-cuneate, generally emarginate;
stamens 6, the anthers ovoid or oblong, obtuse; ovary sessile, 4-many-ovulate, the
style conic or filiform, very short or elongate, the stigma depressed-capitate;
siliques usually short and broad, ovate or lanceolate, straight or curved, some-
times contorted, 2-celled, 2-valvate, the valves usually flat, the median nerve
inconspicuous; seeds 2-seriate, ovoid or ellipsoid, compressed, usually not winged,
not mucilaginous when wet; radicle slender, the cotyledons ovate, equaling the
radicle.

Species about 250, mostly in cold or temperate regions, widely
distributed. Only the following is known in Central America.

Draba volcanica Benth. PI. Hartweg. 82. 1841.

Alpine among rocks or in rock crevices on or near mountain
summits, chiefly on the summits of the higher volcanoes, sometimes
in alpine meadows, rarely along the borders of small alpine streams,
3,600-4,200 meters; Sacatepe"quez (type from the crater of Volcan
de Agua, Hartweg 571); Huehuetenango (Sierra de los Cuchu-
matanes); Quezaltenango (Volcan de Santa Maria); San Marcos
(volcanoes of Tajumulco and Tacana). High peaks of central and
Southern Mexico.

Plants biennial or perennial, erect or decumbent, usually from a thick tap
root, solitary or clustered, sometimes 35 cm. long but usually much shorter, simple
or often much branched, leafy, sparsely or rather densely pubescent with partly
simple and partly branched hairs; basal leaves forming dense rosettes, oblong-
spatulate or oblanceolate, 2-5 cm. long, obtuse, entire, narrowed to the base,
densely white-ciliate; cauline leaves smaller, oblong, entire or remotely denticulate,
glabrate above, covered with scattered, mostly 2-furcate hairs beneath; racemes
at first short and dense, in age many-flowered and often much elongate, the pedicels
2-5 mm. long; flowers yellow, the sepals 2 mm. long, sparsely pubescent, often
purplish; petals about equaling the sepals; ovary 4-16-ovulate, sparsely pubescent

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 369

or glabrous, the style conic, usually very short; fruiting pedicels spreading at a
right angle, the pods ovate or lanceolate, 4-7 mm. long, acute, the valves nervose-
striate; seeds ovoid, dark brown, rugose-striate.

This species belongs to a small group of closely related species of
the highest mountains of Mexico and Guatemala and of the South
American Andes. The Guatemalan material is rather uniform.
Most divergent is a collection (Steyermark 35538) from the summit
of the Volcan de Tajumulco, in which the siliques are exceptionally
small and have a conspicuous style. The plant appears to be a
shade form.

ERYSIMUM L.

Biennial or perennial herbs, the pubescence of 2-parted, appressed, whitish or
grayish hairs; leaves narrow, basal and cauline, entire or dentate; flowers often
large and showy, yellow or dark red, the racemes not bracteate; sepals erect, equal
or the lateral ones gibbous at the base; stamens free, the filaments not dentate;
silique long and narrow, compressed, tetragonous, or subterete, the valves linear,
usually carinate, 1-nerved; style short or elongate, the stigma 2-lobate, capitate,
or emarginate; seeds 1-seriate, oblong, sometimes marginate.

About 90 species, in both hemispheres, chiefly in temperate
regions. Only one has been found in Central America, and in the
western hemisphere the genus finds its southern limit in western
Guatemala.

Erysimum Ghiesbreghtii Donn. Smith, Bot. Gaz. 57: 415.
1914.

Open, often rocky hillsides, 3,000-3,750 meters; Huehuetenango
(Sierra de los Cuchumatanes) ; Quezaltenango (?; between San
Marcos and Ostuncalco, perhaps in San Marcos). Type from
Chiapas.

An erect perennial from a thick, somewhat ligneous caudex, the stems often
several, erect, simple, 30-70 cm. high, leafy, thinly strigose; radical leaves numer-
ous, 5-11 cm. long, 4-7 mm. wide, oblanceolate-linear, acute, long-attenuate to
the base, green, very sparsely strigose; uppermost leaves much shorter, almost
linear; racemes 25 cm. long or shorter, few-many-flowered, the pedicels 5-8 mm.
long; sepals linear-lanceolate, 10-12 mm. long, dull red; petals 16-20 mm. long,
deep red; pods 3.5 cm. long, somewhat tetragonous, very slender, the seeds 12-15,
not emarginate.

Iberis amara L., candytuft, is cultivated rather frequently in
Guatemalan gardens, and is called "llovizna." It is often seen in the
parks, as in the plaza of Huehuetenango, but is rarely grown outside
the higher regions, 1,500 meters or more. It is an erect annual

370 FIELDIANA: BOTANY, VOLUME 24

15-30 cm. high, the narrowly lanceolate leaves dentate toward the
apex; flowers rather large, white, the racemes short, broad, and dense,
but in fruit much elongate. The fruits are compressed, almost as
broad as long, deeply lobate at the apex, the lobes acute; seeds 1 in
each cell, not marginate. The plant is a native of Europe, but is
grown commonly for ornament in most temperate regions.

LAMPROPHRAGMA 0. E. Schulz

Plants perennial or perhaps rather biennial, erect, pubescent below with simple
hairs, glabrous above, the stems solitary or several, simple or much branched;
leaves very narrow, the lower ones pubescent with furcate hairs, entire or repand-
dentate; racemes elongate, the flowers remote, purplish, nutant; sepals suberect,
the inner ones broader, subsaccate at the base; petals little exceeding the calyx;
stamens 6, the filaments linear, the anthers oblong, obtuse; ovary sessile, with very
numerous ovules, the style slender, evident, the stigma depressed; silique linear,
compressed, the valves obscurely 3-nerved; seeds 2-seriate, minute, ellipsoid,
compressed.

The genus consists of a single species, of uncertain status.
Schulz has split the genera of this family often into very small
groups, and it remains to be decided how many of his proposed new
ones are really worthy of recognition. This one is referable to
Thelypodium in its broad sense, and if that is divided, as seems to be
the tendency at present, this could probably be left in Heterothrix.

Lamprophragma longifolium (Benth.) 0. E. Schulz, Pflan-
zenreich IV. 105: 299. /. 63. 1924. Streptanthus longifolius Benth.
PI. Hartweg. 10. 1839. Thelypodium longifolium Wats. Bot. King
Exped. 25. 1871. Heterothrix longifolia Rydb. Bull. Torrey Club 34:
8. 1907.

Probably in open places in forest, 2,300 meters or more; Sacate-
pe*quez (Volcan de Agua) ; Chimaltenango (Calderas). Southwestern
United States; Mexico.

Plants slender, mostly 30-50 cm. high or more, the stems often purplish below;
lowest leaves oblanceolate or linear-oblanceolate, acute, attenuate to the base and
petiolate, irregularly repand-dentate, the upper leaves linear, mostly entire, sessile,
sparsely hispidulous or glabrous; racemes usually many-flowered, the pedicels
about 5 mm. long; sepals 3.5-5 mm. long, glabrous or nearly so, purplish or green;
petals 5-6 mm. long, purple or purplish; pods pendulous at maturity, 4-9 cm. long,
1-1.5 mm. wide, sessile.

LEPIDIUM L. Peppergrass

References: A. Thellung, Die Gattung Lepidium (L.) R. Br., eine
monographische Studie, Mitt. Bot. Mus. Univ. Zurich 28: 1-340.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 371

1906; C. Leo Hitchcock, The genus Lepidium in the United States,
Madrono 3: 265-320. 1936.

Annuals or perennials, herbaceous or rarely somewhat suffrutescent, glabrous
or with pubescence of simple hairs; leaves entire, dentate, or pinnatifid, petiolate
or amplexicaul; flowers in short or elongate racemes, small, white or yellow; sepals
usually pubescent dorsally; petals none or well developed; stamens 2, 4, or 6;
siliques usually more or less rounded, very small, obcompressed and flat, reticulate-
veined or smooth, glabrous or pubescent, retuse, sometimes winged; style none or
well developed; seeds 2.

Species about 120, widely distributed in temperate and warm
regions, the species usually few in the tropics. One other Central
American species has been described from Costa Rica.

Cauline leaves pinnatifid or bipinnatifid L. oblongum.

Cauline leaves merely serrate, or only the lowest pinnatifid.

Stems and leaves densely hispidulous; pedicels compressed; surfaces of the fruit

scarcely reticulate- veined L. lasiocarpum.

Stems and leaves sparsely puberulent or almost or quite glabrous; pedicels not
compressed; surfaces of the fruit delicately reticulate- veined.

L. virginicum.

Lepidium lasiocarpum Nutt. ex Torr. & Gray, Fl. N. Amer. 1:
115. 1838.

Roadside meadows, about 1,950 meters; Huehuetenango (moun-
tains west of Aguacatan, on road to Huehuetenango, Standley
81303). Western and southern United States; Mexico.

Plants annual or biennial, 35 cm. high or less, procumbent or erect, rather
densely hispidulous almost throughout, often much branched; basal leaves rather
large and twice-pinnatifid, the cauline leaves entire, serrate, or the lower ones
pinnatifid; flowers in elongate racemes 3-8 cm. long, the pedicels compressed,
1.5-5 mm. long, ascending or spreading; sepals 1 mm. long; petals equaling the
sepals or absent; stamens 2 or 4; pods oval, elliptic, or rounded, 3-4.5 mm. long,
ciliate in Guatemalan material, the surfaces glabrous, the valves delicately but
evidently reticulate-veined, deeply emarginate, the style very short.

The single Guatemalan collection apparently is referable to var.
typicum. It is rather probable that the plant has been introduced
into Guatemala from Mexico, although this is pure speculation.

Lepidium oblongum Small, Fl. Southeast. U. S. 468, 1331.
1903. Sacabe (Huehuetenango).

A weed in gardens, streets, or waste ground, sometimes on dry
or moist, rocky hillsides, 1,300-3,900 meters; Jalapa; Sacatepe"quez;
Chimaltenango; Huehuetenango; Quezaltenango; San Marcos.
Southern and western United States; Mexico.

372 FIELDIANA: BOTANY, VOLUME 24

Plants annual, erect or procumbent, usually much branched, the stems 20 cm.
long or less, the plants sparsely or often rather densely hispidulous or puberulent;
leaves almost all bipinnatifid or laciniate-lobate, the divisions linear, 2-3 mm. wide
or less, often glabrate; racemes short or elongate, the pedicels scarcely as long as
the fruits, erect or somewhat spreading, compressed; sepals scarcely 1 mm. long,
often purplish, pubescent, often but not always more or less persistent; stamens 2;
pods ovate or ovate-rounded, 2.2-3 mm. long, reticulate-veined or almost smooth,
glabrous but often cilate, shallowly emarginate, the style very short.

This may well be an introduced plant in Guatemala. Hitchcock
believes it to be introduced in North America, probably from South
America, but the Guatemalan and Mexican material is unlike the
South American L. bipinnatifidum Desv., with which it has been
confused. It has been reported incorrectly from Guatemala under
the name L. reticulatum Howell. The plant has been collected on
the very summit of Volcan de Santa Maria, Quezaltenango, which
it, like one or two other plants, probably has reached through human
agency or domestic animals. This species is used variously in house-
hold medicine in Huehuetenango.

Lepidium virginicum L. Sp. PI. 645. 1753. (?)L. Gerloffianum
Vatke ex Thell. Mitt. Bot. Mus. Univ. Zurich 28: 259. 1906 (based
in part on material from Guatemala). Jilipliegue; Mastuerzo;
Lentejuela; Antejuela; Lentejuelilla; Lentejilla; Antejuelilla; Sacabe
(Huehuetenango) .

Open or shaded places, often a weed in waste or cultivated
ground, on open banks, roadsides, moist or dry fields, sometimes on
limestone, 2,450 meters or less; Pete"n; Alta Verapaz; Baja Verapaz;
El Progreso; Jalapa; Jutiapa; Santa Rosa; Escuintla; Guatemala;
Sacatepe"quez; Chimaltenango; Solola; Huehuetenango; Quezalte-
nango. Widely distributed in North America; Mexico; British
Honduras to Costa Rica; South America.

An annual, usually erect and 60 cm. high or less, often much branched, sparsely
pubescent or hirtellous; basal leaves often forming rosettes, pinnatifid or twice
pinnatifid, the lower cauline leaves often pinnatifid, the middle and upper ones
serrate or almost entire; racemes numerous and many-flowered, often much elon-
gate, the slender pedicels terete, erect or spreading, usually somewhat longer than
the fruits; sepals glabrous or slightly pubescent, 1 mm. long; petals white, equaling
the sepals or longer, rarely minute; stamens usually 2; pods glabrous, rounded-
elliptic to almost orbicular, 2.5-4 mm. long, shallowly emarginate, the style
obsolete.

In Salvador sometimes called "mastuerce," "culantrillo," and
"cupapayo"; "putxiu," "putcan" (Yucatan, Maya). This has been
reported from Guatemala as L. lasiocarpum var. tenuipes Watson,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 373

but all Guatemalan material of this alliance probably is referable to
L. virginicum. It is a common weed here, as in many parts of the
United States. The foliage has an acrid but agreeable flavor, and
in the United States it is sometimes cooked and eaten as a pot herb.
We have no information to the effect that the plant is ever eaten
in Central America. A sweetened decoction of it sometimes is
administered to babies suffering from colic, and it is a domestic
remedy for affections of the stomach and intestines. Dieseldorff
states that in Alta Verapaz it is employed for treating inflammation
of the eyes and mouth. In Guatemala the plant behaves as an
introduced weed, but the same may be said of many of the weedy
plants that almost certainly are native in the region. Birds are
fond of the seeds of peppergrass, and in the United States bunches
of the branches with ripe pods often are given to caged birds,
particularly canaries.

LOBULARIA Desvaux. Sweet alyssum

Annual or perennial herbs or low shrubs, usually strigose with pale furcate
hairs; leaves narrow, entire, the stems leafy; flowers small, white, in terminal,
mostly many-flowered racemes, often fragrant; sepals strigose; petals obovate,
entire; filaments slender, not dentate, with 2 small glands at the base; silique
compressed, oval or orbicular, with 1 seed in each cell; seeds marginate;
cotyledons accumbent.

About 4 species, native in the Mediterranean region.

Lobularia maritima (L.) Desv. Journ. Bot. 3: 169. 1814.
Clypeola maritima L. Sp. PL 652. 1753. Alyssum maritimum Lam.
Encycl. 1: 98. 1783. Koniga maritima R. Br. in Denh. & Clapp,
Narr. Exp. Afr. 214. 1826. Llovizna.

Grown commonly for ornament in gardens and parks of Guate-
mala, mostly at middle or rather high elevations, sometimes in the
lowlands. Native of southern Europe.

Plants ascending or procumbent, often forming dense masses of stems, these
usually 30 cm. long or less, the whole plant whitish-strigose; cauline leaves sessile
or nearly so, lanceolate or linear, 1-5 cm. long, acute, attenuate to the base, the
lowest leaves oblanceolate, petiolate; flowers fragrant, 4 mm. broad, the pedicels
ascending, 6-8 mm. long; fruit glabrous, pointed, oval or almost orbicular, 3 mm.
long.

In Salvador sometimes called "no-me-olvides." This is a com-
mon garden flower in both Central America and the United States,
much used as a border for flower beds.

374 FIELDIANA: BOTANY, VOLUME 24

MATTHIOLA R. Brown. Stock

Coarse herbs or shrubs, stellate-tomentose, the pubescence often whitish,
branched, with leafy stems; leaves narrow, entire or sinuate; flowers large, race-
mose, not bracteate, mostly purple or pink; sepals erect, the inner ones saccate
at the base; petals long-unguiculate; siliques long and slender, usually large, terete
or compressed, the septum thick, minutely areolate; stigma lobes erect, connivent,
often thickened or horn-like; seeds 1-seriate, compressed, often marginate.

Species 30 or more, in Europe, Asia, and Africa.

Matthiola incana (L.) R. Br. in Ait. Hort. Kew. ed. 2. 4: 119.
1812. Cheiranthus incanus L. Sp. PI. 662. 1753. Aleli; Alelia.

Cultivated commonly for ornament in the central and western
mountains, especially in the highlands of the Occidente; sometimes
more or less naturalized in the vicinity of dwellings. Native of the
Mediterranean region but cultivated commonly for ornament in
temperate regions.

A coarse biennial or perennial, often becoming woody below and forming
dense bushes a meter high or more, often much branched, finely and closely
stellate-tomentose throughout, the foliage whitish; leaves linear-oblanceolate or
oblong-oblanceolate, obtuse, long-attenuate at the base into a petiole, or sessile,
entire or coarsely and remotely undulate-dentate; flowers white to dark purple,
often double, in short or elongate, dense or lax racemes; sepals narrow, obtuse,
12 mm. long, stellate-tomentose; petals about 2.5 cm. long, long-unguiculate;
pods mostly 6-7.5 cm. long and 4 mm. thick, stellate-tomentose, on stout, erect
or strongly ascending pedicels.

Stocks are common garden plants in most parts of the United
States, but they are even more popular in Guatemala because they
withstand neglect, thrive in the coldest regions, and bloom through-
out the year. They are in flower in Los Altos when most other garden
plants have been killed by the cold. Large bushes often are seen
about the humblest dwellings of the uplands, where they look most
unhappy, trampled by the larger animals and used as perches by
the chickens. The plants apparently endure for many years in this
climate.

NASTURTIUM R. Brown. Watercress

Glabrous aquatic perennial herbs with simple or branched stems; leaves pin-
nately divided, with pungent flavor; flowers small, white, in terminal racemes;
siliques linear or oblong-linear, long-pedicellate, the style short and stout, the
valves convex, nerveless; seeds 2-seriate, the cotyledons accumbent.

The genus consists of a single species. It is questionable whether
this should be treated as a genus distinct from Rorippa, since authors

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 375

vary greatly in their disposition of these plants and also in the specific
name to be used for watercress. It is little to the credit of systematic
botanists, who boast of the defmiteness of Latin names, that they
disagree as to the proper Latin name of watercress, while any child
knows to what the English name "watercress" refers. It is fortunate
indeed that vegetable dealers do not sell vegetables by their Latin
names, else our markets would be in a state of perpetual chaos.

Nasturtium officinale R. Br. in Ait. Hort. Kew. ed. 2. 4: 110.
1812. Sisymbrium Nasturtium-aquaticum L. Sp. PI. 567. 1753.
Rorippa Nasturtium Beck, Fl. Nied. Oesterr. 2: 463. 1892. Radicula
Nasturtium-aquaticum Britt. & Rendle, Brit. Seed PI. 3. 1907.
Berros;Guixocul, Rechsut (Quiche").

In small, cool or cold streams or in marshes, rarely growing on
mud where water has receded, 500-3,300 meters; Alta Verapaz;
Baja Verapaz; Escuintla; Guatemala; Sacatepe"quez; Quiche";
Totonicapan; Huehuetenango; Quezaltenango; San Marcos. Native
of Europe and Asia, widely cultivated and naturalized in most tem-
perate regions; abundantly naturalized in the United States and
Mexico, and in the mountains of Central America.

Plants often much branched and forming dense colonies over the surface of
water, the stems succulent, rooting at the nodes; leaf segments 3-9, the terminal
one larger than the others, ovate, oval, or orbicular, obtuse or rounded at the apex,
more or less undulate or somewhat crenate; racemes short at first, elongate in
fruit, the flowers 4-5 mm. broad; petals twice as long as the sepals; pods 10-30
mm. long, 1 mm. thick or more, spreading and slightly curved upward, the pedicels
about as long as the pods.

Watercress is a popular salad plant in Guatemala and is sold
commonly and often in large amounts in the markets. It is a plant
that is dangerous to eat in the tropics, at least in many regions, and
one that should be avoided in its raw state by visitors. One never
knows in what kind of water it has been grown. Foreigners and some
local people treat it, like lettuce, with a weak iodine solution in order
to kill possible noxious bacteria or other organisms, later rinsing the
plants thoroughly with water, but there always persists a slight taste
of iodine that does not improve the natural flavor of watercress.
In Guatemala, watercress sometimes is cooked like broccoli, and then
is a very good vegetable, tender and of excellent flavor. We have
never seen it cooked in the United States, where it is used extensively
as a salad plant and in some regions is cultivated for market on a
large scale. Watercress grows with great luxuriance in the regions of
Almolonga and Zunil in Quezaltenango, and there are large colonies

376 FIELDIANA: BOTANY, VOLUME 24

in the cold, swift water of the great spring at Aguacatan, Huehuete-
nango. Some of the watercress in the Quezaltenango market is very
large and robust, larger than any we have seen in the United States.
It is said that the plant is much eaten by cattle. It gives its name to
the settlement of El Berro, a caserio in San Marcos.

RAPHANUS L.

Annual or biennial herbs, erect, usually branched; stems leafy, the leaves
lyrate-pinnatifid; flowers rather large and showy, in mostly elongate racemes,
pink, purple, or yellow; pubescence none or of simple hairs; sepals erect, the inner
ones subsaccate at the base; petals unguiculate; filaments not dentate; silique
elongate, terete, continuous or moniliform, smooth or costate, coriaceous or corky,
continuous within or constricted into several cells; style slender, the stigma
emarginate; seeds pendulous, globose, the cotyledons conduplicate, sometimes
complicate.

About 6 species, in Europe and Asia.

Flowers yellow, fading white or purplish; silique longitudinally costate, 4-10-

seeded R. Raphanistrum.

Flowers pink, purple, or white; pods smooth 2-3-seeded R. sativus.

Raphanus Raphanistrum L. Sp. PI. 669. 1753.

A weed in cultivated or waste ground, about 1,800 meters;
Chimaltenango (flax field near Patzicia) ; Quiche". Native of Europe
and Asia; occasionally naturalized in temperate North America, and
sometimes becoming a troublesome weed.

Annual or biennial, erect or ascending from a slender root, a meter high or
less, sparsely hispidulous, especially below; basal and lower leaves deeply lyrate-
pinnatifid, 10-20 cm. long, with a large terminal lobe and 4-6 pairs of smaller
lateral ones, the segments crenate or dentate; upper leaves small, oblong; flowers
12-15 mm. broad, yellow or sometimes purplish, fading to white with purple
veins; pedicels 6-15 mm. long; pods 2.5-3.5 cm. long, 6-10-seeded, cylindric,
constricted between the seeds, longitudinally multicostate, the slender beak 1-2
cm. long.

Raphanus sativus L. Sp. PI. 669. 1753. Rabano. Radish.

Native of Asia, but cultivated in almost all parts of the earth
for its edible roots; grown generally in Guatemala at almost all
elevations, especially in the mountains; sometimes found as a weed
in waste or cultivated ground.

A tall annual, the roots usually much enlarged, but in wild plants slender;
similar in most respects to the preceding species, but the large flowers white, pink,
or purple; pods fleshy or spongy, smooth, ovoid-oblong or lance-oblong, 2-3-seeded,
pale green, the long conic beak often as long as the body of the fruit.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 377

One of the common garden vegetables of Guatemala and all
Central America, thriving particularly well at middle and high
elevations. While most of the radishes grown in Guatemala are of
the small globe variety most popular in the North, large ones, either
red or white, often are seen on sale in the markets. The roots are
eaten mostly sliced in salads, dressed with oil and vinegar.

ROMANSGHULZIA 0. E. Schulz

Reference: Reed C. Rollins, A tentative revision of the genus
Romanschulzia, Contr. Dudley Herb. 3: 216-226. 1942.

Annual, biennial, or perennial herbs, the pubescence none or of simple hairs,
the stems usually branched; leaves mostly cauline, sessile, auriculate, sagittate;
racemes terminal, often much elongate, the flowers usually numerous, small;
sepals early deciduous, not saccate at the base; petals narrow, spreading; filaments
dilated at the base, usually with well-developed nectar glands; siliques terete or
slightly compressed, erect, spreading, or deflexed, stipitate or subsessile; seeds not
winged, 1-2-seriate.

Eight species, in Mexico and Central America. Two other
Central American ones are known, in Costa Rica and Panama.

Petals white; sepals greenish; siliques 1-2 cm. long R. arabiformis.

Petals purple; sepals purple; siliques 1.8-5 cm. long.

Siliques 1.8-3 cm. long, spreading; pedicels spreading; petals and sepals sub-
equal R. guatemalensis.

Siliques 4.8-5 cm. long, strongly ascending; pedicels ascending in fructification;
petals conspicuously longer than the sepals R. alpina.

Romanschulzia alpina Standl. & Steyerm., sp. nov.

Grassy alpine slopes, 3,300-3,700 meters; Huehuetenango
(Sierra de los Cuchumatanes; type collected between Tojquia and
Caxin bluff, Steyermark 50144 in Herb. Chicago Nat. Hist. Mus.;
also near Tunima, Steyermark 48293).

An erect herb, probably annual, the stems simple or sparsely branched,
glabrous, 15-70 cm. high; cauline leaves remote, glaucous, lanceolate, 2-7 cm.
long, 5-11 mm. wide, acuminate, auriculate-clasping or sagittate at the base,
glabrous; racemes terminal, ebracteate, remotely few-flowered, 25 cm. long or
shorter, the pedicels at anthesis 4-5 mm. long, in fruit 5-10 mm. long, ascending;
flowers 5.5-6 mm. long, the sepals lilac, scarious-marginate, oblong or elliptic-
oblong, obtuse or rounded at the apex, 3.2-3.5 mm. long; petals purplish, sub-
linguiform, abruptly contracted at the base, rounded at the apex, 5.5-6 mm. long;
filaments ampliate at the base, glabrous, 3 mm. long; young siliques ascending,
about 5 cm. long, 1.5 mm. wide, sessile; seeds 1-seriate, rufescent, 1.2 mm. long,
reticulate-striatulate.

Herba erecta glabra, simplex vel sparse ramosa; folia caulina remota glauca
lanceolata acuminata, basi auriculato-amplexicaulia vel sagittata; racemi ebrac-

378 FIELDIANA: BOTANY, VOLUME 24

teati ramote pauciflori usque ad 25 cm. longi, pedicellis ad anthesin 4-5 mm. longis,
in statu fructifero 5-10 mm., adscendentibus; flores 5-6 mm. longi, sepalis lilacinis
scarioso-marginatis, oblongis vel elliptico-oblongis, 3.2-3.5 mm. longis; petala
purpurascentia sublinguiformia; filamenta basi dilatata glabra; siliquae adscen-
dentes ca. 5 cm. longae, 1.5 mm. latae, sessiles; semina uniseriata.

The distinguishing characters of this species are its uniseriate,
strongly ascending pods and fruiting pedicels, the sublinguiform
petals much longer than the sepals, and the completely glabrous
stems and leaves. The siliques are much longer than is usual in
other species of Romanschulzia. Although the calyx seems not to
fall at anthesis as is normal in that genus, the ampliate bases of the
filaments and the general habit of the plant would seem to ally it
more closely with Romanschulzia than with Thelypodium or other
genera rather dubiously separable from it.

Romanschulzia arabiformis (DC.) Rollins, Contr. Dudley
Herb. 3: 221. 1942. Nasturtium arabiforme DC. Syst. 2: 200. 1821.

At 3,300 meters; Quezaltenango (Volcan de Santa Maria, A. F.
Skutch 866). Central Mexico.

Plants probably annual, erect, 2 meters high or less, branched, glabrous or
sometimes hirsute near the base; leaves sessile, auriculate and sessile at the base,
narrowly lanceolate or oblong-lanceolate, obtuse or acute, entire or remotely
denticulate, 10-20 cm. long, green above, glaucous beneath; racemes much elon-
gate; sepals glabrous, greenish, 2.5-3.5 mm. long; petals linear, white, 3-4 mm.
long; pedicels spreading or ascending, 6-10 mm. long; pods terete, usually obtuse
at each end, glabrous, stipitate or almost sessile, 1-2 cm. long; style less than 1 mm.
long.

Rollins believes that the Skutch collection cited may represent
an undescribed species, but the material, unfortunately, is not in good
enough condition for description.

Romanschulzia guatemalensis (Standl.) Rollins, Contr.
Dudley Herb. 3: 223. 1942. Sisymbrium guatemalense Standl. Journ.
Wash. Acad. Sci. 17: 251. 1927. R. Loeseneri 0. E. Schulz, Bot.
Jahrb. 66: 101. 1933 (type from Todos Santos, Huehuetenango, C.
& E. Seler 3110).

Open places in forest or in moist or wet thickets, 2,500-3,700
meters; endemic; Chimaltenango; Quich£ (type from San Miguel
Uspantan, -Heyde & Lux 3079); Huehuetenango; Quezaltenango
(Volcan de Zunil).

An erect annual 1-1.5 meters high, or often lower, the stems branched,
glabrous; leaves sessile and amplexicaul, oblong to broadly lanceolate, irregularly
denticulate, glabrous, the auricles rounded; inflorescence large, the racemes much

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 379

elongate, rather dense; sepals purple, oblong, 3.5-4.5 mm. long, early deciduous;
petals purple, linear, 3.5-5 mm. long; pedicels slender, spreading, 10-15 mm. long;
siliques terete, straight or slightly curved, somewhat moniliform, 1.8-3 cm. long,
short-stipitate or subsessile.

RORIPPA Scopoli

Annual or perennial herbs, glabrous or with pubescence of simple hairs;
leaves simple or pinnately lobate or dissected; flowers small, yellow, in short or
elongate racemes; sepals spreading; stamens 1-6; siliques short, terete or nearly
so, not stipitate, the valves nerveless or 1-nerved ; style short or slender and some-
what elongate, the stigma 2-lobate or subentire; seeds turgid, minute, 2-seriate or
rarely 1-seriate; cotyledons accumbent.

About 50 species, chiefly in the north temperate zone. No others
are known in Central America.

Petals none; cauline leaves mostly simple; fruit 2-3 cm. long R. indica.

Petals present; cauline leaves all pinnately parted; fruit 1.5 cm. long or usually
shorter R. mexicana.

Rorippa indica (L.) Hochr. Candollea 2: 370. 1923. Sisymbrium
indicum L. Mant. PI. 1: 93. 1767. Nasturtium indicum DC. Reg.
Veg. Syst. 2: 199. 1821.

Waste ground, sometimes a weed in streets; Alta Verapaz
(Coban) ; Guatemala. Native of Asia and Africa, rarely naturalized
in tropical America; found also in Costa Rica.

Plants perennial, erect or ascending, branched, glabrous, generally 20 cm.
high or less; cauline leaves ovate to oblong, simple, the lower ones with a few small
lobes at the base, acute or obtuse, irregularly dentate, the uppermost sessile, the
lower petiolate; flowers greenish yellow, scarcely 2 mm. long, in short or elongate
racemes, the pedicels much shorter than the pods; pods 2-3 cm. long, very slender,
spreading, the style very short and thick.

Rorippa mexicana (Moc. & Sess^) Standl. & Steyerm. Field
Mus. Bot. 23: 54. 1944. Nasturtium mexicanum Moc. & Sess6 ex
DC. Reg. Veg. Syst. 2: 193. 1821.

Moist or wet pastures or cultivated fields, often a weed in gardens,
streets, or waste ground, sometimes in sand or gravel along streams,
1,000-2,550 meters; Alta Verapaz; Baja Verapaz; Jalapa; Santa
Rosa; Sacatepe"quez; Chimaltenango; Solola; Quezaltenango. Mex-
ico; Honduras; Costa Rica.

Plants annual or perhaps also perennial, erect or decumbent, branched, the
stems 30 cm. long or less, glabrous, usually very leafy; leaves pinnately parted or
twice pinnatifid, the segments usually numerous, narrow or broad, obtuse, usually
sinuate-lobate; flowers yellow or greenish yellow, 1.5-2 mm. long, the racemes

380 FIELDIANA: BOTANY, VOLUME 24

short or often much elongate; pods 10-15 mm. long, 2 mm. thick, straight or
somewhat curved, often somewhat torulose; pedicels about half as long as the pods.

In Costa Rica sometimes called "platanillo," because the fruit-
ing racemes suggest stems of bananas. The plant has been reported
from Guatemala as Nasturtium palustre DC. var. brevipes DC. It
is decidedly weedy, and is most often seen in waste ground about
dwellings.

TOVARIACEAE

Large herbs, perhaps sometimes suffrutescent, strong-scented, glabrous,
branched; leaves alternate, without stipules, 3-foliolate, the leaflets membrana-
ceous, entire; flowers perfect, regular, in long terminal racemes; sepals 8, lance-
subulate, imbricate, deciduous; petals 8, oblong-lanceolate, sessile; torus very
short; stamens 8, free, the filaments pilose at the base; ovary subglobose, on a very
short stipe, 6-8-celled, the septa membranaceous; ovules very numerous, the
placentae spongious, axillary, binate in each cell, the stigma sessile, 8-radiate;
fruit small, baccate, globose, the pericarp membranaceous; seeds very numerous,
minute, the testa crustaceous, granulate; embryo curved; endosperm present.

The family consists of a single genus, with the characters given
above, both species natives of tropical America. Only one occurs in
Central America.

TOVARIA Ruiz & Pavon

Tovaria diffusa (Macfad.) Fawc. & Rendle, Fl. Jam. 3, pt. 1:
246. 1914. Bancroftia diffusa Macfad. Fl. Jam. 1: 112. 1837.

Dense wet thickets along streams in the mountains, 1,500-2,800
meters; Alta Verapaz (mountains east of Tactic); Suchitepe'quez ;
Quich^ ; San Marcos (volcanoes of Tacana and Tajumulco) . Southern
Mexico; Costa Rica; Jamaica; Colombia and Venezuela to Peru.

Plants erect or weak and somewhat reclining, about 1.5 meters tall, almost
wholly glabrous; leaves on very long, slender petioles, the 3 leaflets lanceolate,
long-acuminate, 6-15 cm. long, entire, paler beneath; flowers pale green or rather
bright yellow, long-pedicellate, in very long, lax racemes; petals about 7 mm. long;
fruit about 1 cm. in diameter.

The plant seems to be rare and local in Guatemala, but it is more
plentiful in Costa Rica.

GAPPARIDACEAE. Caper Family

Herbs, shrubs, or trees, glabrous or pubescent, sometimes glandular or lepi-
dote, the stems and branches terete, the sap watery; leaves alternate or rarely
opposite, with or without stipules, simple or palmately 1-5-foliolate, the leaves

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 381

or leaflets entire, rarely serrate or lobate; stipules, when present, setaceous,
herbaceous, or spinescent; flowers mostly perfect, fasciculate or solitary, or ter-
minal and corymbose or racemose, regular or irregular, often showy, the pedicels
ebracteate, or sometimes bracteate at the base; sepals 4-8, free or connate, 1-2-
seriate, subequal or the anterior one larger, the 2 innermost sometimes much
smaller, imbricate or valvate; petals 4, rarely none, sessile or unguiculate, imbricate
or open in bud, very rarely valvate; torus short or elongate, symmetric or asym-
metric, sometimes disk-like, often appendaged, depressed or attenuate into a long
or short stipe; stamens inserted at the base or apex of the torus, suberect, spreading,
or declinate, few or many, equal or unequal, all fertile or some of them without
anthers; filaments usually filiform and free, sometimes connate with the torus,
inflexed or contorted in bud; anthers oblong, dorsifixed near the base; ovary sessile
or stipitate, usually ovoid, 1-celled, sometimes with false septa; style usually short
or none, the styles sometimes 3 and sessile, the stigma usually orbicular and sessile;
ovules numerous, anatropous, affixed to the parietal placentae in 1-many series,
rarely solitary; fruit capsular or baccate, rarely drupaceous, when capsular usually
siliquiform, elongate, compressed, and many-seeded; seeds adnate to the placentae
or septa, reniform, with coriaceous testa, and roughened in the genera with capsular
fruit, angulate or reniform in the baccate fruits and often surrounded by pulp;
endosperm none or scant; embryo arcuate or incurved, the cotyledons incumbent,
plicate, convolute, or induplicate, rarely flat.

About 40 genera, widely distributed, chiefly in tropical regions.
No other genera are known in Central America.

Fruit capsular; plants herbaceous.

Filaments united below with the gynophore, this in fruit bearing a scar left by
the deciduous free filaments a short distance above the base of the gyno-
phore; leaflets 5-9 Gynandropsis.

Filaments free.

Stamens 4 or 6; petals not yellow Cleome.

Stamens 8-many ; petals yellow '..... Polanisia.

Fruit baccate or drupaceous; trees or shrubs.
Leaves, at least most of them, 3-foliolate.

Petals 4; fruit long-stipitate Crataeva.

Petals none; fruit sessile Forchhammeria.

Leaves simple.

Sepals distinct or nearly so Capparis.

Sepals connate almost to the apex.

Calyx tubular-campanulate, scarlet; leaves not peltate, thin. . . Steriphoma.

Calyx campanulate, not scarlet; leaves mostly peltate just above the base,
thick-coriaceous. . . .Morisonia.

CAPPARIS L.

Trees or shrubs, sometimes armed with prickles or spines, glabrous, lepidote,
or tomentose; leaves simple, petiolate, coriaceous to membranaceous, the stipules
spinose or subulate; inflorescence various in form, the flowers mostly white and
bracteate; sepals normally 4, free or connate only at the base, rarely united in bud

382 FIELDIANA: BOTANY, VOLUME 24

and irregularly rupturing, naked within or glandular or sometimes ligulate, val-
vate, imbricate, or open in bud; petals 4, imbricate; torus short; stamens usually
numerous, the filaments filiform, free; ovary long-stipitate, 1-4-celled, the pla-
centae 2-6, the ovules numerous; stigma sessile; fruit baccate, stipitate, globose
to cylindric, often much elongate, rarely dehiscent; seeds numerous, imbedded in
pulp, the testa crustaceous or coriaceous; embryo convolute.

Species about 150, in both hemispheres, mostly in the tropics.
A few additional ones occur in southern Central America. Best-
known member of the genus is the caper (alcaparro] of the Mediter-
ranean region, whose product is not unknown in Guatemala. Capers
are the flower buds and young fruits of Capparis spinosa L., preserved
with salt and vinegar.

Indument of scales or of branched hairs.

Lower surface of the leaves covered with stellate hairs.

Branches stellate-pilose with coarse brownish hairs; leaf blades mostly 7-10

cm. wide, long-acuminate C. Steyermarkii.

Branches minutely stellate-pubescent with gray hairs; leaf blades mostly 2-3

cm. wide, acute C. incana.

Lower surface of the leaves lepidote.

Sepals valvate in bud C. cynophallophora.

Sepals open in bud.

Calyx disk-like, the sepals short, triangular, spreading C. Lundellii.

Calyx deeply lobate, the sepals narrow, erect.

Petioles mostly 1 cm. long or shorter; leaf blades chiefly 1.5-3 cm. wide.

C. indica.
Petioles mostly 1.5-3.5 cm. long; leaf blades chiefly 5.5-7 cm. wide.

C. calciphila.
Indument none or of simple hairs.

Stamens 6; plants glabrous.

Leaves long-acuminate C. Heydeana.

Leaves obtuse or merely acute C. hexandra.

Stamens numerous.

Petioles very unequal, most of them elongate, the uppermost leaves sessile
or on very short petioles, the blades large, mostly 15-30 cm. long, coria-
ceous, chiefly oblong-lanceolate C. Baducca.

Petioles subequal, those of the lower leaves not much longer than those of the
upper leaves.

Leaf blades emarginate or subcordate at the base C. verrucosa.

Leaf blades acute or obtuse at the base.

Leaves emarginate, rounded or obtuse, rarely subacute, at the apex,
coriaceous C, flexuosa.

Leaves all or mostly acute or acuminate, membranaceous or chartaceous.
Sepals ovate or oblong-ovate, 3-3.5 mm. long; pedicels 5-6.5 cm. long.

C. Tuerckheimii.

Sepals broadly oval or orbicular, 5 mm. long; pedicels 2-4 cm. long.

C. quiriguensis.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 383

Capparis Baducca L. Sp. PI. 504. 1753. C. frondosa Jacq.
Enum. PL Carib. 25. 1760. C. stenophylla Standl. Journ. Wash.
Acad. Sci. 13: 437. 1923 (type from San Vicente, Salvador).

Moist or dry thickets or forest, 350 meters or less; Pete"n; Santa
Rosa; Suchitepe'quez ; Retalhuleu; San Marcos. Southern Mexico;
Salvador to Panama; West Indies; northern South America.

A shrub or small tree, 2-7.5 meters high, glabrous throughout; leaves mostly
crowded near the ends of the branches, the petioles very unequal, those of the
lower leaves long and slender, of the uppermost leaves very short or almost none;
leaf blades oblong-elliptic to linear-lanceolate, 10-30 cm. long or even larger,
subobtuse to long-attenuate, somewhat narrowed to the usually emarginate base,
coriaceous, paler beneath, the veins closely reticulate and somewhat prominent
on both surfaces; flowers racemose, white, the racemes short and few-flowered,
the flowers on short stout pedicels; sepals suborbicular, imbricate in bud; fruit
borne on a stipe 1-1.5 cm. long, subterete and somewhat torulose, oblong, 2-5 cm.
long, about 1 cm. thick, dark purple-red or purple-brown, smooth, the seeds large,
tuberculate.

Called "quita-calzon" in Salvador. In some regions the fruit
has the reputation of being poisonous. C. stenophylla has relatively
longer and narrower leaves than in typical forms of the species,
but it is probably no more than an extreme variant of C. Baducca.
The Maya names of Yucatan are reported as "xcabachuloc" and
"cabachulob."

Capparis calciphila Standl. & Steyerm. Field Mus. Bot. 23:
158. 1944.

Wet to dry forest or thickets, 1,300 meters or less; Alta Verapaz
(type collected along knife-edge of a limestone ridge, Cerro Chinaja,
between Finca Yalpemech and Chinaja, above source of Rio San
Diego, Steyermark 45616); Guatemala (Lago de Amatitlan; sterile
and determination uncertain). British Honduras (Jacinto Hills,
W. A. Schipp S-655).

A tree of 12 meters, the branches stout, densely brown-lepidote, the internodes
short; leaves firm-coriaceous, on stout petioles 1.5-2.5 or sometimes as much as
4 cm. long, elliptic or usually obovate-elliptic, 8.5-13 cm. long and 4.5-6.5 cm.
wide or sometimes larger, rounded or very obtuse and apiculate at the apex, obtuse
or broadly cuneate-obtuse at the base, glabrous and lustrous above, the nerves and
veins prominulous, laxly reticulate, yellowish brown beneath, rather densely
lepidote, not at all pilosulous, the costa slender, strongly elevated, the nerves and
veins prominent and laxly reticulate; inflorescences axillary, cymose, few-flowered,
the peduncles as much as 9 cm. long, the pedicels stout, 8-14 mm. long, densely
brown-lepidote; calyx 3.5 mm. long, lobate almost to the base, very densely brown-
lepidote, the lobes open in bud, narrowly triangular, acute, appressed; petals white
within, very densely stellate-tomentose outside, 1 cm. long in bud.

384 FIELDIANA: BOTANY, VOLUME 24

Capparis cynophallophora L. Sp. PI. 504. 1753. C. jamaicen-
sis Jacq. Enum. PL Carib. 23. 1760. Zic (Pete"n, Maya).

Moist or rather dry forest or thickets, 300 meters or less; Peten.
Southern Florida; Yucatan Peninsula of Mexico; British Honduras;
Costa Rica; Panama; West Indies; northern South America.

A shrub or small tree 2-7 meters high, the branchlets brownish-lepidote;
leaves short-petiolate, thick-coriaceous, elliptic or oblong-elliptic, 4-12 cm. long,
obtuse or acute, glabrous and lustrous above, the costa strongly impressed, densely
brown-lepidote beneath; racemes few-flowered, the flowers fragrant, white, on
stout pedicels; sepals valvate in bud, densely brown-lepidote, 8-11 mm. long;
petals 10-13 mm. long, lepidote on the outer surface; stamens numerous, 2-3
times as long as the sepals, purplish, with yellow anthers; fruit siliquiform, terete,
borne on a long gynophore, sometimes 30 cm. long, torulose, lepidote, in age
rupturing irregularly.

Capparis flexuosa L. Sp. PI. ed. 2. 722. 1762. Morisonia
flexuosa L. Amoen. Acad. 5: 398. 1760. Potal (Pete"n, fide Lundell).

Dry or moist thickets, usually on plains, 660 meters or less;
Pete"n; El Progreso; Zacapa; Chiquimula; reported from Escuintla;
Retalhuleu; Quiche". Mexico; Salvador; Panama; West Indies;
South America.

A glabrous shrub or small tree, commonly 4 meters high or less; leaves short-
petiolate, coriaceous, oblong to obovate, 3-6.5 cm. long, retuse to obtuse or sub-
acute at the apex, rounded to subacute at the base, with conspicuous lateral nerves;
flowers few at the ends of the branches, white, showy, fragrant; sepals imbricate,
rounded, 5-10 mm. long; petals 1.5 cm. long; stamens numerous, 3 times as long
as the petals; fruit siliquiform, torulose or continuous, 7-15 cm. long, 1-1.5 cm.
thick, in age more or less 2-valvate, the gynophore 4-9 cm. long; seeds "numerous,
2-seriate, imbedded in scarlet pulp.

The root has a flavor resembling that of horse-radish (Armoracia) .
The Maya name of Yucatan is variously recorded as "xbayunac,"
"xpayumac," and "xpayunac."

Capparis hexandra Blake, Proc. Biol. Soc. Wash. 33: 117. 1920.
Esquisuchil.

Known only from the type locality, Finca Capetillo, Antigua,
Sacatepe"quez, Wilson Popenoe 875; collected there also by Jorge
G. Salas, no. 1399; apparently in cultivation.

A small tree, glabrous throughout; petioles 1-2 cm. long; leaf blades obovate-
oblong, 5.5-10 cm. long, 2-3 cm. wide, rounded to subacute at the apex, somewhat
narrowed to the obtuse base; flowers solitary in the leaf axils, white, fragrant, the
pedicels 1.5-2 cm. long; sepals imbricate, rounded, about 1 cm. long; petals 3.5
cm. long, spatulate-oblanceolate; stamens 6, the filaments equaling the petals;
ovary borne on a long slender gynophore, 2-celled.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 385

Closely related to C. Heydei, and perhaps only a form of it
modified by cultivation. It is strange that this plant should be
called "esquisuchil," since that name is given commonly, especially
about Antigua, to Bourreria.

Capparis Heydeana Donn. Smith, Bot. Gaz. 18: 197. 1893;
20:2. pi. 1. 1895.

Known in Guatemala only from the type, Laguna de Ayarza,
Jalapa, 2,520 meters, Heyde & Lux 4112. Salvador; Costa Rica.

A tree of 9-13 meters, glabrous or nearly so; leaves slender-petiolate, obovate-
oblong to elliptic-oblong, 10-20 cm. long, acuminate or long-acuminate, acute at
the base; flowers in short few-flowered bracteate terminal racemes, the pedicels
longer than the petioles; sepals oblong, 1.5-2 cm. long; petals obovate-spatulate,
5-6.5 cm. long; stamens 6, about equaling the petals; gynophore greatly elongate;
ovary falsely 2-celled; fruit oblong, 2.5 cm. long, coarsely verrucose.

Called "polvora" in Salvador, because it is said the bark smells
like burnt gunpowder.

Capparis incana HBK. Nov. Gen. & Sp. 5: 94. 1821.
Dry thickets, 200-660 meters; Zacapa; Chiquimula. Southern
Mexico.

A shrub or small tree, the branchlets minutely stellate-tomentulose with gray
or rusty hairs; leaves petiolate, chiefly elliptic or ovate, 4-8 cm. long, usually
acute at each end, rather thin, gray-green and glabrous above, densely and
minutely stellate-tomentose beneath with gray tomentum; flowers small, white,
in small few-flowered umbelliform axillary racemes, slender-pedicellate; sepals
open in bud, linear, erect; petals about 5 mm. long, densely pubescent outside;
stamens few, slightly longer than the petals; gynophore equaling or shorter than
the fruit, this globose or oblong, 1-2.5 cm. long, densely stellate-tomentulose.

The Maya names of Yucatan are "bocanche" and "xcoche."
The fruit is reputed poisonous. The Guatemalan material, all from
sterile and mostly small bushes, shows extraordinary variation in the
leaves. On vigorous sterile branches these are sometimes linear,
with a small divaricate lobe on each side near the base. The suc-
ceeding leaves have a broad obcuneate basal portion and a short or
long, linear terminal one, while the ultimate leaves on the branches
are ovate or lance-ovate.

Capparis indica (L.) Fa we. & Rendle, Journ. Bot. 52: 144.
1914. Breynia indica L. Sp. PI. 503. 1753. C. amygdalina Lam.
Encycl. 1: 608. 1785. Fruto de garza (fide Aguilar).

Dry forest or thickets, 250-1,300 meters; El Progreso; Zacapa;
Guatemala; Quiche"; Huehuetenango. Western and southern

386 FIELDIANA: BOTANY, VOLUME 24

Mexico; Honduras; Salvador; Panama; West Indies; Colombia
and Venezuela.

A shrub or small tree 2-5 meters high, the bark smooth, grayish, the branchlets
densely lepidote; leaves short-petiolate, linear to obovate, 5-8 cm. long, acute or
obtuse, obtuse at the base, glabrous above, densely or sparsely lepidote beneath;
flowers small, white, in pedunculate umbelliform few-flowered racemes from the
ends of the branches, fragrant, long-pedicellate; calyx open in bud, the sepals
subulate or lanceolate, 2-3 mm. long; petals elliptic, 10-12 mm. long, tomentose
within, lepidote outside; stamens about 16, almost twice as long as the petals;
ovary falsely 2-celled, borne on a very long gynophore; fruit siliquiform, 6-25 cm.
long, 1 cm. or less in diameter, terete, somewhat torulose, densely brownish-
lepidote, finally 2-valvate; seeds surrounded by scarlet pulp.

Called "guacoco" and "curumo" in Salvador; "taiche" (Yucatan,
Maya). The wood is white when first cut, turning pink upon
exposure.

Capparis Lundellii Standl. Carnegie Inst. Wash. Publ. 461:
57. 1935.

Known only from the region of the type, San Andre's, Petdn,
C.L.LundellSUS.

Branchlets densely silvery-lepidote, sometimes complanate; leaves short-
petiolate, coriaceous, the petiole 5-9 mm. long; leaf blades cuneate-obovate, 7-12
cm. long, 3.5-5 cm. wide, abruptly acuminate, cuneately narrowed to the base, the
base itself narrowly rounded, glabrous above, the costa impressed, beneath densely
and minutely whitish-lepidote; inflorescence cymose-paniculate, 5-6 cm. wide,
dense and many-flowered, borne on a peduncle 4-5 cm. long, the branches com-
planate, the pedicels subumbellate, 6-8 mm. long; calyx disk-like, open in bud,
almost 3 mm. wide, the sepals triangular, acute, subreflexed; petals obovate,
8-10 mm. long, rounded at the apex, stellate-tomentose outside, with a large
orbicular glabrous gland at the base; stamens numerous, the filaments 2.5 cm.
long or more.

Capparis quiriguensis Standl. Proc. Biol. Soc. Wash. 37: 52.
1924.

Wet forest or thickets, often along stream banks or swamps,
400 meters or less; Pete"n; Alta Verapaz; Izabal (type from Quirigua,
Standley 24048). British Honduras.

A shrub or tree, sometimes 12 meters high with a trunk 22 cm. in diameter,
the crown dense and spreading, the branchlets puberulent or glabrous; petioles
slender, 3-12 cm. long, glabrous or sparsely puberulent; leaf blades thin, elliptic
to elliptic-oblong, 11-23 cm. long, 4.5-14 cm. wide, usually acute or acuminate,
rarely obtuse, subacute to rounded at the base, bright green, glabrous or sometimes
hirtellous beneath along the nerves; flowers greenish white, in terminal racemes
4-10 cm. long, the rachis usually puberulent, the flowers few or numerous, the
pedicels 2-4 cm. long, generally puberulent; sepals imbricate in bud, rounded,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 387

5 mm. long, usually puberulent and ciliate; petals 1 cm. long, glabrous; stamens
very numerous, about 3 times as long as the petals; fruit subglobose or oblong,
3.5 cm. long or probably even longer, 1.5 cm. broad, smooth or nearly so, borne
on a gynophore 2-2.5 cm. long.

This has been reported from both Pete"n and British Honduras
as C. Tuerckheimii Donn. Smith.

Capparis Steyermarkii Standl. Field Mus. Bot. 22: 140. 1940.

Known only from the type, Rio Dulce, between Livingston and

6 miles up the river, on the north side, near sea level, Steyermark
39387.

A small tree, the branches densely stellate-pilose with coarse brown hairs;
leaves thick-membranaceous, the stout petioles 1.5-3 cm. long; leaf blades obovate
or oblong-obovate, 18-21 cm. long, 6-10 cm. wide, abruptly acuminate or long-
acuminate, cuneately narrowed to the obtuse base, at first stellate-tomentose above
but soon glabrate, softly stellate-pilose beneath; flowers white, umbellate-racemose
or simply umbellate, axillary, 13 cm. long or less, long-pedunculate, the slender
pedicels 17 mm. long or less; calyx open in bud, the sepals linear, 5-6 mm. long;
petals broadly obovate, 7-8 mm. long, densely stellate-pilose outside with
appressed hairs; stamens 8; ovary linear, 5 mm. long, borne on a gynophore 7 mm.
long.

Gapparis Tuerckheimii Donn. Smith, Bot. Gaz. 46: 100. 1908.

Baja Verapaz (type collected near Panzal, 1,200 meters, Tuerck-
heim 11.1746); Sacatepe"quez(?). Honduras.

A shrub or small tree, sometimes 8 meters high, glabrous throughout; leaves
thin, bright green, mostly on long slender petioles; leaf blades oblong-lanceolate
to oblong-elliptic, 7-18 cm. long, 3-7 cm. wide, usually rather abruptly long-
acuminate, acute or obtuse at the base; flowers white, in short terminal racemes,
the slender pedicels 2-6.5 cm. long; sepals ovate or oblong-ovate, 3-3.5 mm. long,
recurved, open in bud; petals 1.5 cm. long, glabrous; stamens numerous, 3 cm.
long; ovary cylindric-ellipsoid, 5 mm. long, the gynophore 4-4.5 cm. long; fruit
(only broken fruits seen) apparently large and globose.

Capparis verrucosa Jacq. Stirp. Amer. 159. pi. 99. 1763.
Naranjillo (Pete*n, fide Lundell).

Mostly in dry thickets, 660 meters or less; reported from Pete"n;
Chiquimula; Retalhuleu. Western and southern Mexico; Costa
Rica; Panama; West Indies; Colombia and Venezuela.

A shrub or small tree, seldom more than 5 meters high, the branchlets usually
puberulent or scabrous; leaves almost sessile, subcoriaceous, oblong to oblong-
elliptic or obovate-oblong, 4-8 cm. long, acute, subacute or emarginate at the base,
commonly scabrous or hispidulous beneath, at least along the costa, but some-
times glabrous; flowers white, mostly in short racemes, these terminal or in the

388 FIELDIANA: BOTANY, VOLUME 24

upper leaf axils, the pedicels short or elongate; sepals imbricate, rounded, 4 mm.
long; petals glabrous, broad, about 1.5 cm. long; stamens very numerous, 2.5 cm.
long; fruit oblong, densely and obtusely tuberculate, 2.5-6 cm. long, 2 cm.
thick, longer than the gynophore.

CLEOME L.

Herbs, or the plants sometimes suffrutescent, glabrous or glandular, some-
times scandent; leaves simple or palmately 3-7-foliolate, the leaflets entire or
serrulate; flowers solitary or racemose, white to purple or yellow; calyx 4-dentate
or 4-parted, persistent or deciduous; petals subequal, sessile or unguiculate, con-
volute, imbricate, or open in bud; torus short, sometimes appendaged dorsally;
stamens 6 or rarely 4, inserted on the torus, all or only 2 antheriferous; filaments
generally unequal and declinate; ovary sessile or stipitate, the ovules numerous,
the style very short or the stigma sessile; fruit capsular, short or usually elongate
and siliquiform, sessile or stipitate, 1-celled, the valves membranaceous; seeds
reniform, usually roughened or pubescent.

About 75 species, chiefly in tropical regions, in both hemispheres.
A few additional species are known from southern Central America.

Leaves simple, linear C. ephemera.

Leaves digitately compound, the leaflets broader than linear.
Plants armed with axillary spines.

Gynophore equaling or longer than the pedicel C. spinosa.

Gynophore much shorter than the pedicel.

Leaflets 5 in all or most of the leaves, acuminate C. Houstoni.

Leaflets 3, obtuse C. aculeata.

Plants unarmed.

Leaflets 3 C. serrata.

Leaflets 5 or more.

Gynophore shorter than the pedicel; petals less than 1 cm. long. . . C. pilosa.
Gynophore several times as long as the pedicel; petals 2-2.5 cm. long.

C. parvisepala.

Cleome aculeata L. Syst. Nat. ed. 12. 232. 1768.

Moist brushy places, 200 meters or less; Pete"n. Mexico; Hon-
duras; West Indies; South America.

An erect annual, less than a meter high, branched, glandular-puberulent,
armed with stipular spines; leaves long-petiolate, 3-foliolate; leaflets thin, elliptic
to obovate or elliptic-ovate, 1.5-6 cm. long, obtuse or subacute, unequal at the
obtuse base, glandular-puberulent; upper leaves reduced to simple broad green
bracts, these sessile; flowers solitary in the axils of the bracts, white; sepals lanceo-
late or oblong, 2-3 mm. long; petals 5 mm. long, unguiculate; stamens 6; capsule
cylindric, 2.5-5.5 cm. long, 3-4 mm. broad, short-stipitate, striate; pedicels 1-2.5
cm. long; seeds 2.5 mm. broad, reticulate-tuberculate.

Called "flor de caballero" in Yucatan.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 389

Cleome ephemera Brandeg. Proc. Calif. Acad. Sci. II. 3: 112.
1891.

Dry rocky hillsides, 200 meters; Zacapa (near Rosalia, Steyer-
mark 29281). Western Mexico.

A slender delicate annual, 30 cm. high or less, simple or sparsely branched,
glabrous throughout; leaves linear or subulate, 2-3 cm. long, scarcely 1 mm. wide;
flowers solitary in the upper leaf axils or few in terminal naked racemes, 15 mm.
long, the filiform pedicels 5 mm. long or less; sepals very small, linear; petals
yellow, short-unguiculate; stamens 8, only 4 of them perfect, equaling or shorter
than the petals; ovary short-stipitate, about 20-ovulate; capsule almost sessile,
2-2.5 cm. long, 2 mm. wide, short-stipitate; seeds muricate.

Very unlike most members of the genus because of its exceed-
ingly narrow, simple leaves.

Cleome Houston! R. Br. in Ait. Hort. Kew. ed. 2. 4: 131. 1812.

Wet thickets, at or little above sea level; Izabal. Panama;
Greater Antilles.

A coarse erect annual a meter high or less, branched, armed with short stout
yellowish stipular spines, the stems glandular-puberulent and sparsely glandular-
pilose ; leaves long-petiolate, the petioles aculeate ; leaflets 5 in all except the upper-
most leaves, lanceolate to lance-ovate, thin, 3-9 cm. long, long-acuminate, acute
at the base, glandular-puberulent or glabrate, often aculeate beneath on the costa;
flowers in terminal racemes, white or purple and white, the racemes 10-25 cm. long,
the bracts large, ovate or oblong, green, cordate, sessile; pedicels 1.5-2.5 cm. long,
or longer in fruit, much longer than the gynophore; sepals linear-lanceolate, green;
petals unguiculate, 1 cm. long or shorter; stamens 6, shorter than the petals;
capsule linear, 5-9 cm. long, 3-5 mm. broad; seeds muriculate dorsally.

Cleome parvisepala Heilborn, Arkiv Bot. 23A, no. 10: 12. 1931.

Moist or wet forest or thickets, 2,700-2,850 meters; Huehue-
tenango(?); Quezaltenango; San Marcos. Southern Mexico.

A slender but coarse, sparsely branched herb or shrub 2-5 meters high, very
densely glandular-puberulent throughout, often also glandular-pilose; leaves on
very long petioles, the leaflets 5-11, oblong-oblanceolate, mostly 6-14 cm. long,
acuminate, attenuate to the base and short-petiolulate; flowers large, greenish
cream, in greatly elongate, terminal, bracteate racemes; bracts large and foliaceous,
broadly ovate, cordate, sessile or the lowest petiolulate, mostly obtuse; pedicels
3.5 cm. long or less; sepals triangular-ovate, obtuse, glandular-puberulent; petals
2-2.5 cm. long; filaments purple, greatly elongate; gynophore becoming 2-3 times
as long as the pedicels, recurved; capsule linear, 7-14 cm. long, 3 mm. thick,
longitudinally striate, glandular-pubescent; seeds conspicuously and irregularly
cristate.

A rather conspicuous but not at all handsome plant, disagree-
ably clammy when handled; apparently rare in the Occidente.

390 FIELD IANA: BOTANY, VOLUME 24

Cleome pilosa Benth. Bot. Voy. Sulph. 65. 1844. Alcachofla
(Santa Rosa) ; Alcachofa de monte.

Chiefly in moist thickets, sometimes in wet forest or on dry
rocky slopes, often a weed in fields or along roadsides, 250-2,000
meters; Alta Verapaz; Izabal; Zacapa; Chiquimula; Jutiapa; Santa
Rosa; Escuintla; Guatemala; Sacatepe"quez ; Chimaltenango; Hue-
huetenango; Quezaltenango ; San Marcos. Southern Mexico; Costa
Rica; Panama; Colombia and Venezuela.

An erect annual, a meter high or less, often much branched, the stems sparsely
or densely glandular-pilose, unarmed; leaves long-petiolate; leaflets 5, oblanceolate,
mostly 5-10 cm. long, acuminate, attenuate to the base, sessile or short-petiolulate,
sparsely pilose with short gland-tipped hairs; racemes elongate but few-flowered,
the bracts ovate-lanceolate, acute at the base, green, the pedicels long, almost
filiform; sepals minute, linear-lanceolate; petals pale purple or greenish, 1 cm.
long or usually shorter, long-unguiculate; stamens 6, longer than the petals, all
fertile; capsule short-stipitate, 5-8 cm. long, 5 mm. broad, glandular-pilose or
glabrous; seeds lustrous brown, bearing few scattered obtuse tubercles.

Cleome serrata Jacq. Enum. PL Carib. 26. 1760. Mirame-
linda (Pete"n, fide Lundell).

Moist or wet thickets or in waste or cultivated ground, a common
weed in banana plantations, 500 meters or less; Pete"n; Alta Verapaz;
Izabal; Santa Rosa; Escuintla. Southern Mexico; British Hon-
duras to Panama; West Indies; South America.

A glabrous erect annual, a meter high or less, sparsely branched; leaves long-
petiolate, the 3 leaflets lanceolate or narrowly elliptic, 4-14 cm. long, obscurely
serrulate, acute or acuminate at each end; flowers white, often tinged with purple,
in lax few-flowered terminal racemes, the racemes not bracteate; pedicels 1-1.5
cm. long; petals 1 cm. long or shorter, long-unguiculate; sepals green, lanceolate,
serrulate; stamens 6, equaling the petals; capsule linear, 5-7 cm. long, 3 mm.
thick, sessile or short-stipitate; seeds sparsely and minutely muriculate dorsally.

Known in Veracruz by the names "ejotillo," "chispa," and
"chilpate." A common weedy plant of the whole Atlantic coast
region of Central America.

Cleome spinosa Jacq. Enum. PI. Carib. 26. 1760.

Open slopes, sandy thickets, often in sand or gravel along stream
beds, or a weed in waste ground, 1,050 meters or less; Izabal; Zacapa;
Jutiapa; Santa Rosa; Retalhuleu. Mexico; British Honduras to
Salvador and Panama; West Indies; South America.

A coarse erect glandular-pubescent herb 1.5 meters high or less, armed with
short yellowish stipular spines; leaves long-petiolate, the petioles usually aculeate;
leaflets 5-7, oblanceolate or lanceolate, 4-10 cm. long, acute or acuminate, often

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 391

aculeate beneath along the costa; flowers in elongate racemes, the bracts large
and conspicuous, ovate or oval, subcordate at the base, sessile; sepals small, green,
linear; petals purple or whitish, about 2 cm. long, glandular outside; stamens 6,
crimson, long-exserted ; gynophore much longer than the pedicel, sometimes equal-
ing the capsule; stigma sessile; capsule linear, glabrous or puberulent, 5-12 cm.
long, 3-4 mm. thick; seeds almost smooth.

Known in Salvador as "alheli" or "alelia." Although sometimes
grown in the United States for ornament, the plant is not an attrac-
tive one, and it has a strong, far from pleasant odor.

CRATAEVA L.

Shrubs or trees, glabrous or pubescent, the branches with conspicuous pale
lenticels; leaves 3-foliolate, long-petiolate, the leaflets entire, thin; flowers corym-
bose, the corymbs axillary and terminal, often polygamous; calyx 4-parted, the
lobes deciduous, imbricate in bud; petals 4, long-unguiculate, open in bud; torus
hemispheric, lobate; stamens 8-20, inserted on the margin of the torus, the fila-
ments filiform, elongate; ovary ovoid, long-stipitate, 1-2-celled, with 2 placentae;
ovules numerous, multiseriate; stigma sessile, discoid; fruit baccate, globose or
ovoid, 1-2-celled; seeds few-many, reniform, surrounded by pulp, the testa mem-
branaceous; cotyledons incumbent-convolute, the radicle conic.

Species about 10, in the tropics of both hemispheres. Only the
following are known in continental North America.

Leaflets glabrous C. Tapia.

Leaflets densely puberulent beneath C. Palmeri.

Crataeva Palmeri Rose, Contr. U. S. Nat. Herb. 1: 301. 1895.
Cadeno.

Dry brushy plains or hillsides, 200-500 meters; El Progreso;
Zacapa; Chiquimula; Suchitepe"quez(?). Western Mexico.

A large shrub or small tree, about 6 meters high, with a broad crown; leaves
on long slender petioles, obliquely ovate or elliptic, 3.5-6 cm. long, acute or acumi-
nate, rounded to acute at the base, green above and glabrous or nearly so, paler
beneath, densely and finely puberulent; flowers usually produced when the tree is
leafless, purplish, in short dense many-flowered racemes, the pedicels about 3 cm.
long, glabrous; sepals 4 mm. long, ovate, acute, contracted below, costate; stamens
5-6 cm. long or longer, the anthers 6 mm. long; fruit 3.5-5 cm. long.

This is a common and conspicuous small tree in the Zacapa
area. All the Guatemalan specimens are sterile but probably
referred here correctly.

Crataeva Tapia L. Sp. PI. 444. 1753. Matasanillo, Granadillo
(fide Aguilar); Tortugo (Izabal).

392 FIELDIANA: BOTANY, VOLUME 24

Moist or wet forest or thickets, 1,400 meters or less, usually at
or near sea level; Pete"n; Izabal; Santa Rosa; Guatemala. Mexico;
British Honduras to Salvador and Panama; West Indies; South
America.

A tree, 6-18 meters high, glabrous throughout, the trunk 45 cm. or less in
diameter, the bark grayish brown; leaves deciduous, long-petiolate, the leaflets
petiolulate, ovate to oblong-elliptic, 5-15 cm. long, acute or acuminate, green
beneath; flowers long-pedicellate, crowded in chiefly terminal corymbs; sepals
oblong, 5-7 mm. long; petals white or greenish, 1-2 cm. long, oblanceolate or
oblong, long-unguiculate; stamens 2.5-6 cm. long, purple or purplish, the anthers
yellow; fruit borne on a gynophore 3-6 cm. long, globose or ovoid, 2-5 cm. long;
seeds 8 mm. in diameter.

Known in British Honduras as "waika bead" and "yuy";
"cachimbo" (Honduras); "anonillo," "granadillo macho" (Salva-
dor); "cascaron" (Tabasco); "colocmax" (Maya), "cascoron," "Tres
Marias" (Yucatan). The bark has a disagreeable odor. The roots
are acrid, and it is stated that the juice, when in contact with the
skin, produces blisters. The wood has an odor suggestive of garlic;
it is white or yellowish, only moderately hard, of medium texture,
fairly easy to work, brittle, not durable. It is suitable for paper
pulp and minor carpentry. Some authors, as Fawcett and Rendle
in Flora of Jamaica, divide the material here referred to C. Tapia
into two species, C. Tapia and C. gynandra L., but we are unable to
find characters by which the two can be separated clearly, and the
variations in the whole series of specimens are not greater than might
be expected in the case of a tropical tree. There is some question,
indeed, as to whether C. Palmeri is more than a variety of C. Tapia.

Crataeva Tapia var. glauca (Lundell) Standl. & Steyerm.
Field Mus. Bot. 23: 55. 1944. C. glauca Lundell, Bull. Torrey Club
69: 389. 1942.

British Honduras; Honduras; southern Mexico, the type from
Palisada, Campeche. Distinguished by having the lower surface
of the leaflets glaucous, otherwise in all respects like the typical
form of the species. Called "crucito" in Campeche. Although
described as a species, this form of rather wide distribution in Mexico
and Central America seems to have only a single character by which
it can be distinguished. Even this is not too definite, and it is some-
times difficult to determine whether the leaflets are really glaucous
or only green beneath.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 393

FORCHHAMMERIA Liebmann

Trees or shrubs, glabrous or pubescent; leaves alternate, simple or digitately
3-foliolate, petiolate, coriaceous, entire; stipules none or minute; flowers dioecious,
small, apetalous, racemose or paniculate; calyx minute, 4-8-dentate, the teeth
unequal; stamens numerous in the staminate flower, inserted on a low fleshy torus;
disk of the pistillate flower very short, produced into 8-12 deciduous teeth; ovary
2-celled, the stigmas connate to form an orbicular-peltate, obscurely 2-lobate
disk; fruit small, ovoid or globose, by abortion usually 1-celled, drupaceous,
indehiscent, the stigmas becoming lateral as the fruit develops; seeds 1 in each
cell, the testa subcoriaceous.

About 9 species, in Mexico, northern Central America, and
Hispaniola. Only the following are known from Central America.

Fruiting panicles usually 15-50 cm. long; pedicels mostly equaling the fruit or
longer F. Matudai.

Fruiting panicles mostly 8-12 cm. long or shorter; pedicels usually shorter than
the fruits F. trifoliata.

Forchhammeria Matudai Lundell, Lloydia 2: 87. 1939.
Comida de pasha.

Moist forest, 1,000-1,300 meters; Sacatepe'quez; Chimaltenango;
Quezaltenango. Chiapas, the type from Mount Ovando.

A large shrub or a tree, sometimes 14 meters high, the trunk as much as 45
cm. in diameter, glabrous throughout; leaves 2-3-foliolate or rarely simple, mostly
3-foliolate, the compound leaves long-petiolate; leaflets chartaceous or almost
coriaceous, narrowly oblong-oblanceolate, widest at or near the middle, 10-20
cm. long, 2-4 cm. wide, attenuate-acuminate, attenuate to the base; fruiting
panicles many-flowered, lax, pendent, 15-50 cm. long, the slender pedicels mostly
8-15 mm. long; fruit subglobose, 8-9 mm. in diameter.

Material reported from Salvador as F. trifoliata probably is to
be referred here, but we have not seen the specimens on which the
record is based.

Forchhammeria trifoliata Radlk. Field Mus. Bot. 1: 399.
1898. Tres Marias.

Mixed limestone forest, 600 meters or less; Pete*n; Alta Verapaz.
Yucatan Peninsula of Mexico; British Honduras.

A glabrous shrub or tree, sometimes 12 meters high; leaves usually trifoliolate,
on short or long petioles; leaflets obovate-oblong or oblanceolate-oblong, mostly
10-16 cm. long and 3.5-6 cm. wide, obtuse or acute, often abruptly acute, cuneate-
attenuate to the base, subcoriaceous, paler beneath; fruiting panicles mostly about
as long as the petioles, sometimes longer, racemiform, few-flowered, the pedicels
rather stout, nearly always shorter than the fruit; fruit subglobose, asymmetric,
about 1 cm. in diameter.

394 FIELDIANA: BOTANY, VOLUME 24

Known in British Honduras as "bastard dogwood" and "wild
craboo."

GYNANDROPSIS De Candolle

Annual herbs, glabrous, pilose, or glandular-pubescent, unarmed; leaves
digitately 3-7-foliolate, the leaflets entire; flowers small or large, white or purple,
in leafy-bracteate racemes; sepals spreading, deciduous; petals entire or crenulate,
obovate, unguiculate, imbricate or open in bud; torus hemispheric or depressed,
narrowed into an elongate gynophore; stamens 6, all fertile, the filaments connate
below into a tube adnate to the gynophore; ovary stipitate, elongate, with 2
placentae, many-ovulate; style short or elongate, the stigma small, capitate,
bilobate; fruit capsular, sessile or stipitate, compressed or subterete, usually linear;
seeds reniform or orbicular, compressed, the testa rugose or tuberculate; cotyledons
incurved, accumbent.

About 15 species, in the tropics of both hemispheres. Several
other species are known from Central America and one, G. gynandra
(L.) Briq., differing from the following in its small flowers and
3-foliolate bracts, is to be expected in Guatemala.

Gynandropsis speciosa (HBK.) DC. Prodr. 1: 238. 1824.
Cleome speciosa HBK. Nov. Gen. & Sp. 5: 84. pi. 436. 1821. Aleli;
China silita.

Moist thickets, open fields or waste ground, sometimes on gravel
bars along streams, 1,800 meters or less; Pete"n; Alta Verapaz; Izabal
(cultivated); Jalapa (planted); Santa Rosa; Escuintla; Guatemala;
Solola; Suchitepe"quez. Mexico; Honduras and Salvador to Panama;
West Indies; South America.

A coarse annual, often a meter high or more, branched, the stems short-
villosulous; leaves long-petiolate, the leaflets 5-9, narrowly elliptic or oblanceolate,
7-18 cm. long, long-acuminate, attenuate to the base, glabrous or nearly so;
racemes often much elongate and many-flowered, the bracts large and conspicuous,
foliaceous, ovate or oval, sessile, entire; flowers on long, almost filiform pedicels;
sepals small, green, linear-lanceolate, glabrous; petals rose or rose-purple, rarely
white, 2.5-3 cm. long, glabrous; stamen tube about 8 mm. long, appearing in fruit
as a scar near the base of the very long and slender gynophore; filaments greatly
elongate and filiform; capsule subterete, linear, 7-9 cm. long, 3 mm. thick, longi-
tudinally nerved; seeds brown, 1.5 mm. broad.

Known in Salvador as "alelia," "flor de Mayo," and "barba del
rey." A rather showy plant, often grown for ornament in Guate-
mala. We do not know whether it is really native in this area or
merely an escape from cultivation. The plants are scarce, and
usually only one or two are found in a locality.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 395

MORISONIA L.

Unarmed shrubs or small trees, glabrous or variously pubescent; leaves simple,
petiolate, coriaceous, often more or less peltate, entire; flowers corymbose, axillary
and terminal, many-flowered, the flowers rather large or small; calyx campanulate
or ventricose, bilobate or 2-4-fid, with 4 glands within at the base; petals 4,
unguiculate, obtuse, alternating with the glands of the calyx; torus produced into
an elongate gynophore, this bearing the stamens; stamens 6-20, shorter than the
corolla, the filaments subulate; ovary ovoid, stipitate, 1-celled or finally 4-celled,
many-ovulate; stigma discoid, sessile; fruit large, baccate, globose; seeds numer-
ous, surrounded by pulp, the testa crustaceous; cotyledons foliaceous-carnose,
convolute, the radicle fusiform.

About 4 species, in tropical America. Only one is known from
continental North America.

Morisonia americana L. Sp. PI. 503. 1753.

Dry brushy plains or open forest, 120 meters or less; Retalhuleu
and perhaps elsewhere. Western Mexico; Lesser Antilles; northern
South America.

A shrub or small tree, 7 meters high or less, with sparse or very dense, usually
minute and appressed, stellate or stellate-lepidote pubescence on almost all parts,
the leaves at maturity glabrous or nearly so; leaves long-petiolate, oblong or ovate-
oblong, 12-25 cm. long, obtuse or acute, usually rounded and obviously peltate at
the base, sometimes epeltate, often thick-coriaceous, lustrous; flowers rather large,
in lateral corymbs, white; fruit globose, 3.5-6 cm. in diameter, brownish and rough,
sessile or nearly so.

Although rather widely distributed in Mexico and Guatemala,
this shrub seems to be rare or at least local, and it is seldom collected.
In general appearance it suggests the genus Capparis, but it may be
distinguished from Guatemalan species of that genus by its usually
peltate leaves.

POLANISIA Rafinesque

Erect annual herbs, glandular-pubescent and ill-scented; leaves mostly digi-
tately 3-9-foliolate, the uppermost leaves reduced to foliaceous bracts; sepals
lanceolate, free or connate at the base, deciduous; petals sessile or unguiculate,
entire, equal or unequal, imbricate in bud; torus small, depressed, sometimes with
a posterior gland ; stamens 8 or more, inserted at the base of the torus, some of them
occasionally sterile, declinate in an thesis; filaments filiform; ovary sessile or stipi-
tate, usually glandular, the ovules very numerous; style elongate or the stigma
subsessile; capsule linear, sessile or stipitate, cylindric or compressed; seeds reni-
form, transversely rugose or reticulate; cotyledons incumbent, incurved.

Species about 15, in warmer regions of both hemispheres. Only
one is known in Central America.

396 FIELDIANA: BOTANY, VOLUME 24

Polanisia viscosa (L.) DC. Prodr. 1: 242. 1824. Cleome viscosa
L. Sp. PL 672. 1753.

Dry or moist plains or fields, sometimes on gravel bars along
streams, 200-500 meters; Zacapa. Oaxaca; British Honduras;
Salvador; Nicaragua; West Indies; Old World tropics.

A coarse annual, a meter high or less, densely glandular-pubescent throughout;
leaves small, long-petiolate, the leaflets 3-5, obovate or elliptic, 1-4 cm. long,
obtuse or acute, sessile or nearly so, thin; flowers solitary in the axils of the upper
leaves or bracts, long-pedicellate; sepals narrowly oblong, 5-6 mm. long, decidu-
ous; petals yellow, obovate, 1 cm. long; stamens 12-20, usually shorter than the
petals; capsule linear, terete, sessile, 6-8 cm. long, 3 mm. thick, densely glandular-
pubescent, the style 4 mm. long; seeds cochleate, transversely cristate.

Called "tabaquillo" in Salvador. The plant appears to be rare
in continental North America.

STERIPHOMA Sprengel

Unarmed shrubs with stellate pubescence; leaves long-petiolate, simple,
entire; flowers large, showy, in terminal racemes, the pedicels thickened at the
apex, decurved; calyx cylindric-campanulate, 2-4-lobate at the apex, rupturing
irregularly, with 4 small scales within at the base; torus very short, forming an
annular disk; petals 4, sessile, inserted on the torus, the 2 anterior ones slightly
larger; stamens 6, inserted with the petals, ascending, the 2 posterior ones shorter;
filaments long-exserted, the anthers large; ovary ovoid or oblong, 2-celled, the
ovules numerous, 2-seriate; stigma sessile; fruit baccate, globose or angulate;
seeds numerous, surrounded by pulp, angulate; cotyledons spirally convolute.

About 6 species in tropical America. One other Central American
species has been described from Panama.

Steriphoma clara Standl. Field Mus. Bot. 22: 21. 1940.

Dry brushy plains, 120 meters or less; endemic; Retalhuleu
(type collected in thickets near Nueva Linda, halfway between
Retalhuleu and Champerico, Standley 66552).

A slender shrub 1-3 meters high, sometimes subscandent, with few branches,
the young branches covered with a dense subappressed brown stellate tomentum,
later glabrate; leaves membranaceous, the slender petioles 4-9 cm. long; leaf blades
oblong-elliptic or obovate-elliptic, 8-16 cm. long, 3.5-7 cm. wide, acute or rather
abruptly acuminate, obtuse or narrowly rounded at the base, glabrous above, at
least in age, minutely stellate-pubescent beneath or glabrate; racemes dense and
many-flowered, 18 cm. long or shorter, the bracts filiform, caducous; pedicels erect,
decurved at the apex, mostly 2.5-3.5 cm. long, densely covered with a scarlet
stellate tomentum; calyx campanulate, 1.5 cm. long, rounded at the base, very
densely covered with a scarlet stellate tomentum, the lobes broadly ovate, obtuse,
5 mm. long; petals pale yellow, narrowly oblong, obtuse, shortly exserted from the
calyx, stellate-tomentulose on the outer surface; filaments long-exserted, pale
green, the anthers 7 mm. long.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 397

This plant has been reported from Guatemala as S. paradoxa
Endl., a South American species from which it is distinct. It is an
abundant shrub on the plains of Retalhuleu, flowering during the
height of the dry season when many shrubs are more or less dormant.
The fire-red flowers make it very conspicuous at this time, and rather
handsome. The racemes are spire-like in form, standing above the
foliage, and in both form and coloring are strikingly suggestive,
from a distance, of the inflorescences of Combretum Cacoucia of the
Caribbean coast.

RESEDACEAE. Mignonette Family

Annual or perennial herbs, rarely suffrutescent; leaves alternate or fasciculate,
entire to lobate, the stipules gland-like; flowers small, perfect, asymmetric; calyx
4-7-parted, somewhat unequal; petals usually 4-7, entire or cleft, hypogynous;
disk fleshy, hypogynous, one-sided; stamens 3-40, inserted on the disk, the fila-
ments usually unequal; ovary 1, compound, of 3-6 carpels; styles or sessile stigmas
3-6; ovules numerous in the cell; fruit usually capsular, 3-6-lobate; seeds small,
reniform, without endosperm, the cotyledons incumbent.

About 6 genera and 65 species, native chiefly in the Mediter-
ranean region, none of them American.

RESEDA L. Mignonette

Annual or perennial herbs, erect or decumbent; leaves entire or pinnatifid;
flowers small, white, yellowish, or greenish, spicate or racemose; petals 4-7,
dentate or cleft; disk cup-shaped, glandular; stamens 8-40, inserted on one side
of the flower on the inner surface of the disk; capsule 3-6-lobate, with short horn-
like projections at the apex, opening at the apex before the seeds mature.

About 55 species, all native in the Old World. R. Luteola L.,
cultivated in some regions as the source of a yellow dye, has become
well established in Mexico but it has not been noted in Central
America.

Leaves entire R. odorata.

Leaves pinnatifid R. alba.

Reseda alba L. Sp. PI. 449. 1753.

Frequent in gardens of the Occidente, particularly about Quezal-
tenango. Native of southern Europe.

Plants stout, erect, apparently perennial, often much branched and some-
times hard and suffrutescent below, glabrous, with pale stems, the whole plant
often glaucous; leaves crowded, pinnate or deeply pinnatifid, the numerous seg-
ments short, linear or oblong, subobtuse, entire or undulate; flowers white or

398 FIELDIANA: BOTANY, VOLUME 24

whitish, almost sessile, forming short or long, spike-like racemes; petals 6 or 5,
3-cleft at the apex; stamens 12-15; capsule ovoid-oblong, 1 cm. long.

The flowers have a rather unpleasant odor and the plant, at
least as it grows in Guatemala, is not an attractive one for cultiva-
tion. It is noteworthy about Quezaltenango for its ability to bloom
during the coldest months and often is seen in gardens of even the
poorest dwellings, where, neglected and mistreated, broken by
domestic animals and covered with dust, it often appears most
forlorn.

Reseda odorata L. Syst. Nat. ed. 10. 1046. 1759. Reseda.
Mignonette.

Native of northern Africa, but grown in gardens in many remote
parts of the earth. Frequently planted in gardens of Guatemala for
its sweet-scented flowers.

A branched annual, at first erect, in age decumbent; leaves spatulate or
oblanceolate, obtuse, entire or essentially so; flowers yellowish white, in spike-like
racemes, these becoming lax and open in age.

Mignonette flowers are offered for sale in the markets of Guate-
mala and other cities, although the flowers have little besides their
odor to recommend them.

MORINGACEAE

Unarmed trees, the root with a pungent odor, the bark exuding gum; leaves
deciduous, alternate, 2-3-pinnate, the pinnae and pinnules opposite, the leaflets
entire, caducous; stipules none or reduced to glands; flowers perfect, irregular, in
puberulent axillary panicles, rather large, white or reddish; calyx tube short,
cup-like, the limb 5-parted, the lobes unequal, spreading-reflexed, imbricate;
petals 5, similar to the sepals, the 2 upper ones smaller; disk lining the calyx tube,
the margin very short and free; stamens inserted on the edge of the disk, declinate,
5 of them perfect, alternating with as many sterile ones, the filaments free, rather
thick; anthers dorsifixed, oblong, 1-celled, anteriorly dehiscent; ovary stipitate,
terete, villous, 1-celled, the 3 placentae parietal; style terminal, slender, truncate
at the apex; ovules numerous, biseriately affixed to the placentae, pendulous,
anatropous; capsule silique-like, large and much elongate, rostrate, 3-6-angulate,
torulose, 1-celled, 3-valvate, many-seeded; seeds large, ovate, 3-winged or wingless,
the wings membranaceous; embryo without endosperm, orthotropous, the radicle
very short, superior.

The family consists of a single genus.

MORINGA Jussieu

Four species are recognized, natives of northern and eastern
Africa and western Asia. One species is cultivated in most tropical
regions.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 399

Moringa oleifera Lam. Encycl. 1: 398. 1783. Guilandina
Moringa L. Sp. PI. 381. 1753. M. pterygosperma Gaertn. Fruct. &
Sem. 2: 314. 1791. M. Moringa Millsp. Field Mus. Bot. 1: 490. 1902.
Perlas; Paraiso bianco; Marengo.

Native of eastern Africa and perhaps of the East Indies, planted
generally in tropical America for ornament; cultivated commonly
in the warmer parts of Guatemala, chiefly in the tierra caliente, and
naturalized in many localities; Pete"n; Zacapa; Chiquimula; El
Progreso; Jutiapa; Santa Rosa; Escuintla; Guatemala; Retalhuleu;
San Marcos; doubtless in most of the other departments.

A large shrub or small tree, rarely as much as 10 meters high, the bark whitish,
the trunk usually thick and irregular, the crown small and dense, the branchlets
and leaves puberulent or glabrate; leaflets numerous, thin, oblong to obovate,
1-2 mm. long, obtuse, entire, pale; flowers numerous, fragrant, white, on pedicels
5-10 mm. long; sepals linear to linear-oblong, 9-13 mm. long; petals slightly larger
than the sepals; capsule linear, obtusely trigonous, pendent, 20-45 cm. long, 1-2
cm. thick; seeds broadly winged, 2.5-3 cm. long.

The English name is "horse-radish tree." In British Honduras
it is called "maranga" and "maranga calalu"; in the Yucatan Pen-
insula, "paraiso de Espafia" and "paraiso bianco"; in Salvador,
"teberinto," "terebinto," "teberindo," and "marango." The thick,
fleshy roots have the flavor and odor of horse-radish (Armoracia) ,
for which they have been substituted at times. The wood has been
reported to yield a blue dye. In India the young leaves, pods, and
flowers are cooked and eaten. From the seeds is obtained ben oil
of commerce, used for lubricating watches and other delicate
machinery. Being odorless and never becoming rancid, it has been
found useful in manufacture of perfumes. Although so common in
the warmer parts of Central America, the tree is neither particularly
handsome nor desirable in cultivation for ornament or shade. The
trees usually are irregular in form ; neither the foliage nor the flowers
are especially attractive, at least in old trees, and the weak branches
are easily broken.

DROSERACEAE. Sundew Family
Reference: L. Diels, Droseraceae, Pflanzenreich IV. 112. 1906.

Low herbs, usually provided with gland-tipped hairs, especially on the leaves;
leaves mostly basal; stipules present or absent; inflorescence lateral or terminal,
cymose or racemose, the branches often elongate and recurved; bracts present or
absent, the pedicels naked; flowers usually small, perfect, 5-parted; sepals more or
less connate at the base, imbricate, persistent; petals hypogynous, imbricate,
marcescent; stamens 5-20, the filaments generally free, filiform or nearly so;

400 FIELDIANA: BOTANY, VOLUME 24

anthers 2-celled, the cells dehiscent by extrorse slits; disk none; ovary 5-3-carpel-
late, free, superior, 1-celled, the placentae parietal or basal; styles 5-3, usually free,
simple or divided; fruit a membranaceous capsule, loculicidally dehiscent; ovules
usually numerous, anatropous; endosperm carnose.

Four genera, two confined to the Old World, one to the eastern
coast of the United States, and one, containing all but three mem-
bers of the family, widely distributed. One of the most remarkable
plants of the family is the Venus flytrap of the coast of the Carolinas,
Dionaea muscipula Ellis. Its leaf blades are "hinged" down the
middle, the margins set with long spine-like bristles. When irritated
by contact with an insect, the blades fold together, trapping the
insect, which is used as food by the plant.

DROSERA L. Sundew

Mostly perennial herbs, the stems often scape-like; leaves alternate or often
all radical, glandular and provided with slender irritable tentacles; stipules present
or absent; inflorescence simple or branched, the flowers usually secund, small;
sepals mostly 5, connate at the very base; petals spatulate or cuneate-obovate;
stamens as many as the petals; styles 3-5, free or coherent at the base, simple or
usually divided; capsule 3-5-valvate, the seeds usually numerous.

More than 80 species, chiefly in the southern hemisphere, but
represented in most temperate and tropical regions of the earth.
Several species are native in North America, but only the following
in continental tropical North America.

Drosera capillaris Poir. in Lam. Encycl. 6: 299. 1804.

Wet savannas or about the margins of pools, at or little above
sea level; British Honduras; southern Mexico; West Indies; British
Guiana.

Stems very short; leaves numerous, forming a dense rosette at the base of the
plant, most of them lying flat on the ground, the inner ones erect or ascending, on
petioles 10-20 mm. long, the blades spatulate-obovate, rounded at the apex,
narrowed at the base, 3-7 mm. long, glabrous beneath, covered above with long
gland-tipped reddish hairs; peduncles scape-like, simple, naked, very slender,
5-15 cm. long, glabrous or nearly so, the flowers 4-10, on pedicels 1.5 mm. long
or shorter; sepals oblong-elliptic, minutely glandular or almost glabrous, 4-5 mm.
long; petals white or pink, 6-7 mm. long; seeds obovoid, costulate and papillose.

Called "spider plant" in British Honduras. This is an insecti-
vorous plant, which captures insects by the viscous exudate from
the glands that tip the hairs of the leaves. It probably will be found
in some of the pine or savanna areas of Izabal or Pete"n.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 401

PODOSTEMONACEAE

Reference: George V. Nash, Podostemonaceae, N. Amer. Fl.
22: 3-6. 1905.

Immersed annual or perennial herbs, usually creeping and closely adherent to
rocks in swift streams, the rootstocks branched or disk-like, the stems often fleshy;
leaves variable in form, in Central American plants small and scale-like or large
and divided into slender segments; flowers very small, perfect, naked or at first
enclosed in a spathe, this ruptured by the elongating pedicel and persistent at its
base; perianth none or of a few minute scales, sometimes large, membranous, and
3-dentate or 5-parted; stamens 1-many, hypogynous, when numerous the filaments
free in complete or incomplete verticels, or more or less united at the base, per-
sistent in fruit; anthers with 2 parallel cells; ovary free, sessile or stipitate, mostly
2-3-celled; styles 1-3, distinct or short-connate at the base, linear to foliaceous;
ovules numerous, inserted on central or parietal placentae; capsule 2-3-celled and
septicidally dehiscent, or sometimes 1-celled, the valves generally with obvious
nerves, 2 and equal or unequal, or 3 and equal; seeds minute, numerous, sessile;
endosperm none.

About 20 genera, in both hemispheres, chiefly in tropical regions.
One other genus, Blandowia, is represented in southern Central
America and may well reach Guatemala. Since the plants grow in
places often difficult of access where few or no other phanerogams
are found, they are neglected by most collectors, and the available
collections are therefore limited in number. The family has received
no serious monographic attention during the present century, and
the taxonomy of the American species, at least, is in an almost
chaotic state.

Flowers with a perianth; spathe none; stamen 1; leaves minute, scale-like, entire.

Tristicha.

Flowers without a perianth, or the perianth much reduced; spathe present; stamens
few-many; leaves not scale-like, dissected into narrow, often filiform seg-
ments Marathrum.

MARATHRUM Humboldt & Bonpland

Plants growing on rocks in usually swift water, attached by thickened fleshy
disks or by short dichotomous rhizomes; leaves in Central American species well
developed and rather large, decompound, divided into very numerous, short,
mostly linear and flaccid segments; flowers small, at first enclosed in a spathe,
this irregularly ruptured by the elongating pedicel; perianth rudimentary, of
minute scales; stamens 5-25, forming a complete whorl about the ovary, persistent
in fruit and suggesting rigid linear perianth segments; ovary 2-celled; styles 2,
free or somewhat united below; capsule dehiscent, of 2 equal valves, these con-
spicuously nerved longitudinally, persistent after dehiscence.

Species about 10, or perhaps much more numerous, in tropical
America. At least two other species are recorded from Central
America.

402 FIELDIANA: BOTANY, VOLUME 24

Capsule acute, 4-5 mm. long M. oxycarpum.

Capsule obtuse.

Pedicel conspicuously thickened at the apex into a cup-like hypanthium; capsule

4-5 mm. long M. Schiedeanum.

Pedicel not or scarcely thickened at the apex; capsule 2.5-3 mm. long.

M. modestum.

Marathrum modestum (Wedd.) Nash, N. Amer. Fl. 22: 4.
1905. M. Schiedeanum var. modestum Wedd. in DC. Prodr. 17: 54.
1873. M. minutiflorum Engler, Bot. Jahrb. 61, Beibl. 138: 4. 1927
(based in part on a Bernoulli collection from Mazatenango, Suchi-
tepe'quez). Piniju de piedra; Paxte de piedra; Muzgo.

On rocks in streams, 200-1,300 meters; Escuintla; Suchitepe"quez;
Retalhuleu; here may belong sterile material' from Chiquimula and
Retalhuleu. British Honduras; Nicaragua.

Plants rather small, forming dense colonies on rocks, the rhizome disk-like,
lobate; leaves mostly 3-10 cm. long but sometimes as much as 25 cm., petiolate,
repeatedly divided into very numerous small segments, these mostly linear or
nearly so and 1-2 mm. long, usually subacute; spathe 1 cm. long or less; mature
pedicels slender but stiff, 1.5-3.5 cm. long, not or scarcely thickened at the apex;
stamens 6-8, in age about 3 mm. long and exceeding the capsule, the anthers
deciduous, 1 mm. long; capsule 2.5-3 mm. long, oval or globose-obovoid, con-
spicuously nerved.

The plant is abundant in streams along the Pacific foothills,
often forming large colonies on great rocks in the swiftest parts of
the current, where it is nearly or quite impossible to reach them.
The distribution of this and other species in Guatemala is obscure,
since many of the collections are sterile and therefore not determin-
able except by sheer guess. This is probably the plant, at least in
part, that has been reported from Guatemala as M. foeniculaceum
Humb. & Bonpl. In Costa Rica plants of this genus, called there
"pasacarne," are sometimes important forage for cattle during the
dry season. The animals learn to wade into the streams and pull
the plants from the rocks, submerging their heads in doing so.

Marathrum oxycarpum Tulasne, Ann. Sci. Nat. III. 11: 94.
1849.

Reported from Guatemala by Nash, the locality not indicated ;
reported also from Mazatenango, Suchitepe'quez. Honduras;
Nicaragua; Panama.

Plants arising from a fleshy lobate rhizome; leaves petiolate, sometimes 40
cm. long, several times dissected into short flaccid filiform divisions, these often
3 mm. long or more, obtuse or subacute; fruiting pedicels mostly 4 cm. long or

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 403

more, slender, scarcely dilated at the apex, stiff; stamens 8-10, the filaments
subulate, slightly shorter than the capsule; capsule about 5 mm. long and 2 mm.
broad, ellipsoid, acute, conspicuously 8-costate; stigmas united below.

Marathrum Schiedeanum Chamisso, Linnaea 9: 504. 1835.

Submerged on rocks in swift streams, 200-600 meters; Suchite-
pe"quez; Retalhuleu. Southern Mexico; Costa Rica; Colombia.

Plants usually rather stout, from thick elongate rhizomes; leaves mostly 20-40
cm. long, petiolate, several times divided into small, linear, mostly obtuse seg-
ments, these spreading or ascending, rather firm; pedicels in fruit 2-8 cm. long,
slender but stiff, abruptly enlarged at the apex into a cupular hypanthium 1.5-2.5
mm. broad; stamens 6-8, the filaments subulate or narrowly linear, slightly shorter
or longer than the capsule; capsule ellipsoid, obtuse, 5 mm. long, obscurely or
conspicuously 8-costulate.

TRISTIGHA Thouars

Plants small and moss-like, firmly attached to large rocks beneath the water,
or often emersed for part of the year, the stems short, usually densely cespitose;
leaves very small, sessile, 3-ranked, crowded; flowers minute, mostly binate at
the ends of the branches; perianth membranous, 3-lobate, the lobes subimbricate;
stamen 1, the filament filiform; ovary 3-celled, with a central placenta; styles 3,
linear, short, erect; capsule septicidally 3-valvate, the valves equal, 3-nerved.

Three species, in tropical America and Africa. Only one occurs
in North America.

Tristicha hypnoides (St. Hil.) Spreng. Syst. Veg. 4, pt. 2: 10.
1827. Dufourea hypnoides St. Hil. Me"m. Mus. Paris 10: 472. 1823.
Pashtillo (Jutiapa).

On submerged rocks in usually swift streams, 1,400 meters or
less; Alta Verapaz; Izabal; Zacapa; Jalapa; Jutiapa; Santa Rosa;
Suchitepe'quez; Retalhuleu; Quezaltenango. Southern Mexico;
Honduras; Costa Rica; Cuba; tropical South America.

Plants small and moss-like, usually forming very dense and often large colonies
on smooth rocks, the slender stems densely crowded, mostly 1-3 cm. long but some-
times longer, rather stiff and rigid; leaves crowded, elliptic to broadly ovate, obtuse,
entire, commonly 3-ranked, 1-2 mm. long, dull green; pedicels 3-15 mm. long;
perianth segments oblong-elliptic; capsule ellipsoid, 1.5 mm. long, obtuse, scarcely
nerved.

Unless examined closely, this plant is likely to be taken for a moss
by those unacquainted with it, because the leaves and capsules
strongly suggest some of the acrocarpous mosses.

404 FIELDIANA: BOTANY, VOLUME 24

CRASSULACEAE. Stonewort Family

References: N. L. Britton and J. N. Rose, Crassulaceae, N. Amer.
Fl. 22: 7-74. 1905. A. Berger, Crassulaceae, in Engler & Prantl,
Pflanzenfam. ed. 2. 18a: 352-483. 1930.

Annual or usually perennial herbs, sometimes suffrutescent, the leaves usually
fleshy, often very thick, variously arranged, mostly simple and entire or dentate,
rarely compound; stipules none; flowers mostly small, sometimes rather large and
showy, cymose, racemose, or rarely solitary, regular, symmetrical, generally per-
fect; calyx hypogynous, persistent, commonly 4-5-parted or 4-5-1 obate; petals as
many as the calyx segments, free or more or less united, usually persistent, rarely
none; stamens as many or twice as many as the petals, the anthers longitudinally
dehiscent; receptacle usually with a scale at the base of each carpel; carpels of the
ovary and fruit as many as the sepals, distinct or united below; styles subulate or
filiform; ovules numerous, 2-seriate along the ventral suture of the carpel; seeds
small or minute, the endosperm carnose; embryo terete, the cotyledons short,
obtuse.

Different authors have divided the family very differently into
genera. Berger recognizes 33, a number that will be acceptable to
most botanists. The number recognized by Britton and Rose was
relatively much larger, 25 being recognized in North America alone.
Only the following genera are represented in Central America, and
in Central America below Guatemala only two species are native.
In North America the great majority of species are found in Mexico.
The family is a most unsatisfactory one to study from dried speci-
mens since the plants are so fleshy and succulent that they change
form greatly when dried and thus their true characters often are
very difficult to determine.

Leaves mostly compound, pinnate; calyx much inflated Bryophyllum.

Leaves simple, entire; calyx not at all inflated.

Stamens as many as the calyx lobes; plants very small and slender, the flowers

barely 2 mm. long Tillaea.

Stamens twice as many as the calyx lobes; plants often large, the flowers always
much more than 2 mm. long.

Petals distinct Sedum-

Petals conspicuously united near the base.

Leaves linear, terete or subterete; petals united into a distinct but short

tube Villadia.

Leaves broad, flat, more or less spatulate; petals very shortly connate at
the base. . . .Echeveria.

BRYOPHYLLUM Salisbury

Erect, often large, fleshy, perennial herbs with usually branched stems, the
stems leafy; leaves very thick and succulent, opposite, simple or pinnate; flowers
large, perfect, nutant, in cymes or panicles; sepals 4, united to form a large inflated

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 405

calyx; corolla subcampanulate to urceolate, the limb 4-lobate, the lobes spreading;
stamens 8, biseriate, adnate to the corolla tube, the filaments filiform; disk bear-
ing 4 oblong glands; carpels of the ovary 4, distinct or partially united; ovules
numerous; follicles of the fruit 4.

About 20 species, natives of Madagascar, one of them often
cultivated and naturalized in many tropical regions of the earth.

Bryophyllum pinnatum (Lam.) Kurz, Journ. Asiat. Soc.
Bengal 40, pt. 2: 52. 1871. Cotyledon pinnata Lam. Encycl. 2: 141.
1786. B. calycinum Salisb. Parad. Lond. pi. 3. 1805. Hoja del aire;
Hoja del norte; Flor del aire.

Often planted in gardens; abundantly naturalized in thickets or
on banks at various elevations from the lowlands high into the
mountains, frequently forming large and dense colonies; Alta
Verapaz; Izabal; Zacapa; Chiquimula; Jutiapa; Santa Rosa;
Escuintla; Sacatepe"quez ; Retalhuleu; Quezaltenango; Huehue-
tenango. Mexico; British Honduras; frequently naturalized in
other parts of Central America.

Plants erect, glabrous, succulent, sometimes as much as 1.5 meters high but
usually scarcely half as tall; leaves very thick, 10-30 cm. long, petiolate, simple or
the larger ones pinnate, the few leaflets petiolulate, elliptic to oblong, obtuse or
rounded at each end, coarsely crenate; flowers usually forming large panicles, the
individual flowers pendulous; calyx reddish green, oblong-campanulate, much
inflated, 3-3.5 cm. long; corolla reddish brown, longer than the calyx, the lobes
lanceolate or ovate-lanceolate, acute.

Called "hoja de la vida" in Honduras; "sanalotodo" (Salvador);
"sisalxiu," "tzitzalxiu" (Yucatan, Maya). The plant probably is
used in domestic medicine, since it is planted commonly about
Indian dwellings or in other places where there are few ornamental
plants and since it is not very ornamental, although the flowers are
bizarre and somewhat interesting. They last a long time when cut;
so they often are used for decorating altars and roadside crosses.
One often sees Indian cargadores returning from the coast carrying
bunches of the flowers to the highlands. Below Zunil, plants of this
species were observed growing on the grass thatch of a small dwelling.
They are remarkably tenacious of life, growing for a long time when
withdrawn from the soil and placed upon a wall or in some similar
situation. If a leaf is placed on the soil, young plants are produced
at each of the notches of the margin. The West Indians of Panama
use the leaves to determine the fidelity of their women. The men
place one of the leaves above the door, and if a new plant grows from
each of the notches of the leaf, the virtue of the woman thus tested is
beyond question.

406 FIELDIANA: BOTANY, VOLUME 24

ECHEVERIA De Candolle

Reference: Karl von Poellnitz, Zur Kenntnis der Gattung
Echeveria DC., Repert. Sp. Nov. 39: 193-270. 1936.

Perennial plants, usually herbaceous, rarely suffrutescent below, simple or
branched, usually glabrous, sometimes pubescent; leaves mostly flat but very thick
and succulent, often glaucous, spirally arranged, often forming lax or dense
rosettes, frequently with red margins, generally pointed at the apex; inflorescence
usually a simple, rather lax spike or raceme, sometimes paniculate, frequently
secund; sepals 5, united below, generally unequal, the lobes mostly long and
narrow, erect or spreading; corolla usually conspicuously 5-angulate, very broad
at the base, the petals united near the base, commonly erect but sometimes with
spreading tips, often bright pink or red; stamens 10, five of them attached near
the middle of the petals, the other 5 either free or inserted lower on the corolla,
the anthers oblong; scales of the receptacle large, thick; carpels of the ovary free,
the follicles of the fruit oblong, erect, tapering into the slender styles; seeds
numerous.

About 90 species, most of them in Mexico, a few occurring in
western South America. One other, E. australis Rose, occurs in
Costa Rica. Because of their handsome foliage, these plants long
have been favorites in cultivation and the majority of them have
been introduced into cultivation at one time or another. The
characters in this genus appear to be indefinite and variable, or else
too many species have been recognized. Although rather ample
material is available for study, the division of the Guatemalan
species is unsatisfactory. Poellnitz's key to the species is evidently
worthless, at least in large part, and the following one is perhaps no
more trustworthy. The genus was named for Atanasio Echeverria,
artist of the Sesse" and Mocifio botanical expedition to Mexico and
Guatemala.

Leaves abundantly pubescent E. macrantha.

Leaves glabrous.

Plants acaulescent, even when fully developed.

Inflorescence an elongate spike-like raceme, usually many-flowered; pedicels
short and thick, in age about 3 mm. long; leaves mostly acute.

E. huehueteca.

Inflorescence short, few-flowered, the lower pedicels mostly 4-12 mm. long
and relatively slender; leaves rounded or very obtuse at the apex and

usually apiculate E. Steyermarkii.

Plants with elongate leafy stems, the leaves sometimes collected in rosettes at

the top of the main stem.

Leaves densely crowded at the apex of the stem and forming distinct rosettes.
Peduncles and leafy stems stout; leaves 4-7 cm. long, 2-3 cm. wide, usually

blue- or glaucous-green E. nuda.

Peduncles and leafy stems slender; leaves mostly 2-5 cm. long, 0.5-1.2 cm.

wide, usually dark dull green or purplish red E. Maxonii.

Leaves rather evenly distributed along the stems, not forming a distinct
rosette.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 407

Leaves mostly 2-4.5 cm. long and about half as wide, rounded or very obtuse
at the apex E. guatemalensis.

Leaves mostly 5-10 cm. long, narrow, more than twice as long as wide,
acute or conspicuously pointed at the apex E. Pittieri.

Echeveria guatemalensis Rose, Contr. U. S. Nat. Herb. 12:
395. pi. 47. 1909.

Usually epiphytic on trees in moist or wet forest, 1,500-3,200
meters; endemic; Jalapa; Sacatepe"quez (type from Volcan de Agua,
at 2,700-3,000 meters, W. R. Maxon 3726); Guatemala; Chimal-
tenango; Solola; Totonicapan; Quezaltenango; San Marcos.

Plants erect, sparsely branched, 25 cm. high or less, the branches very thick
and fleshy, sometimes suffrutescent below, the plants glabrous throughout; leaves
not forming rosettes but densely crowded along the upper part of the branches,
the older ones gradually deciduous, spreading, spatulate, mostly 2-4 cm. long and
0.5-1.5 cm. wide, rounded or very obtuse at the apex, grayish green, sometimes
with narrow red margins; scapes 30 cm. long or usually shorter; flowers as many as
20 or even more but sometimes fewer, the pedicels mostly 3-4 mm. long, stout;
bracts 6-8 mm. long; calyx short, the segments linear, narrowed to an obtuse apex;
corolla about 1 cm. long, red or salmon below, yellowish above, the lobes lanceo-
late, acute, somewhat outcurved above; follicles obliquely oblong-ovoid, 8 mm.
long, conspicuously rostrate.

Echeveria huehueteca Standl. & Steyerm. Field Mus. Bot. 23:
159. 1944. GaUinita.

On limestone bluffs or in moist or dry soil, sometimes growing
with Juniperus, 2,000-3,500 meters; endemic; Huehuetenango (type
from Cumbre Papal, on south-facing bluffs between Cuilco and
Ixmoqui, Steyermark 50934; collected also on pine-forested slopes
along Rio Selegua opposite San Sebastian H., and about Tunima).

Plants glabrous, acaulescent, the roots fleshy-fibrous; leaves numerous, form-
ing a rosette, sessile, ascending or spreading, fleshy, oblong-lanceolate to subelliptic
or broadly cuneate-obovate-oblong, 2-5 cm. long, 1-2.5 cm. wide, subacuminate
at the apex to obtuse or rounded and cuspidate-apiculate, very broadly cuneate
at the base, green on both surfaces or sometimes purplish; stems erect, strict,
20-30 cm. tall, simple, remotely leafy, the leaves greatly reduced, lance-linear to
oblong-lanceolate, 18 mm. long or shorter, acute or acuminate, sessile; flowers
laxly racemose, the racemes 7-13 cm. long, remotely 5-11-flowered, the pedicels
very stout, 2-3 mm. long; sepals strongly unequal, broadly linear to oblong-ovate,
obtuse or subacute, carnose, green, subappressed or somewhat spreading, 6-8 mm.
long; corolla vermilion-red, 8-10 mm. long, the petals almost free, narrowly
lanceolate, suberect, slightly excurved at the apex, narrowly long-attenuate,
dorsally carinate; follicles 8 mm. long, gradually attenuate into a subulate beak.

Echeveria macrantha Standl. & Steyerm. Field Mus. Bot. 23:
159. 1944. GaUinita.

408 FIELDIANA: BOTANY, VOLUME 24

Known only from the type, at 2,000-2,200 meters, Jalapa, dry
rocky slopes, Montana Miramundo, at Buena Vista, Steyermark
32808.

Plants erect, suffrutescent, sparsely branched, the branches as much as 1 cm.
thick, leafy only at the tips; leaves densely rosulate at the ends of the branches,
sessile, rounded-cuneate, 3 cm. long, 2-2.8 cm. wide, broadly rounded or sub-
truncate at the apex and very shortly apiculate, very broadly cuneate at the base,
thick-carnose, abundantly but not very densely hirtellous on both surfaces, pale
yellowish green, the margins rose-colored; peduncle stout, 4.5 cm. long, about 3-
flowered, the fruiting pedicels 8 mm. long, thick, densely hirtellous; sepals almost
free, 8 mm. long, lance-oblong, gradually narrowed to the obtuse apex, densely
hirtellous; corolla in age persistent, pubescent outside, 2 cm. long, the petals
narrowly lanceolate, gradually attenuate to the apex, acutely carinate dorsally;
follicles 15 mm. long, sparsely pubescent, the body lance-oblong, 8 mm. long,
attenuate into a slender beak of about the same length.

The species is noteworthy for its exceptionally large flowers and,
of course, among Guatemalan species for its pubescence. Eric
Walther considers it synonymous with E. Pringlei Rose, a species of
Mexico.

Echeveria Maxonii Rose, Contr. U. S. Nat. Herb. 12: 395.
pi. 48. 1909. Siempreviva.

On exposed or shaded rocks or more often epiphytic, 2,200-3,300
meters; endemic; Baja Verapaz (type from Chuacus, W. R. Maxon
3406); El Progreso; Totonicapan; Quezaltenango.

Plants glabrous, caulescent except when very young, erect or decumbent,
80 cm. long or less; leaves forming rosettes at the ends of the stout branches or
often scattered along the branches and spreading, spatulate, mostly 2-5 cm. long,
0.5-1.2 cm. wide, rounded and apiculate to subacute at the apex, broadly cuneate
at the sessile base, green above, green or silvery green beneath, sometimes tinged
with red or purple, the edges often red; peduncles sometimes 50 cm. long but usually
much shorter, the inflorescence racemose or sometimes subpaniculate, few-many-
flowered, short or much elongate; pedicels mostly 4-6 mm. long or in age even
longer; bracts small, about equaling the pedicels; sepals shortly united below,
semiterete, acute or obtuse, unequal, 3-5 mm. long, often spreading in age; corolla
about 1 cm. long, coral-red and often partly bright yellow, the petals erect but
somewhat excurved at the apex.

A common plant in many parts of the Guatemalan mountains.
The material referred here is ample and somewhat variable. Further
study of a greater number of collections may devise a means of
segregating some of the apparently minor forms.

Echeveria nuda Lindl. Gard. Chron. 280. 1856. A species of
central Mexico, represented in Guatemala by the following variety:

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 409

Echeveria nuda var. montana (Rose) Poellnitz, Repert. Sp.
Nov. 39: 224. 1936. E. montana Rose, Bull. N. Y. Bot. Gard. 3: 6.
1903. Gallina (San Marcos).

Usually on shaded cliffs, 1,850-3,400 meters; Solola; Quezalte-
nango; San Marcos; Huehuetenango. Southern Mexico.

Plants glabrous, suffrutescent, branched, sometimes 1.5 meters high, generally
lower, the stems as much as 2.5 cm. thick; leaves forming dense rosettes at the ends
of the branches, broadly obovate to almost orbicular, mostly 4-7 cm. long, 2-3 cm.
wide, sometimes oblong-obovate, rounded or very obtuse at the apex, broadly
cuneate to almost rounded at the base, thick and succulent, bluish green with red
edges; inflorescences usually long-pedunculate, very stout, racemose, not secund,
usually many-flowered, very dense to somewhat remotely flowered, the pedicels
usually very short and thick; sepals fleshy, green, linear or narrowly lanceolate,
about equaling the corolla, usually spreading; corolla 10-12 mm. long, rose-red
to orange-red or coral-red, yellow within.

The inflorescences of this and other species of Echeveria are much
used in the highlands for decorating altars, especially about Christ-
mas time. They are very handsome, brightly colored, and showy,
and of course last for a long time out of water. This plant is or was
considered a good species by Eric Walther, who has given the genus
much study. Poellnitz, who certainly divided the species finely
enough, considers it merely a variety.

Echeveria Pittieri Rose, Contr. U. S. Nat. Herb. 13: 296. 1911.

On rocks or more often epiphytic, usually in shaded situations
but sometimes growing in the open, 1,000-2,400 meters; Zacapa;
Chiquimula (type from lake on Volcan de Ipala, H. Pittier 1880);
El Progreso; Jutiapa; Solola; Huehuetenango.

Plants caulescent, glabrous, usually branched, commonly about 30 cm. high
but varying in size, the branches somewhat woody, very thick and succulent;
leaves forming rosettes at the ends of the branches or regularly scattered along
them, oblanceolate and 3-8 cm. long, variable in shape and size, mostly subacute,
gradually attenuate to the base, green or often tinged with red or purple; peduncles
10-20 cm. long, the bract-like leaves similar to the lower ones of the plant but
smaller; inflorescence not secund, dense and short, about 4-6 cm. long, or often
more open and longer, the flowers usually numerous, on very short, thick pedicels;
corolla salmon-red or sometimes yellowish tinged with red, 12-13 mm. long, the
petals erect but usually with excurved tips; sepals shortly united at the base,
linear, 6-8 mm. long; follicles 5 mm. long, attenuate into the beak-like styles, these
4 mm. long.

Echeveria Steyermarkii Standl. Field Mus. Bot. 23: 160. 1944.

Epiphytic or on rocks, 1,300-3,700 meters; endemic; Zacapa
(type collected between Santa Rosalia de Marmol and San Lorenzo,

410 FIELDIANA: BOTANY, VOLUME 24

f

Steyermark 43145); Solola (volcanoes of Tollman and Santa Clara);
San Marcos (between Sibinal and Ichiguan).

Plants glabrous, acaulescent, solitary or cespitose, the roots fibrous; leaves
usually very numerous and forming a dense rosette, spreading or ascending, green,
sometimes tinged with pink or purple, narrowly or very broadly oblong-spatulate,
2.5-6.5 cm. long, 1-2 cm. wide, rounded or very obtuse at the apex and obtusely
short-apiculate, carnose but not very thick, broadly cuneate at the base; scapes
solitary or few, 5-20 cm. high, the flowers few, short-racemose or subcorymbose:
leaves of the scapes few and inserted near the base or more numerous and continued
to the inflorescence, linear or oblong, the largest 2 cm. long, obtuse, ascending;
flowers 3-10, long-pedicellate, the pedicels slender, mostly 8-15 mm. long, the
bracts oblong or almost linear, much shorter than the pedicels; sepals unequal,
green, fleshy, 5-8 mm. long, oblong or ovate-oblong, obtuse, appressed or some-
what spreading; petals rose-red or vermilion, 8-11 mm. long, lanceolate or narrowly
lanceolate, erect but excurved at the apex, attenuate-acuminate; follicles 7-8 mm.
long, suberect, long-rostrate.

It may be that more than a single species is represented by the
five specimens referred here for they exhibit some variations, but
it is believed that these probably result from varying conditions of
moisture and exposure.

Some of the species of Kalanchoe, natives of tropical Africa and
Asia, are cultivated rarely in Guatemalan gardens for ornament.
In this genus the corolla is gamopetalous and usually has a con-
spicuous, often slender tube.

SEDUM L.

Reference: Harald Froderstrom, The genus Sedum L., Act. Hort.
Gotoburg. 10, App., pp. 6-262, with 115 plates. 1935.

Plants chiefly perennial, succulent, glabrous or nearly so, erect or decumbent,
the stems often branched, generally very leafy; leaves alternate, often imbricate,
entire or dentate, terete to flat; flowers small or rather large, variously colored,
perfect, in terminal cymes, the flowers often secund along the more or less elongate
branches; calyx 4-5-lobate or 4-5-parted; petals 4-5, distinct or barely united at
the very base; stamens 8-10, perigynous, the alternate ones generally attached
to the petals, the filaments filiform or subulate; scales of the receptacle entire or
emarginate; carpels of the ovary 4-5, distinct or united at the base, the styles
slender, usually short; ovules numerous; follicles of the fruit containing few or
numerous seeds.

A large genus, with 125 or more species, widely distributed in
both hemispheres but most numerous in China and Mexico. One
other Central American one is known, in Salvador. In the tropics
the species are confined to mountain regions.

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 411

Leaves linear or oblong, terete or subterete, sometimes compressed but then very

thick, usually broadest at or below the middle.

Flowers sessile; stems smooth, not papillose; plants cultivated or rarely becoming
naturalized near dwellings; flowers yellow S. mexicanum.

Flowers pedicellate, the pedicels rather long or often very short; stems papillose;

native plants.

Flowers pink, conspicuously pedicellate; petals acuminate. . .S. guatemalense.
Flowers yellow, on very short pedicels or appearing sessile; petals obtuse.

Leaves almost linear when dried, terete in the living state; inflorescence
few-many-flowered S. australe.

Leaves oblong, flat or compressed; inflorescence only 2-flowered . . . S. Triteli.
Leaves spatulate or obovate, broadest above the middle, flat.

Plants small and slender, the stems (when dry) about 1 mm. thick; leaves mostly

5-10 mm. long.

Sepals conspicuously calcarate at the base; inflorescence several-few-flowered.

S. Batesii.

Sepals not calcarate at the base; inflorescence 2-flowered S. Triteli.

Plants rather large and stout, the stems thick and stout, when dry mostly 3-6
mm. thick; leaves mostly 2 cm. long or longer.

Sepals 1.2-2.5 mm. long; petals more than twice as long as the sepals, often

3-4 times as long S. praealtum.

Sepals 5-6 mm. long; petals equaling or little longer than the sepals.

S. Millspaughii.

Sedum australe Rose in Britt. & Rose, Bull. N. Y. Bot. Gard.
3: 41. 1903.

Usually on mossy rocks in moist or rather dry forest, sometimes
on exposed or shaded banks, occasionally on limestone, 2,500-4,000
meters; endemic, so far as known, but to be expected in Chiapas;
Quezaltenango (type from Volcan de Santa Maria, E. W. Nelson
3707); San Marcos (volcanoes of Tajumulco and Tacana); Totoni-
capan (above Totonicapan) ; Huehuetenango (Sierra de los Cuchu-
matanes).

Plants perennial, often with very numerous stems and forming large mats or
dense colonies, the stems procumbent and rooting, often somewhat woody below,
rather stout and thick, tuberculate; leaves usually very densely crowded and often
more or less imbricate, semiterete or subterete, mostly 6-8 cm. long, very succulent,
often tinged with rose-red, very obtuse, sessile, glabrous; inflorescence usually
very dense, subcapitate to corymbose, few-many-flowered, the flowers sessile or
nearly so; calyx lobes green, half as long as the petals, not calcarate at the base;
petals about 7 mm. long, yellow, often tinged with red especially in withering,
mucronate; carpels of the fruit usually dark red, spreading in age, long-rostrate.

This is a very common plant in the high mountains of the
Occidente, often growing in alpine meadows but most plentiful on
large rocks. It is not particularly ornamental or handsome but it
sometimes is planted for ornament in gardens of the area where it is
native.

412 FIELDIANA: BOTANY, VOLUME 24

Sedum Batesii Hemsl. Diag. PL Mex. 1: 12. 1878. Altamiranoa
Batesii Rose, N. Amer. Fl. 22: 49. 1905.

Type collected in Guatemala by Bates, the locality not indicated;
reported from Carrizal, Santa Rosa, 1,700 meters; sterile material
collected between Democracia and Santa Ana Huista, Huehuete-
nango, 800-1,000 meters, probably belongs here. Southern Mexico.

Plants probably annual, slender, often much branched from the base, the
slender branches erect or diffusely spreading, 15 cm. long or less, rather sparsely
leafy; leaves oblanceolate or spatulate, mostly 6-12 mm. long, obtuse, narrowed
below into a petioliform base, thin when dried, flat; inflorescence rather laxly
corymbose, or sometimes elongate and few-flowered, the flowers 4-5-parted, sub-
sessile; bracts oblanceolate, obtuse, 2.5 mm. long; sepals calcarate at the base,
oblanceolate or oblong, obtuse, 3-4 mm. long; petals shortly united below, oblong
or lanceolate, obtuse, apparently not mucronate, slightly longer than the sepals,
white; carpels of the fruit united one-third their length, slender, suberect, 4.5
mm. long, 5-6-seeded, the subulate persistent styles elongate.

Sedum guatemalense Hemsl. Diag. PL Mex. 1: 11. 1878.
Colchdn de nino.

Shaded moist banks or most often epiphytic on the trunks or
branches of trees or on logs or stumps, often in forest of Cupressus
or Alnus, 2,000-4,000 meters; endemic; Jalapa; Chimaltenango
(type from "summit above Calderas," on dead stump, Salvin &
Godman 78); Solola; Suchitepe'quez; Quezaltenango; San Marcos;
Totonicapan ; Huehuetenango.

Plants perennial, rather stout, often woody below, the stems simple or
branched, papillose, 15-30 cm. long, procumbent and rooting below or often
pendent from branches; leaves numerous, semiterete or subterete, 5-14 mm. long,
obtuse, narrowed into a petioliform base, usually numerous but not crowded,
spreading, green; inflorescence laxly corymbose, few-many-flowered, the flowers
on long slender papillose pedicels; bracts oblanceolate, obtuse; sepals not calcarate,
linear, obtuse, 3-5 mm. long; petals pink or reddish, free almost to the base, sub-
ovate, subobtuse and mucronate, 5-6 mm. long; carpels of the fruit shortly united
at the base, broad and turgid, with short spreading styles, 5 mm. long; follicles
many-seeded; seeds very long, linear, smooth, 2 mm. long.

This is a very common plant in many parts of the Guatemalan
highlands. We have collected it at the type locality, on the slopes
above Calderas, where it is abundant.

Sedum mexicanum Britton, Bull. N. Y. Bot. Gard. 1: 257.
1899. Colchdn de nino; Chisme.

Perhaps native of Mexico, but the native habitat somewhat
uncertain; planted commonly in Guatemalan parks and gardens,
rarely naturalized, as on roadside banks near Tactic (Alta Verapaz).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 413

Plants perennial, herbaceous, the stems slender, decumbent and rooting, the
aerial branches erect, densely leafy, mostly 10 cm. long or less; leaves alternate or
subverticillate, linear or nearly so, 5-20 mm. long, subcalcarate at the base, obtuse,
terete or subterete; flowers sessile along the slender spreading branches of the
corymb, the bracts lanceolate, subobtuse, 3-4 mm. long; sepals lanceolate, green,
subobtuse, very unequal, calcarate at the base, 3-5 mm. long; petals lanceolate,
5-6 mm. long, subobtuse, short-mucronate, bright yellow; follicles of the fruit
turgid, 5 mm. long.

This is a very common plant in gardens of Central America, at
least in Guatemala and Salvador, but there is no reason for believing
that it is native anywhere in the region. It is often used as a border
plant about flower beds, and sometimes it is planted to make formal
designs. At Coban plants were seen growing on a tile roof.

Sedum Millspaughii Hamet, Field Mus. Bot. 2: 378. 1913.

Known only from the type, Lago de Amatitlan, Guatemala,
1,200 meters, W. A. Kellerman 6559.

Apparently a large and coarse, branched perennial, erect, the branches succu-
lent, about 4 mm. thick; leaves flat, obovate, very obtuse or subacute, contracted
below and narrowed into a petioliform base, about 2.5 cm. long; inflorescence
laxly corymbose, the flowers numerous, short-pedicellate; sepals not calcarate,
oblong, subacute or obtuse, almost equaling the petals; petals probably white,
obovate, obtuse and aristate-mucronate, 5-6.7 mm. long; carpels of the fruit erect,
2.5 mm. long, long-rostrate.

The species is known only from very poor material, much broken
probably as a result of transport by foreign mail. It is almost certain
that a synonym of this is S. salvadorense Standl. Journ. Wash. Acad.
Sci. 13 : 438. 1923, the type of which was collected at Finca Colima,
Sierra de Apaneca, Salvador.

Sedum praealtum DC. PI. Rar. Geneve 10: 21. 1847. Siempre-
viva; Santa Polonia (Coban).

Open or wooded mountain sides, in moist or rather dry situations,
2,300-3,000 meters; Sacatepe"quez; Solola; Quezaltenango; Quiche";
often cultivated in other regions, as in Alta Verapaz. Mexico.

Plants erect, coarse, often much branched, a meter high or less, the branches
very thick and succulent, often woody below; leaves flat, spatulate or obovate,
mostly 1.5-4 cm. long, rounded at the apex, very thick and fleshy, green, narrowed
below into a petioliform base; inflorescence corymbose or subpaniculate, many-
flowered, often 6-10 cm. long and 4-6 cm. broad, the numerous flowers sessile or on
short thick pedicels; sepals short and broad, not calcarate, 1.2-2.5 mm. long;
petals bright yellow, free almost to the base, lanceolate to oblong, subobtuse,
short-mucronate, 5-8 mm. long; stamens slightly shorter than the petals; carpels
of the fruit erect or spreading, 5-6 mm. long; seeds ovoid, 1 mm. long.

414 FIELDIANA: BOTANY, VOLUME 24

This plant often is grown in pots in patios, and also planted in the
ground. It has been reported from Guatemala as S. dendroideum
Mocino & Sesse", a Mexican species from which it is perhaps not
distinct. About Coban the juice of the leaves is employed frequently
for treating inflammation of the eyes and mouth, particularly the
mouth affection called fuego de la boca.

Sedum Triteli Hamet, Field Mus. Bot. 2: 379. 1913.

Known only from the type, collected in the "Sierra Madre" of
Guatemala at 400 meters, F. C. Lehmann 1528.

Plants perennial, with fibrous roots, the stems subrepent, the tips erect, gla-
brous and papillose; leaves alternate, sessile, obovate or almost linear, 5-7 mm. long,
obtuse; inflorescence 2-flowered, the pedicels very short; calyx lobes 5, not cal-
carate, linear^deltoid, about 5 mm. long; petals yellow, longer than the calyx,
about 6 mm. long, subobtuse; follicles of the fruit few-seeded, divergent, not
gibbous, the styles 2 mm. long.

We have seen no material of the species. It is uncertain in what
part of Guatemala it may have been collected, and it is not apparent
what may be meant by the Sierra Madre. The elevation is a remark-
ably low one for a plant of this genus.

Semper vivum lector um L., native of Europe, known in the United
States as "hen-and-chickens," often is grown in Guatemala in
gardens or pots, being known as "gallina con polios," "rosa verde,"
and "conchita." It is an acaulescent plant with a basal rosette of
leaves, producing short lateral stolons that end in globose cabbage-
like heads of small pale succulent leaves.

TILLAEA L.

Very small, branched herbs, mostly annuals, aquatic or terrestrial, commonly
glabrous, slightly succulent; leaves very small, opposite, terete or flat, entire;
flowers minute, axillary, solitary or cymose-aggregate, white or reddish, often
fasciculate; calyx 3-5-lobate or 3-5-parted; petals 3-5, free or connate at the very
base; stamens 3-5, the filaments filiform, the anthers didymous; scales of the
receptacle 3-5 and linear, or none; carpels of the ovary 3-5, free, attenuate to the
short subulate styles, the stigmas minute; ovules 1 or more in each carpel; follicles
many-seeded or rarely 1-seeded.

About 20 species, widely distributed in both hemispheres. Only
the following is known from Central America. By Berger the genus
is united with Crassula, but most American and other botanists
have treated it as a distinct group.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 415

Tillaea connata Ruiz & Pavon, Fl. Peruv. 1: 70. pi. 106, /. a.
1798.

Collected in Guatemala but once; Quezaltenango, near Zunil,
on a dry mud wall, the plants withered, 2,280 meters, Standley
83194. Mexico; western South America.

Plants small, sometimes 9 cm. long but usually much smaller, erect or pros-
trate, much branched; leaves lanceolate, connate at the base, acute, 1-4 mm. long;
flowers usually clustered in the leaf axils, sometimes solitary, on short slender
pedicels or sessile; calyx lobes oblong-lanceolate or ovate-lanceolate, acutely
acuminate, 2 mm. long; petals similar to the calyx segments but shorter; seeds 1-2.

Perhaps the plant is more widely dispersed in western Guate-
mala, but the plants are so small that they are easily overlooked,
and probably they grow only during the wet months.

VILLADIA Rose

Perennial herbs, the roots fleshy or somewhat tuberous, the stems elongate,
leafy, simple or branched; leaves alternate, short, terete or semiterete, turgid;
flowers small, white, reddish, or orange, forming an elongate equilateral raceme or
spike or a very narrow and compact panicle; flowers 5-parted, the sepals subequal;
corolla terete, the segments thin, united below into a short but distinct tube, the
lobes more or less campanulate-connivent; stamens 10, the anthers short and
broad; scales of the receptacle thin, conspicuous; carpels erect, the styles very
short.

About 25 species, in Mexico and western South America. Only
one is found in Central America.

Villadia guatemalensis Rose, Contr. U. S. Nat. Herb. 12: 396.
1909. Altamiranoa guatemalensis Walther, Cact. & Succ. Journ. 10:
24. 1938.

Known only from the type, near Chuacus, near Salama, Baja
Verapaz, W. R. Maxon 3411; plants were grown and flowered in
Washington in the United States.

Plants much branched, spreading, usually procumbent, the flowering branches
erect or ascending; leaves close together, divergent at almost a right angle, pale
yellowish green, terete, 1.5-2 cm. long, pointed; flowers few, the upper ones ter-
minal, the others from the leaf axils, all sessile; sepals ovate, green, almost free;
corolla lemon-yellow, the tube short but definite, the lobes spreading; styles
slender; carpels erect, even in age.

We have seen no material of this species. We have done little
collecting in Baja Verapaz, except on its borders, and this plant
quite probably is a strictly localized species.

416 FIELDIANA: BOTANY, VOLUME 24

SAXIFRAGACEAE. Saxifrage Family

Mostly perennial herbs, shrubs, or small trees, very varied as to habit, some-
times armed with spines or prickles; leaves alternate, rarely opposite, without
stipules, various in form but usually simple; flowers perfect, usually small or
medium-sized; hypanthium usually well developed and more or less adnate to the
base of the gynoecium; sepals normally 5; petals as many as the sepals or none;
stamens as many or twice as many as the sepals, borne on the edge or inner surface
of the hypanthium, rarely numerous, the filaments subulate or clavate, sometimes
broad and bidentate; anthers subglobose or narrow, 2-valvate, with lateral or
inner valves; carpels usually fewer than the sepals and more or less united, some-
times free or of the same number as the sepals, the placentae parietal, axial, or
basal; ovary partly or wholly inferior; styles as many as the carpels, free; fruit
capsular or follicular, rarely baccate.

About 75 genera, the species most numerous in temperate or
arctic regions, those of the tropics mostly somewhat abnormal and
perhaps to be referred to distinct families, as they have been placed
by some authors. Only the following genera are known in Central
America. They are ill-assorted groups diverse in character and hav-
ing little in common.

Leaves palmate-nerved, usually cordate at the base; plants sometimes armed with

prickles; fruit baccate Ribes.

Leaves penninerved or triplinerved, not cordate at the base; plants unarmed.
Flowers borne on the upper surface of the leaf near its apex; fruit baccate.

Phy llonoma.
Flowers arising from the branches; fruit dry.

Petals valvate, very small; flowers in corymbs; stamens 8 Hydrangea.

Petals convolute or imbricate, large; flowers mostly solitary in the leaf axils
or in small cymes; stamens 25 or more Philadelphus.

Deutzia gracilis Sieb. & Zucc., native of Japan, is in cultivation in
Coban, Alta Verapaz, but is very rare in Central America, and we
have not seen it elsewhere. It is a shrub about a meter high with
oblong-lanceolate, stellate-pubescent, sharply serrate leaves and
small, pure white flowers. It is not uncommon in cultivation in the
United States but is not a very attractive shrub, and scarcely merits
cultivation when there are so many other better ornamental plants.

Escallonia floribunda HBK. is sometimes cultivated about Guate-
mala City, as in the Jardin Botanico and La Aurora Park. It is
native in Costa Rica and northwestern South America. It is an
almost glabrous shrub or small tree with entire, coriaceous, narrowly
lance-oblong leaves 4-9 cm. long, the small white flowers in dense
terminal panicles.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 417

HYDRANGEA L.

Trees, shrubs, or woody vines, often epiphytic; leaves opposite, deciduous or
persistent, entire, dentate, or rarely lobate, the pubescence often of branched
hairs; flowers perfect, small, or the marginal ones sterile and much enlarged,
usually white or pink, in corymbose or paniculate cymes; sepals 4-5, minute, in
the sterile flowers large and petaloid; petals 4-5, valvate; stamens 8-10, the fila-
ments filiform or nearly so, the anthers didymous; ovary inferior, completely or
partially 3-4-celled; styles subulate or the stigmas sessile; ovules numerous, on
axial placentae; capsule 2-4-celled, small, membranaceous or coriaceous, opening
at the apex between the stigmas; seeds numerous, minute.

About 40 species, in North and South America and in eastern
Asia south to Java. Three other species are known from Costa
Rica and Panama. Four species, some of them with rather showy
flowers, are native in the eastern United States.

Plants scandent and epiphytic, native; pubescence of stellate hairs; inflorescence
in bud surrounded by an involucre of 4 large membranaceous bracts.

H. Steyermarkii.

Plants erect shrubs, the sparse pubescence of simple hairs; inflorescence not
bracteate H. macrophylla.

Hydrangea macrophylla (Thunb.) DC. Prodr. 4: 15. 1830.
Viburnum macrophyllum Thunb. Fl. Japon. 125. 1784. Hortensia
opuloides Lam. Encycl. 3: 136. 1789. Hydrangea opuloides K. Koch,
Dendrol. 1 : 353. 1869. H. Hortensia Sieb. Act. Acad. Leop. Carol. 14,
pt. 2: 688. 1829. Hortensia.

Native of Japan, but widely cultivated for ornament; common
in gardens of Guatemala; rarely naturalized, but found in wet thicket
near a stream along Rio Tacana above San Antonio, San Marcos,
2,450 meters.

Usually a shrub of 1-2 meters, almost glabrous throughout; leaves short-
petiolate, thick, obovate to elliptic or broadly ovate, 7-15 cm. long, abruptly
short-acuminate, cuneate at the base, paler beneath, coarsely serrate; corymbs
pedunculate, sparsely appressed-pilose, globose and 15-20 cm. broad or larger;
flowers blue, pink, or white.

The hortensia is one of the favorite ornamental plants of Guate-
mala and other portions of Central America, and is planted at various
elevations but especially in the mountains. There is even a caserio
called Las Hortensias in the Department of Guatemala. The shrub
grows well in the open and also may be seen as a pot plant in patios
where the climate is severe. Particularly famous is the large planting
of thrifty bushes at San Rafael on the road between Guatemala and
Antigua. This place, now maintained as a resort hotel or roadhouse,
has been a famous stopping place for travelers for many decades,

418 FIELDIANA: BOTANY, VOLUME 24

especially in times of the old diligencias. This hydrangea is much
grown as a pot plant in the colder parts of the United States, or out
of doors where the climate is sufficiently mild.

Hydrangea Steyermarkii Standl. Field Mus. Bot. 22: 233. 1940.

Wet mixed forest, 1,500-2,800 meters; Alta Verapaz; Suchite-
p£quez; Huehuetenango; San Marcos (type from Quebrada Canjula,
between Sibinal and Canjula, Volcan de Tacana, Steyermark 36044).
Known certainly only from Guatemala but probably extending into
Mexico, perhaps to Veracruz.

A large scandent epiphytic shrub, the stems ferruginous, abundantly furnished
with linear scale-like trichomes and also laxly stellate-tomentose; leaves on petioles
1-1.5 cm. long, coriaceous, cuneate-obovate to obovate-oblong, 6.5-17 cm. long,
3.5-9 cm. wide, rounded or very obtuse at the apex and often apiculate, cuneately
narrowed to the base, the margin closely or remotely denticulate, glabrous above,
at least when mature, paler beneath, tomentose at first but soon glabrate, the
lateral nerves about 9 pairs; inflorescences mostly axillary, about 4.5 cm. high
and 9 cm. broad, with 5-8 rays, dense and many-flowered, stellate-pilose with
mostly brownish and stipitate hairs, the basal bracts 2 cm. long, densely brown-
tomentose, the flowers sessile or short-pedicellate, secund; hypanthium semi-
globose, 3 mm. broad, glabrous; sepals semiorbicular; petals caducous.

In this species there are none of the showy sterile flowers that are
found in some of the species of Costa Rica and Panama. The tropi-
cal American hydrangeas are very unlike the United States species in
habit, being scandent epiphytes that often climb to the tops of tall
trees, where they produce their flowers, usually high above the ground.
H. Oerstedii Briq. of Costa Rica has very large inflorescences with
many large, bright pink sterile flowers, and is perhaps quite as hand-
some as any of the well-known cultivated species. The large mature
plants of H. Steyermarkii are hard to locate but the juvenile plants
are plentiful on low tree trunks or even on fence posts in the forests
of San Marcos. They are slender vines with almost thread-like
stems that creep closely on the bark by their many delicate roots.
Their leaves are small, usually only 1.5-3 cm. long. Unless one
knows the true nature of these juvenile plants, one is likely to be
puzzled as to their relationship and spend much time searching for
their flowers. Their true relationship, however, is exposed when
the stellate hairs are observed.

PHILADELPHUS L.

Slender shrubs, the stems arching or sometimes scandent; leaves small, oppo-
site, dentate or entire; flowers white, often fragrant, showy, solitary or in 3-9-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 419

flowered cymes at the ends of the branches; sepals 4, valvate, persistent, usually
tomentulose within; petals 4, convolute, rounded or retuse at the apex; stamens
commonly 25-60, the filaments flat, subulate, more or less united at the base,
the anthers short, didymous; ovary more than half inferior, 4-celled; styles elon-
gate and equaling the stamens or short, nearly distinct or united to the apex;
ovules numerous, pendulous, multiseriate, imbricate; capsule obovoid, ligneous
or coriaceous, loculicidal; seeds numerous, the testa membranaceous, reticulate;
endosperm carnose.

About 40 species, in North America, southern Europe, Caucasus,
and eastern Asia. Only the following are known in Central America.
Many of the species, known in the United States as "mock-orange"
or "syringa," have been introduced into cultivation because of their
handsome, usually fragrant flowers, produced in the North in spring
or early summer.

Petals glabrous within, glabrous outside or sparsely pilose near the base and along

the median line P. mexicanus.

Petals copiously villous or pilose on both surfaces P. myrtoides.

Philadelphia mexicanus Schlecht. Linnaea 13: 418. 1839.
P. Matudai Lundell, Contr. Univ. Mich. Herb. 4: 6. 1940 (type from
Volcan de Tacana, Chiapas, 1,400 meters, Matuda 2916). Mosqueta.

Cultivated commonly for ornament in the mountains of Guate-
mala, and probably wild in San Marcos; often becoming naturalized
about plantations. Central and southern Mexico.

A tall slender arching shrub, often forming dense, much interlaced thickets,
the bark brownish or grayish, the branchlets strigose; leaves on short slender
petioles, lance-ovate or ovate, 3-7 cm. long, long-acuminate, rounded at the base,
triplinerved, remotely denticulate or subentire, strigose, sometimes glabrate
above, pale beneath; branchlets bearing usually 1-5 or sometimes more fragrant
flowers; hypanthium and calyx whitish-strigose, the sepals lance-ovate, 8-10 mm.
long, acute or acuminate; petals obovate, 12-15 mm. long, glabrous within,
sparsely pilose or villous outside near the base; capsule rounded-obovoid, 1 cm.
long.

Cultivated commonly in the cooler parts of Central America,
often to form dense hedges, and known everywhere by the name
"mosqueta." It has given its name to La Mosqueta, a caserio in the
Department of Quezaltenango. Most if not all the cultivated plants
of Guatemala and other parts of Central America seem to belong
to this species, which probably was imported from Mexico. It is
strange that the equally showy, indigenous P. myrtoides has not
been planted in local gardens, but probably it is disdained because
it is a wild plant.

420 FIELDIANA: BOTANY, VOLUME 24

Philadelphia myrtoides Bertol. Fl. Guat. 421. 1840. P. tri-
chopetalus Koern. ex Regel, Gartenflora 16: 73. 1867. Mosqueta.

Moist or wet thickets or mixed forest, 1,500-3,000 meters;
Guatemala; Sacatepe"quez (type from Volcan de Agua, Velasquez);
Solola; Quich^ ; Huehuetenango; Quezaltenango; San Marcos.
Probably extending into Chiapas; reported from Costa Rica, but
not native there.

An arching shrub or often climbing over shrubs and trees, the young branches
strigose; leaves short-petiolate, ovate or elliptic-ovate to lance-ovate, 3-9 cm.
long, acuminate, rounded at the base, remotely denticulate, 3-5-plinerved, hirsute-
strigose, green above, pale beneath; flowering branchlets bearing usually 1-3
flowers, these pedicellate; hypanthium and calyx grayish-strigose, the sepals
ovate, acuminate, 10-15 mm. long, acuminate; petals broadly obovate or sub-
orbicular, 2 cm. long, finely villous on both surfaces; capsule 1 cm. long.

This has handsome large flowers, but in the wild state, at least,
they are not produced in large numbers. P. myrtoides is too closely
related to P. mexicanus, of which probably it is only a form or
variety.

PHYLLONOMA Willdenow

Slender shrubs or trees; leaves alternate, simple, denticulate or subentire,
petiolate; flowers small, in small cymes arising from the upper surface of the leaf
blade near its apex; hypanthium turbinate; epigynous disk broad, covering the
bases of the petals and filaments; sepals 5, persistent; petals 5, valvate, acute;
stamens 5, inserted with the petals, the filaments subulate, recurved ; ovary 2-car-
pellate, 1-celled, with 2 parietal placentae, the ovules 2-seriate; stigmas 2, sessile,
recurved; fruit small, baccate, 1-celled, few-seeded; seeds oblong, obtuse at each
end, the testa coriaceous; embryo small, straight, the endosperm copious.

About 7 species, or perhaps fewer, ranging from southern Mexico
to Peru. Three species not enumerated here have been recorded or
described from Costa Rica. The species are separated by rather
"feeble" characters, and it is doubtful how many more than one
can be maintained.

Leaf blades narrowly oblong-lanceolate, subentire or remotely and inconspicuously

denticulate P. cacuminis.

Leaf blades mostly broadly elliptic to lanceolate, usually conspicuously serrate.

P. laticuspis.

Phyllonoma cacuminis Standl. & Steyerm. Field Mus. Bot.
22: 334. 1940.

Wet mixed forest, 2,000-2,600 meters; endemic; Zacapa (Sierra
de las Minas; type from Quebrada Alejandria, summit of Sierra de
las Minas, 2,500 meters, Steyermark 29870).

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 421

A glabrous tree, sometimes 14 meters high, the branchlets very slender,
subangulate and striate; leaves on petioles 6-8 mm. long, narrowly oblong-lanceo-
late, 5-8 cm. long, 1.3-2.2 cm. wide, rather abruptly and narrowly long-caudate-
acuminate, the acumen 1.5 cm. long or less, often falcate, acute at the base, lustrous
above, the margin entire or remotely and minutely serrulate; inflorescence arising
at the base of the acumen, small and few-flowered, the pedicels 1 mm. long;
hypanthium 1 mm. long, the sepals triangular-ovate, obtuse, reflexed.

This may be only a form of P. latiscuspis, but it seems to be as
distinct or well marked as most species that have been proposed in
the genus.

Phyllonoma laticuspis (Turcz.) Engler in Engl. & Prantl,
Pflanzenfam. 3, 2a: 87. 1890. Dulongia laticuspis Turcz. Bull. Soc.
Nat. Moscou 31, pt. 1: 454. 1858. Cerecillo, Uvillo (fide Aguilar).

Wet mixed forest, 1,800-2,800 meters; Zacapa; Quiche"; Que-
zaltenango; San Marcos. Southern Mexico.

A slender tree 4-9 meters tall, usually with few branches, glabrous throughout;
leaves subcoriaceous, on slender petioles 5-13 mm. long; leaf blades broadly
elliptic to lanceolate, mostly 4-8 cm. long and 1-2.5 cm. wide, abruptly contracted
at the apex and prolonged into a linear acumen often 2.5 cm. long, usually obtuse
or rounded at the base, coarsely and often closely or sometimes remotely and
inconspicuously dentate or serrate, bright green and lustrous above, slightly
paler beneath; inflorescence small, arising at the base of the acumen, the pedicels
1-2 mm. long; flowers purple-brown or greenish, the ovate petals 1 mm. long;
stamens shorter than the petals; berry subglobose, 3-4 mm. broad, white.

This species is occasional in forests of the Occidente, but the
tree is an inconspicuous one and likely to be overlooked unless
the strange inflorescences are noted. These are most remarkable,
appearing as they do on the upper side of the leaf blade, where it
is contracted and produced into the long acumen. No other plant
of Central America has similar inflorescences, although the same
arrangement is found in certain South American and African groups
of other families.

RIBES L.

Shrubs, unarmed or prickly; leaves alternate, petiolate, simple, usually pal-
mate-lobate, folded or rarely convolute in bud, without stipules; flowers small or
rather large, perfect or dioecious, 5-merous, in few-many-flowered racemes;
hypanthium cylindric to rotate, often colored, like the sepals; petals mostly smaller
than the sepals; stamens shorter or longer than the sepals; ovary inferior, 1-celled,
many-ovulate; styles 2, more or less connate; fruit baccate, juicy, usually many-
seeded; seeds with endosperm, the embryo minute, terete.

About 150 species in colder and temperate regions of the northern
hemisphere, occurring in tropical America only in the higher moun-

422 FIELDIANA: BOTANY, VOLUME 24

tains, extending southward to Patagonia. Some of the species are
cultivated for their showy flowers. To this genus belong the currants
and gooseberries cultivated extensively for their fruits in the United
States and Europe. They are not grown anywhere in Central
America, as far as we know. Only the following species have been
found wild in Central America.

Plants unarmed R. ciliatum.

Plants abundantly armed with sharp stout spine-like prickles . . . R. microphyllum.

Ribes ciliatum Humb. & Bonpl. ex Roem. & Schult. Syst. Veg.
5: 500. 1819.

Damp or moist thickets or forest, often in Pinus-Abies forest,
sometimes on limestone in Juniperus forest, 3,000-3,800 meters;
Totonicapan; Huehuetenango; Quezaltenango; San Marcos. Cen-
tral and southern Mexico; Costa Rica.

A slender shrub 2-5 meters high, sparsely branched, the young branches
glandular-pubescent; leaves slender-petiolate, suborbicular in outline, mostly
3-6 cm. wide, deeply cordate at the base, 3-5-lobate and crenate-dentate, green
and glabrate above, the venation impressed, paler beneath and glandular-pubes-
cent, the lobes obtuse or subacute; racemes as long as the leaves, about 10-flowered,
finely pubescent and glandular, the pedicels 4-7 mm. long, bracteate; hypanthium
pubescent, short-cylindric, 4-5 mm. long, green or whitish; sepals oblong, sub-
obtuse, greenish white, 3-4 mm. long; ovary glabrous; berry globose, 8 mm. in
diameter, black at maturity.

The ripe fruit is sweet and edible but it is produced in only small
amounts and, as far as we know, is not eaten locally. The shrub is
not plentiful in Guatemala but occurs here and there in the highest
coniferous forests, usually in dense shade. At Desconsuelo the
bushes shed their leaves during the cold months, the young, bright
green leaves appearing at the end of January. This species belongs
to the currant group.

Ribes microphyllum HBK. Nov. Gen. & Sp. 6: 62. 1823.

Rocky hillsides under pine trees, 3,300 meters; Huehuetenango
(Sierra de los Cuchumatanes, between the first cumbre and La
Pradera, Standley 81148a). Central and southern Mexico.

A shrub 2 meters high or lower, the branchlets villous, armed with abundant
stout spine-like prickles; leaves slender-petiolate, ovate-orbicular to reniform-
orbicular, 2.5 cm. wide or smaller, 3-5-lobate and incised-dentate, somewhat
pubescent on both surfaces; bracts broad, membranous, glandular and pubescent,
longer than the very short pedicels; ovary glabrous; hypanthium cylindric, yellow
or reddish, pubescent and glandular, 6 mm. long; sepals lanceolate, about equaling

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 423

the hypanthium; petals obovate or spatulate, retuse, shorter than the sepals;
stamens about equaling the petals; berry globose, glabrous, 8 mm. in diameter.

The specific determination of the small sterile specimen is very
uncertain, and it may well be that the Guatemalan plant is an
undescribed species. A single small shrub was found near the road
across the Cuchumatanes, and prolonged search failed to discover
more plants, nor was it found in the extensive explorations of the
junior author. This is the only record of a wild gooseberry in Central
America, and represents a great extension of range for the subgenus
(Grossularia) from its previously known range in southern Mexico.

Saxifraga sarmentosa L., native of eastern Asia, often is grown
as a house plant in Guatemala, generally in hanging baskets. We
have noted it at Quezaltenango, Momostenango, Coban, and numer-
ous places in the central region. It is sometimes seen in the United
States, where it is called "beefsteak geranium." It is a herbaceous
perennial with a cluster of basal leaves and produces long, slender,
pendent stolons. The succulent leaves are hairy, rounded-reniform,
crenate-dentate, green above and reddish or purplish beneath. The
small white flowers are borne in lax panicles.

There are in cultivation in Guatemala City and perhaps else-
where two species of Pittosporum, plants native in eastern Asia and
Australia and belonging to the family Pittosporaceae. They are
shrubs or trees with entire, alternate or apparently verticillate,
rather thick, exstipulate leaves. The regular 5-parted flowers are
white or cream-colored and arranged in umbels or cymes, these
terminal or borne in the axils of the leaves. P. Tobira (Thunb.)
Ait. has very obtuse or rounded leaves; P. undulatum Vent., acute
or acuminate leaves.

BRUNELLIAGEAE

Trees, sometimes armed with prickles, usually tomentose; leaves opposite
or ternate, with stipules, simple, 3-foliolate, or unequally pinnate, the leaflets more
or less coriaceous, entire or serrate; flowers small, dioecious, in axillary or terminal
panicles, bracteate; calyx 4-5-parted, the lobes valvate; corolla none; disk
depressed, hirsute, adnate to the calyx, 8-10-lobate; staminate flower with 8-10
stamens inserted at the base of the disk; carpels 4-5 in the pistillate flower, dis-
tinct, sessile, hirsute, 1-celled; styles subulate, recurved, the stigmas simple;
ovules geminate, collateral; capsules 4-5 or by abortion fewer, spreading, bival-
vate, 1-2-seeded, the endocarp cartilaginous, separating from the exocarp; testa

424 FIELDIANA: BOTANY, VOLUME 24

of the seed crustaceous, the endosperm fleshy; cotyledons plane, ovate, the radicle
superior.

The family consists of a single genus, with the characters of the
family. About 15 species are known, in the mountains of tropical
America. One other Central American species occurs in Costa
Rica.

BRUNELLIA Ruiz & Pavon

Brunellia mexicana Standl. Journ. Wash. Acad. Sci. 17: 166.
1927. Cedrillo; Huel bianco (Huehuetenango) .

Moist or wet, mixed forest, 1,300-3,000 meters; Zacapa; Chiqui-
mula; El Progreso; Huehuetenango; San Marcos. Southern Mexico.

A small tree 6 meters high or more, or sometimes as much as 30 meters high,
with thick branches, the young branchlets glabrous; leaves large, the leaflets
11-17, short-petiolulate, oblong or lance-oblong, 6-14 cm. long, 2-4.5 cm. wide,
acuminate or long-acuminate, somewhat oblique at the base and rounded or obtuse,
appressed-serrulate, deep green above, glabrate, pale beneath or when young
usually glaucous, velutinous-pilose beneath when young, sometimes glabrous or
glabrate in age, the rachis often tinged with red or rose; panicles about 15 cm.
broad, dense and many-flowered, long-pedunculate, the branches densely tomen-
tose, the pedicels 4-7 mm. long; calyx lobes oblong-ovate, 2.5 mm. long, tomentu-
lose; carpels of the fruit 4-5, compressed, 5 mm. long, densely tomentose and
hispidulous; seeds 2 mm. long, dark brown.

The material from Guatemala exhibits considerable variation in
pubescence, but apparently it is all referable to a single species. The
wood in this genus is pale brown throughout, odorless and tasteless,
light and soft to moderately so, rather fine-textured, easy to work;
not durable.

CUNONIACEAE

Shrubs or large trees; leaves opposite or sometimes verticillate, simple or
compound, with stipules; flowers small, generally perfect, sometimes polygamous
or dioecious, variously arranged; hypanthium with a hypogynous or perigynous
disk within; sepals 4-5; petals 4-5, usually no larger than the sepals; stamens as
many as the sepals, twice as many, or more numerous, inserted under the margin
of the disk, the filaments filiform, longer than or equaling the petals, the anthers
short, 2-celled; carpels of the ovary usually 2 and united, sometimes distinct;
ovules commonly numerous and biseriate on the placenta; fruit capsular or follic-
ular; seeds several or numerous in each cell, often winged; embryo small; endo-
sperm present, the cotyledons flat or convex.

About 20 genera, chiefly in tropical mountains of both hemi-
spheres. One other genus is known from North America, Lyono-
thamnus, in California.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 425

WEINMANNIA L.

Shrubs or often large trees, pubescent or almost glabrous; leaves opposite,
odd-pinnate or rarely simple, the rachis often winged between the leaflets; stipules
deciduous; inflorescence racemose, simple or paniculate, terminal or axillary, the
flowers often glomerate, small, perfect or polygamo-dioecious; sepals 4-5, imbri-
cate; stamens 8 or 10, the anthers small, didymous; petals white; ovary superior,
2-celled or rarely 3-celled, the styles subulate, persistent, the stigma simple;
ovules several or numerous, biseriate, pendulous; disk hypogynous, thick, cyathi-
form, 8-10-angulate or 8-10-lobate; capsule small, 2-celled, septicidally bivalvate,
containing few or many seeds; seeds oblong to reniform, usually pilose, with thin
testa.

About 125 species in America, Australia, New Zealand, and the
African and Pacific islands. One other Central American species
occurs in Costa Rica. Only three species are known from North
America.

Leaves pinnate W. pinnata.

Leaves, except sometimes on sterile branches, simple W. Tuerckheimii.

Weinmannia pinnata L. Syst. Nat. ed. 10. 1005. 1759.

Wet, mixed, mountain forest, 2,000-3,100 meters; Zacapa; Hue-
huetenango; San Marcos. Southern Mexico; Costa Rica and
Panama; West Indies; South America.

Usually a small or medium-sized tree but sometimes tall, the branchlets
densely short-pilose or glabrate; leaves small, pinnate, short-petiolate, the rachis
broadly winged; leaflets usually 9-17, oblong to ovate or obovate, 1-2 cm. long,
obtuse, acute at the base, glabrous above, beneath short-pilose or glabrate; racemes
mostly longer than the leaves, up to 8 cm. long, dense and many-flowered, the
pedicels fasciculate, much longer than the calyx; sepals ovate or lance-ovate, acute,
about 1 mm. long; petals obovate, longer than the sepals; stamens 3 times as long
as the sepals; capsule ovoid, glabrous.

In Guatemala this tree is local in distribution, but in Costa Rica
the species is a characteristic and often dominant tree of the wet
forests of the higher mountains.

Weinmannia Tuerckheimii Engler, Pflanzenfam. ed. 2. 18a:
252. 1930.

Wet mixed forest, 1,450-3,000 meters; endemic; type collected
in Alta Verapaz by Tuerckheim, the exact locality not indicated;
Alta Verapaz (swamp near Tactic); El Progreso; Zacapa (Sierra de
las Minas) ; Huehuetenango.

A glabrous tree 5-9 meters high with a narrow crown, rather densely branched,
the internodes short; leaves coriaceous, short-petiolate, elliptic or ovate-elliptic,
2.5-6 cm. long, 1.5-3 cm. wide, narrowed to an obtuse apex, acute at the base,

426 FIELDIANA: BOTANY, VOLUME 24

rather coarsely crenate-serrate; leaves of vigorous young shoots pinnate, with
3-5 leaflets, each leaflet similar to an adult leaf.

No proper description of the species has been published, the
name appearing originally in a running key to the species of the
genus. The junior author found the tree at two or more localities
in the Sierra de las Minas, where evidently it is more plentiful than
in the Tactic region. In this genus the heartwood is brownish to
light reddish brown, merging gradually into the lighter-colored
sap wood; odorless and tasteless; rather light in weight and firm to
moderately heavy and hard; rather fine in texture and uniform; of
fair durability.

HAMAMELIDACEAE. Witch-hazel Family

Trees or shrubs, the pubescence often stellate; leaves mostly alternate,
deciduous or persistent, simple, pinnately or palmately nerved, the teeth some-
times gland-tipped; stipules usually in pairs, often persistent; flowers small,
actinomorphic or zygomorphic, often capitate; calyx tube more or less adnate to the
ovary, the lobes imbricate or valvate; petals 4 or more, rarely none, perigynous or
epigynous, imbricate or valvate; stamens 4 or more, perigynous, 1-seriate, the
filaments free; anthers oblong, 2-celled, opening by longitudinal valves, the con-
nective often produced; disk none, or annular or of distinct glands between the
stamens and ovary; ovary inferior or nearly so, rarely superior, 2-carpellate, the
carpels often free at the apex; styles subulate, free, often recurved; ovules 1 or
more in each cell, pendulous from axile placentae; fruit a ligneous capsule; seeds
various, the endosperm thin, carnose, the embryo straight.

Genera about 20, in Asia, Africa, and North America, chiefly in
temperate regions. Only the following are known in Central America.

Leaves palmate-lobate; flowers in large long-pedunculate globose heads.

Liquidambar.

Leaves not lobate, entire or nearly so, penninerved or 3-nerved; flowers not in
globose heads.

Leaves triplinerved; stamens 20-24 Matudaea.

Leaves penninerved; stamens 2-8 Distylium.

DISTYLIUM Siebold & Zuccarini

Reference: E. H. Walker, A revision of Distylium and Sycopsis,
Journ. Arnold Arb. 25: 319-341. 1944.

Shrubs or trees with persistent leaves, the pubescence stellate; leaves short-
petiolate, entire or dentate, penninerved; stipules deciduous; flowers polygamous
or dioecious, apetalous, in axillary racemes, subtended by small bracts; sepals 1-5
or none; stamens 2-8, spatulate, the filaments short, the anthers flat, convex
outside; pistillate flowers with a superior stellate-tomentose ovary, with several

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 427

stamens or none; styles 2, stigmatic on the inner side; seed 1 in each cell of the
capsule.

Ten species are known in Asia, and two other American ones grow
in Honduras and Sinaloa, Mexico. The isolated species occurring
in Central America and Mexico present a puzzling instance of dis-
continuous distribution, but one that may be matched in some
other groups.

Distylium guatemalense Radlk. ex Harms, Notizbl. Bot. Gart.
Berlin 11: 716. f. 13. 1933.

Known only from the type, Coban, Alta Verapaz, 1,350 meters,
Tuerckheim 11.1613, and another collection made by the same col-
lector on the "Rio Chiu" (Chiuc?) in the same department.

A tree, the branches terete, somewhat flexuous, sparsely stellate-pubescent
when young or sometimes furfuraceous-tomentose; leaves short-petiolate, ovate
or elliptic-lanceolate, 6-14 cm. long, 2.5-6 cm. wide, acute, at the base acute or
subobtuse, on petioles 1-1.5 cm. long, entire or nearly so, chartaceous, the lateral
nerves 4-5 pairs, glabrous above or nearly so, stellate-pubescent beneath; stipules
minute, linear, soon deciduous; inflorescences axillary, spike-like, dense, on
peduncles 5-10 mm. long, the spikes about 1.5 cm. long, the flowers sessile or nearly
so, mostly perfect; sepals 5-6, unequal, obovate-oblong or obovate-lanceolate,
obtuse to acuminate, pubescent outside, 3-4 mm. long; petals none; stamens 5-6,
exserted, the filaments glabrous; anthers broadly oblong, fasciculate-pilose at the
apex, 1.2-1.5 mm. long; ovary sessile, laterally compressed, villous-hirsute, 2-
lobate; styles 2, elongate, divaricate, linear.

The tree probably is rare in Alta Verapaz, for in spite of persistent
search we have been unable to find it. Not much reliance can be
placed on the locality "Coban" on this or others of Tuerckheim's
labels, for many of his plants so labeled came from localities many
miles from that town. The other Central American species is D.
hondurense Standl., known from the vicinity of Siguatepeque,
Comayagua, Honduras, and also from Montana de la Flor, Teguci-
galpa. It is a tree of 6-9 meters, growing in pine-oak forest, and the
Guatemalan species probably will be found in a similar environment.

LIQUIDAMBAR L. Sweet gum

Large trees, deciduous or in the tropics often in leaf almost or quite through-
out the year; leaves slender-petiolate, palmately 3-7-1 obate, serrate, with small
stipules; flowers small, apetalous, usually monoecious, in large globose pedunculate
heads; staminate flowers without a perianth, intermixed with small bracts, forming
a terminal raceme; pistillate flowers in slender-pedunculate globose heads, com-
posed of more or less coherent, 2-rostrate ovaries, subtended by minute scales;

428 FIELDIANA: BOTANY, VOLUME 24

fruiting head globose, somewhat spiny from the persistent styles, consisting of
dehiscent capsules, each with 1-2 winged seeds.

Species 4, all except the following in Asia.

Liquidambar Styraciflua L. Sp. PI. 999. 1753. S. macro-
phylla Oerst. AmeY. Centr. 16. 1863. L. Styraciflua var. mexicana
Oerst. loc. cit. Liquidambar; Ocop, Occob (Quecchi); Estoraque
(Coban); Oc6m (Quecchi); Tzote (Huehuetenango) ; Quiramba
(Tactic, Alta Verapaz).

Moist or wet, often mixed forest, mostly on mountain sides or
along streams, often associated with pine or oak, 900-2,100 meters;
Alta Verapaz; Baja Verapaz; El Progreso; Izabal; Zacapa; Chiqui-
mula; Quiche"; Huehuetenango. Eastern and southern United
States; southern Mexico; Honduras; Salvador; Nicaragua.

A tall tree, sometimes 40 meters high (reported in Guatemala as reaching
35 meters or even more), the bark deeply furrowed, grayish, the trunk often a
meter or more in diameter; young branches red-brown, often with corky ridges
or thick wings; leaves on very long, slender petioles, 10-18 cm. long and wide,
cordate or subcordate, deeply 5-7-1 obate, the lobes oblong-triangular, acuminate,
finely serrate, dark green and lustrous above, paler beneath, glabrous except for
tufts of hairs in the axils of the nerves; fruit heads 3 cm. in diameter.

The usual name for the tree in Guatemala is "liquidambar," a
word said to have been corrupted in Honduras into "diquidambo."
Such corruptions are not at all uncommon in Central America in the
case of long and more or less difficult words. A caserio of the Depart-
ment of Chiquimula has been given the name Liquidambar. The
sap wood is nearly white; heartwood brown or reddish brown, with
satiny luster, sometimes beautifully figured with dark markings;
specific gravity 0.55-0.65, the weight about 35 pounds per cubic
foot; grain usually irregular, the texture fine and uniform; easy to
work, finishes very smoothly; likely to warp badly if not carefully
dried; not highly resistant to decay. In Guatemala, as far as we
have been able to learn, little or no use is made of the wood, in spite
of its often great abundance. In the United States it is much used
for furniture, interior trim, doors and panels, veneers for plywood,
baskets, dishes, vegetable barrels, and many other purposes. In
Mexico it is said to be employed for match sticks and toothpicks.
In many regions this is one of the commonest of Guatemalan trees,
especially about Coban, where it is often the dominant tree. The
trees are conspicuous because of their often tall and spire-like crowns,
but the crowns are sometimes rather broadly pyramidal or in age
even somewhat rounded. In the spring months (of the North) the

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 429

trees are conspicuous also because of their new, bright pale-green
foliage, this characteristic color, so common in the north, being
decidedly unusual in the tropics. Before the old leaves fall they
often are colored in shades of red, but this coloring is less conspicuous
in Central America than in the United States. Branches with full-
grown leaves are used at Coban as decorations during Holy Week.
The largest and handsomest trees we have seen in Guatemala are
those at Finca La Isla, south of Santa Cruz, Alta Verapaz, where
there are many of them, apparently planted long ago, bordering a
large shallow pond in a pasture. Some of the trunks are a meter in
diameter above the base, but many of them are hollow, and the basal
portion, which probably is covered with water during the invierno,
has large irregular openings, so that the trunk is upheld on stilts, as
it were. Inquiry has not revealed a similar growth habit in the
southern United States, where it might be expected. These giant
trees bear flowers, young fruits, and mature fruits at the same time.
They often support many ferns, orchids, and other epiphytes. We
have noted a few Liquidambar trees planted for shade in Guatemala
City.

A resin or balsam obtained from the sap of this tree is little
known in the United States, apparently, but it is much used in
Central America, and large amounts of it have been shipped to
Europe from Central America and Mexico. It is a transparent
yellowish liquid with a peculiar balsamic odor and a bitter, warm,
acrid taste. It hardens upon exposure to air. In Europe the balsam
was employed variously in medicine, and it is much used in domestic
medicine in Guatemala, for treating sores and wounds of people
and domestic animals. The Indians of Alta Verapaz bathe in a
decoction of the leaves and also take the balsam internally as a sup-
posed remedy for gonorrhea. The hardened gum often is chewed
to clean and "preserve" the teeth. In pre-Conquest days Liquidam-
bar was employed as incense in the temples and houses, and for
flavoring smoking tobacco. Its use is mentioned by Bernal Diaz de
Castillo, associate of Cortez, who wrote his account of the Conquest
in a house that is still standing in Antigua, Guatemala.

MATUDAEA Lundell

Small trees with stellate-lepidote pubescence; leaves alternate, short-petiolate,
subcoriaceous, narrow, entire, triplinerved; stipules small, linear, deciduous;
flowers small, perfect, axillary, spicate; calyx ovoid, closed in bud, rupturing at
anthesis into irregular lobes; petals none; stamens 20-24, the filaments stout,
elongate; anthers oblong, acute, the cells longitudinally dehiscent; ovary 2-celled,

430 FIELDIANA: BOTANY, VOLUME 24

the 2 styles stout, recurved, stigmatose within; ovules solitary; capsule ligneous,
oblong-ovoid, 2-cuspidate, 2-valvate at the apex, the valves 2-fid; seeds oblong-
ovoid.

A single species is known.

Matudaea trinervia Lundell, Lloydia 3: 210. 1940.

Wet mixed forest, 1,500-1,600 meters; Alta Verapaz (mountains
along the road between Tactic and the divide on the road to Tamahu,
Standley 91320, 91327). Chiapas, the type from Mount Ovando.

A tree of 10 meters, the branchlets sparsely stellate-lepidote; stipules linear,
1 cm. long or less; leaves on petioles 5-17 mm. long, lanceolate or oblong-lanceolate,
6-15 cm. long, 2-5 cm. wide, acuminate to long-acuminate, rounded or obtuse at
the base and often somewhat unequal, triplinerved, at first rather sparsely stellate-
pubescent on both surfaces, in age almost glabrous but usually barbate in the axils
of the basal nerves; inflorescences short-spicate, head-like, 13 mm. long or less,
short-pedunculate, stellate-lepidote, the flowers few, crowded, sessile or nearly so;
calyx 3 mm. long, stellate-lepidote; filaments glabrous, the anthers stellate-
lepidote; styles as much as 4.5 mm. long; capsule 2-seeded, oblong-ovoid, stellate-
lepidote; seeds black, lustrous, oblong-ovoid, 7.5 mm. long, slightly compressed.

The Guatemalan material is sterile but there is every reason to
believe that it is properly referred to this recently described genus.
The specific determination is questionable, and it is probable that a
second species occurs in Guatemala, since there are rather definite
differences in pubescence and venation of the leaves. It seems
unwise to describe a species from these sterile collections, for there
is a possibility that they do not belong to this genus.

PL AT AN ACE AE. Sycamore Family

Reference: Henry A. Gleason, Platanaceae, N. Amer. Flora 22:
227-229. 1908.

Trees with exfoliating bark; leaves alternate, petiolate, stipulate, usually
broad, palmately 3-5-nerved and lobate, cuneate to cordate at the base; stipules
membranaceous, lobate or entire, connate, deciduous; petioles dilated at the base
and covering the buds; flowers unisexual, in very dense and many-flowered, globose
heads, these racemose or spicate, or solitary and terminal on long peduncles;
flowers small and greenish, monoecious, usually isomerous; sepals 3-4 or rarely 6,
spreading, pubescent outside; petals alternate with the sepals, spatulate, lobate
or emarginate; stamens of the staminate flowers alternate with the sepals, the
filaments short; anthers linear, the connective dilated at the apex and covering
the anthers; staminodia of pistillate flowers caducous; carpels of the ovary 3-4,
distinct, opposite the petals and adherent to them; ovary linear, the style linear,
with a recurved tip, stigmatose on the inner side; ovule usually 1, pendulous,
orthotropous; fruit an achene, indehiscent, linear, 4-sided, surrounded at the base

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 431

by long hairs, truncate, tipped with the persistent style, collected in large, very
dense, globose heads.

A single genus, with about 10 species, ranging in America from
southern Canada to Guatemala, and in the Old World from south-
eastern Europe to India. Only one species extends to Central
America. In Mexico there are 7, more than in all the rest of the earth
combined.

PLAT ANUS L. Sycamore

With the characters of the family. The wood of P. occidentalis L.
of the eastern and southern United States is used for boxes and
crates, furniture, plywood, butchers' blocks, woodenware, and other
articles. It is similar to that of other species, light brown to pinkish,
the sap wood almost white; it has a specific gravity of about 0.50,
is firm, tough, and strong.

Platanus chiapensis Standl. Contr. U. S. Nat. Herb. 20: 212.
1919.

At about 1,100 meters; Quiche" (Finca San Francisco, Cotzal,
A. F. Skutch 1839; Rio Chipal, near Nebaj, A. J. Sharp 45154).
Chiapas, the type from Zincantan.

A giant tree, in Guatemala as much as 45 meters high, with a trunk 2 meters
in diameter, the bark light gray to almost white, peeling off in sheets, the branch-
lets reddish brown, covered at first with a dense felt-like tomentum, soon glabrate;
leaves on stout petioles 2-6 cm. long, broadly ovate to ovate-orbicular, 9-23 cm.
long, 6-19 cm. wide, obtuse to truncate at the base and usually abruptly cuneate-
decurrent, triplinerved, usually 3-lobate to the middle or less deeply, the lobes
acuminate, commonly entire, glabrate in age on the upper surface, densely covered
beneath with a rather lax tomentum of whitish or yellowish, branched hairs;
inflorescences as much as 25 cm. long, the flower heads 3-5, on peduncles 1-2 cm.
long, globose, 2.5-3 cm. in diameter; achene 5-5.5 mm. long, glabrous below,
densely pilose above.

Apparently this is very local in distribution in Guatemala, and
known only from the two collections cited. The species is said to
be common in some of the mountains of Chiapas. Even for the
tropics, this is a very large tree, as the dimensions indicated by Skutch
prove. P. occidentalis L. is the largest broad-leaf tree of the United
States, attaining sometimes a height of 50 meters. It is much planted
as a shade tree.

In Central Park of Guatemala City there have been planted
several trees of Platanus acerifolia (Ait.) Willd., and it is probably
the same form that is growing in La Aurora, on the outskirts of

432 FIELDIANA: BOTANY, VOLUME 24

Guatemala City. Some of the trees are as much as 8 meters high,
and they appear to thrive in £his climate. The leaves are green
beneath, not white-tomentose. These trees are said to have been
imported from California. P. acerifolia is the London plane, much
planted for shade, especially along streets, in the United States and
Europe. It is said to be a hybrid between P. occidentalis and
P. orientalis L., of southeastern Europe and western Asia.

ROSACEAE. Rose Family

Reference: P. A. Rydberg, Rosaceae, N. Amer. Fl. 22: 239-533.
1908-1918.

Herbs, shrubs, or trees, with persistent or deciduous leaves, often armed with
thorns or prickles; leaves alternate or rarely opposite, simple or compound,
usually with conspicuous stipules; flowers usually perfect, rarely unisexual, com-
monly regular, the perianth perigynous, the axis sometimes enlarged to form a
flat to urceolate or conic receptacle or a hypanthium bearing the sepals, petals,
and stamens on its margin, inside usually lined with a glandular disk; sepals 4-5,
imbricate; petals 4-5, imbricate, sometimes wanting; stamens 5-many; carpels
of the ovary 1-many, distinct or united, often connate with the receptacle; carpels
enclosing 1-several erect or pendulous ovules; styles as many as the carpels, some-
times connate; fruit a follicle, achene, drupe, hip (in Rosa), or pome; seeds usually
without endosperm, the cotyledons often carnose and convex, rarely folded or
convolute.

A large family, well represented in almost all parts of the
earth. Probably all the Central American genera are represented in
Guatemala.

Leaves pedately or pinnately compound, with 3 or more leaflets.
Plants armed with prickles.

Fruit of few or numerous drupelets borne on an enlarged receptacle; native

or cultivated plants Rubus.

Fruit of achenes borne on the inside of a fleshy receptacle (hip); cultivated

plants Rosa.

Plants unarmed.

Flowers capitate or in head-like spikes Poterium.

Flowers not capitate or in head-like spikes.

Calyx bearing numerous uncinate prickles; leaves pinnate Agrimonia.

Calyx unarmed.

Fruit armed with barbate spines; leaves pinnate Acaena.

Fruit unarmed.

Petals none Alchemilla.

Petals present.

Receptacle not enlarged and fleshy in fruit; leaves pinnately or

pedately compound, with 3 or more leaflets Potentilla.

Receptacle in fruit enlarged and juicy; leaves with 3 leaflets.

Petals white; fruit sweet, edible Fragaria.

Petals yellow; fruit bitter, inedible : . .Duchesnea.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 433

Leaves simple.

Fruit consisting of 1-5 dehiscent follicles or of achenes and dry, or of several or
numerous fleshy drupelets.

Fruit of drupelets Rubus.

Fruit dry, of 1 or more follicles or achenes.

Fruit of achenes; leaves white-tomentose beneath Holodiscus.

Fruit of 1-5 follicles; leaves tomentose or glabrous beneath.

Fruit of usually 5 follicles; cultivated plants Spiraea.

Fruit of 1 follicle; native plants Guamatela.

Fruit indehiscent, usually fleshy and juicy.
Ovary superior.

Style terminal or nearly so; leaves sometimes serrate Prunus.

Style basilar; leaves entire.
Stamens 3-10.

Petals none or minute Licania.

Petals 5 Hirtella.

Stamens 15-many.

Calyx tube campanulate; inflorescence cymose; ovary sessile in the
bottom of the calyx Chrysobalanus.

Calyx tube elongate; inflorescence racemose or paniculate; ovary

adnate to the throat of the calyx Couepia.

Ovary inferior.

Carpels of the fruit bony at maturity, the fruit containing 1-5 nutlets;

trees armed with thorns Crataegus.

Carpels with leathery or papery walls at maturity; fruit a 1-5-celled pome,

each cell with 1 or more seeds; plants usually unarmed.
Flowers in compound corymbs or panicles.

Leaves rusty-tomentose beneath; carpels of the ovary wholly connate;

cultivated trees Eriobotrya.

Leaves not tomentose beneath except when very young; carpels free

at the apex; native trees . . . Photinia.

Flowers in umbels or racemes, or solitary.

Carpels of the fruit containing 4 or more seeds.

Styles connate at the base; leaves serrate or crenate; petals red.

Chaenomeles.

Styles free; leaves entire; petals white or pink Cydonia.

Carpels of the fruit 1-2-seeded.

Ovary and fruit 2-5-celled, the cells 2-ovulate.

Styles connate at the base; fruit apple-shaped, usually without

grit cells Mains.

Styles free; fruit pear-shaped, containing numerous grit cells.

Pyrus.
Ovary and fruit incompletely 6-10-celled, the cells 1-ovulate.

Amelanchier.

ACAENA Mutis

Plants woody or suffrutescent, usually low; leaves odd-pinnate, the stipules
nore or less adnate to the petioles; inflorescence spicate or racemose, the lower
lowers mostly remote; hypanthium ellipsoid, covered with retrorsely barbed

434 FIELDI AN A: BOTANY, VOLUME 24

prickles, contracted above and usually produced into a short tube; calyx 3-5-
lobate, usually 4-lobate; corolla none; stamens 3-5, inserted in the mouth of the
hypanthium; pistil usually 1, the style terminal, short, the stigma multifid;
achene wholly enclosed in the prickly hypanthium; seed pendent, the radicle
superior.

Species about 40, mostly in South America, southern Africa, and
Australia. In North America three species are known, one in central
California, another in the high mountains of Costa Rica.

Acaena elongata L. Mant. PI. 200. 1771. A. agrimonioides
HBK. Nov. Gen. & Sp. 6: 231. 1823. Mozote; Mozotillo; Tlacha
(Volcan de Santa Maria); Pegapega; Secam (Volcan de Zunil).

Open meadows or hillsides of the higher mountains, often in
moist thickets or in wet Cupressus or Abies forest, sometimes on
limestone, mostly at 2,400-4,000 meters; Jalapa; Sacatepe"quez;
Solola; Chimaltenango; Totonicapan; Quezaltenango; Huehue-
tenango; San Marcos. Central and southern Mexico; Costa Rica;
Colombia.

Plants usually woody, sometimes almost wholly herbaceous, commonly 25-50
cm. tall, sometimes a meter high or slightly more, often forming dense colonies,
the bark brown or purplish; leaflets 9-19, oval or elliptic, acute at each end, 8-15
mm. long, acutely serrate, subcoriaceous or herbaceous, glabrous and lustrous
above, sparsely sericeous-strigose beneath along the veins; stipules linear-lanceo-
late, adnate to the petioles, ciliate; racemes remotely or densely flowered, mostly
10-30 cm. long, the flowers subsessile, 3-bracteate; hypanthium sparsely villous
when young, covered with numerous sharp barbed prickles; calyx lobes ovate,
1 mm. long; fruit short-pedicellate, ellipsoid, nutant, 6-8.5 mm. long, glabrate in
age, the spines brown or vinaceous, 2-3 mm. long, commonly with 3 barbs at the
apex.

Acaena is without doubt one of the most noxious plants of the
highlands, and also one of the most characteristic and common at
higher elevations, particularly in the sheep country. In some regions,
such as the white pine-Abies forests of Totonicapan, it is the domi-
nant low plant, and in many other areas it is scarcely less plentiful.
Acaena and sheep form a natural association. The troublesome burs
that adhere tenaciously to the clothing of people or even to their
skin naturally become entangled with the fleeces of the many sheep
that feed in the highlands. Some of the sheep, in fact, are one matted
mass of wool and Acaena burs. It would be of some interest to know
to what extent the plant has been spread by this agency, but possibly
it was always as abundant in the Guatemalan mountains as now.
Apparently its foliage is not eaten by sheep or other animals. The
leaves, at least during the colder months, usually are tinged with dark

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 435

red or bronze, making patches of the plant conspicuous even from
some distance. The Mexican A. agrimonioides HBK. was main-
tained as distinct by Rydberg, although he reported it only from the
original locality. A fragment and photograph of the type in the
Herbarium of Chicago Natural History Museum indicate that this
supposed species differs in no respect from A. elongata.

AGRIMONIA L.

Perennial herbs with rootstocks; leaves odd-pinnate, the leaflets very unequal,
smaller ones interposed between the larger ones, coarsely toothed; flowers regular
and perfect, in spike-like interrupted racemes; hypanthium hemispheric to
obconic, constricted at the throat and enclosing the achene in fruit, usually 10-
striate vertically, bearing at the margin a ring of uncinate spines; sepals 5, after
anthesis connivent and forming a nipple-shaped beak on the fruit; petals 5, small,
yellow, not unguiculate; stamens 5-15, with slender filaments; pistils 2, the styles
terminal, the stigmas bilobate.

About a dozen species, in cooler regions of the northern hemi-
sphere. The genus is a temperate rather than a tropical one, and
only the following species extends as far southward as Central
America.

Agrimonia macrocarpa (Focke) Rydb. N. Amer. Fl. 22: 392.
1913. A. parviflora var. macrocarpa Focke ex Bonn. Smith, Bot.
Gaz. 16: 3. 1891. Chichicaste (Quezaltenango, a meaningless and
probably erroneous name).

Moist thickets or fields, sometimes in pine-oak forest, 1,200-
2,100 meters, probably endemic, but perhaps extending into southern
Mexico; Alta Verapaz (type from Coban, Tuerckheim 1409); Baja
Verapaz; Chimaltenango; Quiche"; Huehuetenango; Quezaltenango;
Suchitepe"quez; San Marcos.

Plants a meter high or less, simple or branched, hirsute with long, brownish
or yellowish hairs, glandular-granuliferous above; petioles and leaf rachis hirsute;
stipules broadly and obliquely ovate or semicordate, 1-2 cm. long, coarsely
dentate; larger leaflets 7-9, oval to elliptic-lanceolate, 3-6 cm. long, thin, acute
or obtuse, rather densely hirsute, glandular-granuliferous beneath, coarsely
dentate or crenate; smaller leaflets only 3-10 mm. long; racemes 10-20 cm. long,
the pedicels ascending, 2-5 mm. long; sepals lanceolate, acuminate, green,
3-nerved, incurved in fruit, 2-2.5 mm. long; petals obovate, 3-3.5 mm. long;
bristles of the hypanthium in numerous series, the inner ones 3 mm. long, the outer
ones shorter and reflexed.

The plant is a nuisance wherever it grows because of the bur-like
fruits that adhere to clothing. Although widely distributed in
Guatemala, we have not found it plentiful in any locality.

436 FIELDIANA: BOTANY, VOLUME 24

ALCHEMILLA L.

Reference: Lily M. Perry, A tentative revision of Alchemilla
section Lachemilla, Contr. Gray Herb. 84: 3-57. 1929.

Herbs, mostly perennial, most often prostrate or procumbent, the stems leafy;
leaves alternate, usually orbicular and digitately lobate or palmately parted,
sometimes pinnately cleft or parted; stipules sheathing, adnate to the petiole;
flowers minute, generally crowded in dense corymbs, sometimes laxly cymose or
solitary, ebracteate, sessile or pedicellate; hypanthium urceolate, persistent,
constricted in the throat, the limb 8-10-lobate, the lobes 2-seriate, the inner ones
valvate, the outer ones small; petals none; stamens 1-4, inserted in the throat of
the calyx, small, the filaments short, free; disk with a thickened margin, closing
the mouth of the calyx; ovary of 1-4 carpels, these sessile or substipitate in the
bottom of the calyx, free; styles basal or ventral, filiform, glabrous, the stigma
capitellate; ovules solitary, ascending from the base of the cell; achenes 1-4,
included in the calyx, membranaceous.

About 100 species, widely distributed in both hemispheres,
chiefly in temperate regions; in the tropics confined to mountain
regions. A few other species are found in southern Central America.
Many of the species are much alike in general appearance and are
separated only by minute details of pubescence and flower structure.

Leaves pinnately parted A. pinnata.

Leaves simple or palmately parted or cleft.
Basal leaves 5-11-lobate or 5-11-cleft.

Leaves shallowly 9-11-lobate A. pectinata.

Leaves deeply 5-9-cleft, the lobes extending well toward the base of the blade.

A. guatemalensis.
Basal leaves 3-5-cleft or 3-5-parted.

Hypanthium pubescent or villous within; inflorescence usually a rather open

cyme with flowers on pedicels 2-10 mm. long.
Leaves appearing 5-lobate, the lateral segments bifid; stipules lobate or

incised-dentate A. procumbens.

Leaves appearing 3-lobate, the lateral segments not bifid; stipules bifid.

A. vulcanica.

Hypanthium glabrous within; inflorescence usually glomerate, the flowers
mostly sessile or on very short pedicels.

Flowers glabrous A. aphanoides.

Flowers pubescent, sometimes glabrate in age.

Leaves appearing 3-lobate, the lateral lobes not bifid; achenes tapering

to an acute apex A. Pringlei.

Leaves appearing 5-lobate, the lateral lobes bifid; achenes subacute or
subobtuse A. sibbaldiae folia.

Alchemilla aphanoides L. f. Suppl. PL 129. 1781. A. sub-
alpestris Rose, Contr. U.' S. Nat. Herb. 10: 96. 1906. Lachemilla
subalpestris Rydb. N. Amer. Fl. 22: 384. 1908. A. aphanoides var.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 437

subalpestris Perry, Contr. Gray Herb. 84: 39. 1929. Flor de suelo
(Quezaltenango) ; Losan (Huehuetenango).

Moist or dry fields and banks, oak, pine, or Cupressus forest,
often in exposed, sometimes rocky, alpine places, occasionally a
weed in corn fields, 1,800-4,000 meters; Baja Verapaz; Jalapa;
Chimaltenango; Solola; Huehuetenango; Totonicapan; Quezal-
tenango; San Marcos. Mexico; Costa Rica; Andes Mountains of
South America.

Plants perennial, usually prostrate or decumbent, sparsely or densely pilose,
the stems often numerous and much branched; leaves 3-parted, the lower ones
petiolate, pilose or villous, the upper ones sessile, often glabrate, the lobes incised-
lobate or incised-dentate, oblanceolate to cuneate-obovate, 1.5 cm. long or mostly
less than 1 cm. long; stipules 2-4-lobate or 2-4-cleft; flowers aggregate in axillary
and terminal cymes, usually on very short pedicels, subtended by lobate bracts;
hypanthium urceolate, glabrous at maturity and often from the first; calyx lobes 8,
acute, mostly subequal, the bractlets lanceolate or lance-ovate, the sepals ovate;
achenes 1-3.

This, like some of the other species, is a very common plant of
open places in the highlands of Guatemala. The Guatemalan
material is referable to var. subalpestris., the typical form of the
species being confined to the South American Andes.

Alchemilla guatemalensis Rothm. Notizbl. Bot. Gart. Berlin
12: 489. 1935. Fresilla (fide Aguilar).

Moist or wet fields and banks, sometimes in white sand in Alnus
forest, 1,500-2,700 meters; Alta Verapaz (type from Chucaneb,
Tuerckheim 1783); Sacatepe"quez ; Quiche"; Retalhuleu; Quezal-
tenango; San Marcos. Southern Mexico.

A procumbent or prostrate perennial, producing stolons, the stems often
numerous, short- villous, leafy, often much branched; basal and lower cauline
leaves rounded-reniform, 1.5-3.5 cm. wide, thin, 5-9-cleft, shallowly cordate at the
base, the lobes obovate to broadly cuneate-obovate, incised-dentate above the
middle, green and sparsely pilose on the upper surface, paler and villous or hirsute-
villous beneath, especially on the nerves; petioles 1-4 cm. long, villous; stipules
membranaceous, incised at the apex; upper leaves similar to the lower ones but
smaller and short-petiolate, with fewer lobes, the uppermost leaves sessile,
3-parted; flowers axillary or in terminal racemose cymes, short-pedicellate, 2-2.5
mm. long; hypanthium 1.5 mm. long, turbinate-urceolate, densely pubescent;
bractlets lanceolate, acute, slightly narrower than the broadly lanceolate sepals;
pistils 2-4.

This has been reported from Guatemala as A. venusta Schlecht.
& Cham.

438 FIELDIANA: BOTANY, VOLUME 24

Alchemilla pectinata HBK. Nov. Gen. & Sp. 6: 226. 1824.
A. pectinata var. mexicana Perry, Contr. Gray Herb. 84: 15. 1929.
Hoja redonda (San Marcos).

Moist or dry banks or meadows, or in forest of pine, oak, Cupres-
sus, Juniperus, or Alnus, 1,800-4,000 meters; Alta Verapaz; El
Progreso; Guatemala; Sacatepe'quez; Chimaltenango; Huehue-
tenango; Totonicapan; Quezaltenango; San Marcos. Southern
Mexico; Costa Rica; Andes of South America.

A stoloniferous perennial, procumbent or prostrate and creeping, the stems
often elongate and branched; lower leaves rounded-reniform, 1.5-4 cm. wide,
rather thick, shallowly 9-11-lobate, deeply cordate at the base, pectinate-serrate,
green and glabrate above, sericeous and often pale beneath; petioles chiefly 3-6
cm. long, sericeous or sometimes pilose with spreading hairs; stipules large, mem-
branaceous, light brown; upper leaves short-petiolate or sessile; inflorescence
racemose, axillary or terminal, the flowers large, 3.5-4 mm. long, densely sericeous;
hypanthium turbinate-campanulate, the lobes 8 or 10, spreading, ovate-oblong
or ovate, acute; bractlets equaling or slightly narrower than the sepals; carpels
4-6; achenes 2-4.

This has been reported from Guatemala as A. orbiculata Ruiz &
Pavon, a South American species.

Alchemilla pinnata Ruiz & Pavon, Fl. Peruv. 1: 69. 1798.

Moist or wet, alpine meadows, or in open forest of pine and
Abies, 3,000-4,300 meters; Huehuetenango (Sierra de los Cuchu-
matanes); San Marcos (Volcan de Tacana). Southern Mexico;
Andes of South America.

A prostrate perennial, often forming dense mats; basal leaves bipinnatifid,
2-6 cm. long, the petioles 3.5 cm. long or shorter, villous, the leaf blades linear-
oblong; pinnae 8-15 pairs, unequally 2-cleft or 2-parted, the pinnules 3-7 mm.
long; stipules chartaceous, lanceolate or lance-ovate, entire; upper leaves pinnate
or the uppermost 3-parted, their stipules multilobate, sheathing; flowers solitary
and axillary or in terminal glomerules, the pedicels 2-5 mm. long; hypanthium
villous or glabrate, the lobes lance-ovate, acute or subobtuse; achenes 1-2.

A characteristic but uncommon alpine plant of the high moun-
tains of western Guatemala.

Alchemilla Pringlei Fedde, Bot. Jahresb. 36, Abt. 2: 496. 1910.

Moist or rather dry, open, grassy places, sometimes in Cupressus
forest, 1,500-3,800 meters; Sacatepe'quez (locality questionable);
Quezaltenango; San Marcos; Huehuetenango. Mexico.

Plants perennial, the stems procumbent or prostrate, sometimes suberect,
branched, hirsute with short spreading hairs; leaves 3-parted, usually glabrous
above and sparsely hirsute beneath, the lower ones short-petiolate, the upper ones

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 439

sessile, the lobes obovate or cuneate, coarsely serrate-dentate, the lateral lobes
rarely bifid; stipules cleft into 2-4 lance-linear or oblong lobes; flowers in terminal
or axillary glomerules, on very short pedicels; hypanthium broadly urceolate,
1 mm. long, pubescent, the lobes acute; bractlets lanceolate, often slightly shorter
than the ovate sepals; achenes 2-4, sharp-pointed.

Alchemilla procumbens Rose, Contr. U. S. Nat. Herb. 10: 96.
1906. Lachemilla procumbens Rydb. N. Amer. Fl. 22: 382. 1908.
L. costaricensis Dammer, Repert. Sp. Nov. 15: 362. 1918. Alfombra
de llano (fide Aguilar).

Moist or rather dry, open slopes or fields, sometimes in rocky
places, frequently in Alnus or Juniperus forest, 1,900-3,700 meters;
Sacatepe"quez; Chimaltenango; Solola; Quiche"; Huehuetenango;
Totonicapan; Quezaltenango; San Marcos. Costa Rica; Andes of
South America.

Plants perennial, the stems often numerous, usually prostrate and creeping,
often much branched, glabrate or appressed-pilose; leaves 5-25 mm. long, 3-parted,
rather thick and firm, glabrous above, sericeous or appressed-pilose beneath, the
lobes spatulate to obovate, coarsely serrate-dentate, the lateral ones unequally
bifid; petioles 2-10 mm. long, appressed-pilose; stipules foliaceous, oblong, lobate
or incised-dentate; inflorescence laxly cymose, the pedicels 3-10 mm. long; hypan-
thium 1.5-2 mm. long, campanulate, sericeous outside, densely pubescent within;
calyx lobes 8 or 10, the bractlets ovate, acuminate, usually longer than the broadly
ovate, acute sepals; achenes 3-8.

Alchemilla sibbaldiaefolia HBK. Nov. Gen. & Sp. 6: 225. pi.
561. 1824. Lachemilla sibbaldiaefolia Rydb. N. Amer. Fl. 22: 384.
1908. L. Tonduzii Dammer, Repert. Sp. Nov. 15: 362. 1918. A. sib-
baldiaefolia var. Tonduzii Perry, Contr. Gray Herb. 84: 34. 1929.

Moist open slopes or meadows, often in thin or dense forest of
pine, oak, or Cupressus, 1,500-3,400 meters; Baja Verapaz; Jalapa;
Guatemala; Sacatepe"quez; Chimaltenango; Huehuetenango; Totoni-
capan; Quezaltenango. Mountains of Mexico; Costa Rica.

Plants perennial, the stems often numerous, usually decumbent to prostrate,
often much branched, hirsute or pilose with appressed or somewhat spreading
hairs; lower leaves short-petiolate, 3-parted, with the lateral lobes bifid, the upper
leaves subsessile and 3-parted, the segments narrowly obovate to oblanceolate,
incised-serrate, glabrate above, paler and appressed-pilose beneath; stipules
about equaling the petioles, cleft into 2-4 linear-oblong lobes; flowers glomerate
near and at the ends of the stems, the pedicels very short; hypanthium campanu-
late-urceolate, appressed-pubescent, the lobes acute, equal in length; bractlets
lanceolate, the sepals ovate; achenes 2-3.

Some of the Guatemalan material is referred by Miss Perry
(who has determined many of our collections) to var. Tonduzii, which
is not very definitely different from the typical form of the species.

440 FIELDIANA: BOTANY, VOLUME 24

Alchemilla vulcanica Schlecht. & Cham. Linnaea 5: 573.
1830. Lachemilla vulcanica Rydb. N. Amer. Fl. 22: 382. 1908.

Open, grassy, often rocky places, chiefly at high elevations and
often in alpine meadows, sometimes in open forest of pine, Abies, or
Juniperus, occasionally on limestone, 2,700-3,900 meters; Sacate-
pe"quez (Volcan de Agua) ; Chimaltenango (Volcan de Acatenango) ;
Huehuetenango (Sierra de los Cuchumatanes) ; Quezaltenango
(Volcan de Santa Maria) ; San Marcos. High mountains of central
and southern Mexico; Andes of South America.

Plants perennial, often with numerous branched stems, the stems creeping,
appressed-pilose; leaves mostly 1 cm. long or smaller, 3-parted, the lateral lobes
usually simple, somewhat pilose on both surfaces or glabrous above, the veins
impressed, the lobes cuneate, deeply incised into 3-7 linear-lanceolate segments;
petioles 3-5 mm. long; stipules sheathing, bifid, the lobes linear; inflorescence
terminal, cymose, the pedicels 1-5 mm. long, pubescent; hypanthium turbinate-
campanulate, 1.5 mm. long, densely pubescent outside, sparsely pilose within;
bractlets linear to lanceolate, acute; sepals lance-ovate to broadly ovate, obtuse
or subacute; achenes 1-4.

AMELANCHIER Medicus

Shrubs or trees, unarmed; leaves alternate, simple, petiolate, persistent or
deciduous, thin or coriaceous, serrate or entire; flowers white or pinkish, mostly
racemose; hypanthium campanulate, the limb 5-lobate, the lobes narrow, reflexed,
persistent; petals 5, broad or narrow; stamens numerous, inserted in the throat of
the hypanthium, the filaments subulate; styles 2-5, connate, pilose at the base;
ovary wholly or partly inferior, its cavities becoming twice as many as the styles;
ovule 1 in each cavity, erect; fruit a small berry-like pome, containing 4-10 cells;
testa of the seed cartilaginous.

About 25 species, in the north temperate zone. Only one species
extends to Central America, but there are two or three others in
Mexico and a large number in the United States, where the plants
are known by such names as "shadbush," "Juneberry," and "service-
berry." The fruits of some northern species are juicy and edible,
and the shrubs are cultivated occasionally for their sweet fruit. The
fruits of the Rocky Mountain species were an important food among
some of the Indians, who often dried them for use in winter.

Amelanchier denticulata (HBK.) Koch, Dendrol. 1: 183.
1869. Cotoneaster denticulata HBK. Nov. Gen. & Sp. 6: 169. pi.
556. 1823. C. nervosa Decaisne, Nouv. Arch. Mus. Paris 10: 177.
1874. A. nervosa Standl. Contr. U. S. Nat. Herb. 23: 337. 1922.
Malacomeles denticulata G. N. Jones, Madrono 8: 36. 1945. M. ner-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 441

vosa G. N. Jones, op. cit. 38. Membrillito; Membrillo; Cerezo rojo;
Manzanita; Huiton (Huehuetenango).

Usually on dry brushy slopes or in open rocky forest, 1,400-2,300
meters; Baja Verapaz; Zacapa; Guatemala; Sacatepe"quez ; Quiche;
Huehuetenango; Quezaltenango. Southwestern Texas; Mexico.

A densely branched, rigid shrub, usually 1.5-3 meters tall, the branches
blackish or dark reddish brown; leaves petiolate, coriaceous, oval to oval-obovate
or broadly elliptic, mostly 3-4.5 cm. long, rounded to truncate and abruptly
short-pointed at the apex, subacute to rounded at the base, bright green and
glabrous above, sparsely or often very densely whitish-tomentose beneath, in age
rarely glabrate, the margin remotely serrulate or subentire; flowers in short few-
flowered terminal racemes, these often umbel-like, the pedicels short or elongate,
stout, densely tomentose; hypanthium 4-5 mm. long, densely white-tomentose,
the calyx lobes 2.5 mm. long, broadly deltoid, obtuse, reflexed, glabrous within;
petals suborbicular, twice as long as the sepals or longer, glabrous, white; fruit
oval, 1 cm. long or even larger, juicy, more or less tomentose, varying from pink
to pale red or purple.

This shrub is exceedingly abundant in some parts of the valley
of Antigua (Sacatepdquez), where large areas on hillsides are covered
almost exclusively with it. In Huehuetenango it is plentiful, but
mostly as isolated bushes. The fruit, according to experience of the
writers, is barely edible, being unpleasantly bitter. The leaves are
persistent. In general appearance the plant is substantially differ-
ent from most, although not all, of the usual Amelanchier species of
the United States. Recently Jones has separated from Amelanchier
the genus Malacomeles, recognizing two species, M. denticulata and
M. nervosa, both of which, according to his treatment, occur in
Guatemala. We can find no valid reason for maintaining the genus
Malacomeles or for recognizing two species in this group. Ame-
lanchier nervosa might be treated as a variety of A. denticulata, but
it does not seem worth while to give it even that vague distinction.

Chaenomeles lagenaria (Loisel.) Koidz. Japan quince.

An ornamental shrub, native of China, much cultivated in the
United States, where it is one of the first shrubs to bloom in spring,
and showy with its handsome scarlet-red flowers. A plant of this
was seen with a few flowers in Finca La Aurora, Guatemala City.
Several years old, the plant had produced numerous shoots, but these
were only about 30 cm. high.

CHRYSOBALANUS L.

Shrubs or small trees; leaves entire, coriaceous, short-petiolate; flowers small,
white or greenish, perfect, in terminal or axillary cymes or panicles; hypanthium

442 FIELDIANA: BOTANY, VOLUME 24

campanulate or turbinate, the calyx 5-lobate, the lobes subequal, imbricate;
petals 5, deciduous; stamens numerous, the filaments slender; ovary sessile,
1-celled; ovules 2, erect, the style filiform, basal or lateral; fruit a drupe with
juicy pulp, the osseous stone 5-6-costate.

About 3 species, 2 of them American, the other African. A
single species occurs in Central America.

Chrysobalanus Icaco L. Sp. PI. 513. 1753. Icaco.

In coastal swamps or in thickets along sea beaches, at sea level;
Izabal; San Marcos, and probably in all the Pacific coast depart-
ments; often planted inland in fincas or along hedges. Mexico to
British Honduras and Panama; Florida; West Indies; northern
South America.

A shrub or small tree, often 5-6 meters high in cultivation, along seashores
usually lower and often only 1-2 meters high, the bark thin, brownish, the branches
glabrous or nearly so, reddish brown; leaves coriaceous, on very short petioles,
elliptic to obovate or suborbicular, mostly 3-8 cm. long, rounded to obtuse or
emarginate at the apex, broadly cuneate to acute at the base, dark green and
lustrous above, dull beneath, glabrous or nearly so; cymes pedunculate, with few
or many flowers, shorter than the leaves; calyx densely sericeous, the lobes tri-
angular-ovate, acute, 2.5 mm. long; petals white, cuneate-obovate, twice as long
as the sepals; fruit globose or oval, 2-4 cm. long, white to pink or dark purple.

Cultivated frequently in the lowlands of the Pacific slope, whence
the fruit is sent to the Guatemala market. The fruit is edible, but
no one seems to esteem it highly. The flesh is somewhat spongy,
white, very juicy, and insipid in flavor. The English names are
"coco-plum" and "pigeon-plum." The leaves and fruit are reported
to yield a black dye. The large seeds are rich in oil, and it is stated
that the Caribs of the Antilles strung them on sticks and burned
them like candles. The name "icaco" (sometimes written "jicaco"
or "hicaco") is believed to be of Antillean origin. In Florida, jelly
is sometimes made from the ripe fruits. The shrub often forms a
large part of the beach thickets of Central American shores.

COUEPIA Aublet

Trees or shrubs, often white-tomentose; leaves short-petiolate, coriaceous,
entire, the petiole sometimes 2-glandular at the apex; stipules mostly setaceous
and deciduous; flowers racemose or paniculate, large or small, often tomentose,
bracteate; hypanthium elongate and somewhat tubular, subterete, often gibbous
at the base, the 5 sepals imbricate, the throat pilose; petals 5 or rarely none, slightly
longer than the calyx; stamens 10-15 or numerous, inserted in the mouth of the
hypanthium, the filaments more or less united at the base, incurved and flexuous,
the anthers small; ovary inserted laterally in the throat of the hypanthium,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 443

villous, 1-celled, the style elongate, the stigma punctiform; ovules 2, ascending
from the base of the cell, collateral; fruit drupaceous, ovoid or globose, dry or
fleshy, the stone ligneous, 1-seeded.

Species 30 or more, all American and mostly in South America.
One other species is known from Central America, in Panama.

Couepia polyandra (HBK.) Rose, Contr. U. S. Nat. Herb. 5:
196. 1899. Hirtetta polyandra HBK. Nov. Gen. & Sp. 6: 246. pi. 565.
1821. H. dodecandra DC. Prodr. 2: 529. 1825. C. Kunthiana Benth.
ex Hemsl. Biol. Centr. Amer. Bot. 1: 367. 1880. C. dodecandra
Hemsl. in Hook. Icon. PI. 27: pis. 2620, 2621. 1899. C. floccosa
Fritsch, Ann. Naturh. Hofmus. Wien 5: 12. 1890 (type locality
cited in error as Guatemalan, the type really from Guanacaste,
Costa Rica). Suncillo; Moxpin (Suchitepe'quez) ; Uspip, Zuspi
(Pete"n, fide Lundell).

Occasional in damp thickets, most often planted, rather infre-
quent, at 600 meters or less; Pete"n; Alta Verapaz; Santa Rosa;
Escuintla; Suchitepe'quez; Retalhuleu; Solola. Southern Mexico;
British Honduras to Costa Rica.

A large or medium-sized tree with gray scaly bark and spreading branches;
leaves persistent, coriaceous, short-petiolate, the blades mostly oval or elliptic-
oval, 5-8 cm. long, 2.5-5 cm. wide, rounded to obtusely subacuminate at the
apex, obtuse at the base and abruptly contracted, deep green and glabrous above,
covered beneath with a white or rusty, very close and dense tomentum, the lateral
nerves slender, elevated, conspicuous; inflorescence racemose or thyrsoid-panicu-
late, dense and many-flowered, 3-8 cm. long, the rachis and pedicels densely
tomentose, the pedicels 5-7 mm. long; hypanthium tubular or narrow-funnelform,
4 mm. long, appressed-tomentose; sepals ovate, obtuse, 4 mm. long; petals whitish
or pinkish yellow, 5 mm. long, ciliate; stamens 15-20, the filaments glabrous;
ovary densely pilose; fruits on stout thick pedicels, roughly oval and irregular,
6-8 cm. long, 3-4 cm. broad, yellowish green, somewhat verrucose, the pulp juicy,
yellow; stone 5-6.5 cm. long.

Called "baboon cap" and "monkey cap" in British Honduras; in
Honduras, "zapotillo" and "munzap"; "pio" and "uspio" in Tabasco.
Names recorded from Salvador are "uluzapote," "zapote bolo,"
"zapotillo amarillo," and "sunsapotillo." The tree blossoms at
the beginning of the rainy season and matures its fruit early in the
invierno. The fruit is edible but apparently not much esteemed; the
pulp is sweet but very fibrous. C. polyandra and C. dodecandra have
been maintained by some recent authors as distinct species, separated
by the number of stamens, but there appears to be no good basis,
if any at all, for recognizing more than a single species of Couepia
with tomentose leaves in Central America.

444 FIELDIANA: BOTANY, VOLUME 24

CRATAEGUS L. Hawthorn

Shrubs or small trees, often thorny; leaves stipulate, alternate, simple,
petiolate, dentate or lobate; flowers white or pink, in terminal corymbs; hypan-
thium cup-shaped or campanulate, the sepals 5, entire or dentate; petals 5,
spreading, rounded, inserted on the margin of the disk; stamens 5-25, in 1-3
series, the filaments filiform, the anthers white, yellow, or red; ovary inferior,
1-5-celled; styles 1-5, distinct; ovules normally 1 in each cell; fruit a small or
rather large pome, usually yellow or red, containing 1-5 osseous nutlets.

A large and difficult genus, with about 90 species in the Old World;
more than 1,000 have been described from North America, nearly
all from the United States, but large numbers of these have been
reduced by other authors, and 10 per cent of them may be valid
species. Several species are native in Mexico but in Central America
it is questionable whether any species is indigenous.

Crataegus pubescens (HBK.) Steud. Nom. Bot. ed. 2. 433.
1841. Mespilus pubescens HBK. Nov. Gen. & Sp. 6: 213. pi. 565.
1824. M. stipulosa HBK. loc. cit. C. stipulosa Steud. loc. cit.
Manzanilla; Manzanita; Cainum (Cacchiquel).

Common in cultivation in the mountains; often wild in moist or
rather dry, open forest, frequently in pine-oak forest or in thickets,
chiefly at 1,500-2,700 meters; El Progreso; Chiquimula; Jalapa;
Santa Rosa; Guatemala; Sacatepe'quez; Chimaltenango; Quiche";
Quezaltenango; San Marcos. Southern Mexico; cultivated in Sal-
vador and Costa Rica; Ecuador, where probably naturalized from
Mexico.

A very thorny tree, commonly 6-9 meters high, with rounded crown and low
thick trunk, sometimes shrubby and forming dense thickets; leaves short-petio-
late, oblanceolate or obovate, mostly 4-7 cm. long, acute or obtuse, cuneate-
attenuate to the base, green above, sparsely pilose or almost wholly glabrous,
paler beneath, sparsely or densely tomentose or pilose, crenate-serrate, often more
or less lobate; corymbs few-flowered, whitish-tomentose; sepals lanceolate, green,
tomentose, about 5 mm. long, subentire or glandular-serrulate; petals white, 1 cm.
long or less; fruit resembling a small apple, pale orange-yellow, mostly 2-3 cm.
broad.

Crataegus pubescens and C. stipulosa usually have been regarded
as distinct species, largely perhaps because the former was described
from Mexico and the latter from Ecuador. It seems almost certain,
however, that the tree was carried from Mexico to South America
in early colonial days, as happened with Prunus Capuli. The
fruit is a rather important one in Guatemala, where large quantities
are produced and consumed. During the season, in autumn and
early winter of the North, it is offered in large amounts in most of

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 445

the markets, being transported to regions where it is not grown.
The raw fruit is not very appetizing, although of moderately agree-
able flavor; it is cooked in various ways but is used principally in
preparing an intensely sweet preserve that is one of the commonest
desserts. The sirup is employed for flavoring some of the popular
aguas gaseosas, similar to soda pop. The plant is sometimes used in
Guatemala as a stock on which to graft pears and apples. The
trees often bear great quantities of fruits. A few fruits persist upon
the trees as late as February, when the blossoms unfold. Strung on
cords, the rather handsome fruits are used as decorations for Christ-
mas and other holidays. It is uncertain whether the tree really is
native in Guatemala although at times it does occur on the borders
of forest or even in forest itself. We have not noted it, however, in
what could be termed with any assurance primeval forest, and in the
places where it has been noted as possibly native, there probably
were dwellings long ago if not recently. It is possible that the tree
was introduced into Guatemala by the Mexican mercenaries who
accompanied the early conquistador es, or even by traders at some
earlier date. It is probable that in Mexico the tree behaves much as
in Central America, and there it may have been more or less domesti-
cated for a very long time.

CYDONIA Miller. Quince

Deciduous unarmed shrubs or small trees; leaf buds small, pubescent, with
few scales; leaves petiolate, entire, stipulate; flowers terminal, solitary at the ends
of leafy branchlets, white or pale pink; sepals 5, entire, reflexed; petals 5, obovate;
stamens 20; styles 5, free, pubescent below; ovary inferior, 5-celled, each cell with
numerous ovules; fruit a many-seeded pome.

The genus consists of a single species.

Cydonia oblonga Miller, Gard. Diet. ed. 8. No. 1. 1768. C.
vulgaris Pers. Syn. PI. 2: 40. 1807. Membrillo.

Cultivated commonly in the highlands, chiefly at 1,500-2,400
meters; especially frequent about Cantel and Huehuetenango;
planted also in the Oriente. Native of central Asia, but in cultiva-
tion since ancient times, and now widely dispersed in temperate
regions.

A shrub or small tree, sometimes as much as 8 meters high but usually lower,
commonly branched from the base, the young branchlets tomentose; leaves on
rather short, slender petioles, ovate to oblong, acute to rounded at the apex,
rounded or subcordate at the base, 5-10 cm. long, dull green and glabrous above,
abundantly tomentose beneath; flowers 4-5 cm. broad; hypanthium and calyx

446 FIELDIANA: BOTANY, VOLUME 24

tomentose, the sepals oblong-elliptic, obtuse; fruit broadly pear-shaped, fragrant,
yellow when ripe, more or less villous.

The leaves normally turn yellow before falling. The very sour
fruit is used in Guatemala principally in preparation of a thick jelly
or marmalade (called queso de membrillo in Mexico) similar to guava
paste in texture, that is served for desserts, but this conserve is
seldom seen on the table. The bushes in most places apparently
receive little attention after planting, and often spread to form dense
thickets, as in the Cantel region of Quezaltenango.

DUCHESNEA J. E. Smith. Indian strawberry

Perennial herbs, the trailing branches elongate, often rooting at the nodes;
leaves long-petiolate, 3-foliolate; flowers axillary, slender-pedunculate, perfect;
calyx 5-parted, 5-bracteolate, the bractlets larger than the sepals and alternating
with them, dentate or incised; petals 5, yellow, obovate; stamens numerous; pistils
numerous, borne on a hemispheric receptacle, this greatly enlarging in fruit and
becoming pulpy; achenes superficial on the receptacle.

Two species, native in southern Asia.

Duchesnea indica (Andr.) Focke in Engler & Prantl, Pflanzen-
fam. 3, pt. 3: 33. 1888. Fragaria indica Andr. Bot. Rep. pi. 479. 1807.

Noted in Guatemala only as a weed in flower beds of the Jardin
Botanico, Guatemala. Native of Asia, widely naturalized in the
United States and in other temperate regions.

Plants green but somewhat sericeous; leaflets obovate to broadly oval, thin,
crenate or dentate, obtuse, rounded or narrowed at the base; peduncles equaling
or exceeding the leaves; flowers 12-20 mm. broad; calyx lobes ovate or lanceolate,
acuminate, green, spreading; fruit red, ovoid or globose, resembling a small
strawberry.

In general appearance the plant is like a species of Fragaria,
except for the yellow petals. Although the fruit looks exactly like a
strawberry, it is not edible, being insipid or even bitter in flavor.

ERIOBOTRYA Lindley. Loquat

Evergreen trees or shrubs; leaves short-petiolate or almost sessile, the
conspicuous veins ending in marginal teeth; flowers rather large, white, in terminal,
broad, usually lanate panicles; sepals acute; petals 5, oval to orbicular, unguicu-
late; stamens 20; styles 2-5, connate below; ovary inferior, the cells 2-ovulate;
fruit a pome with a thin endocarp, juicy, containing 1-2 large seeds, the sepals
persistent and incurved at its apex.

About 10 species, in eastern Asia.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 447

Eriobotrya japonica (Thunb.) Lindl. Trans. Linn. Soc. 13: 102.
1822. Mespilus japonicus Thunb. Fl. Japon. 206. 1784. Nispero;
Nispero japones; Nispero del Japdn.

Planted abundantly in many parts of the country, chiefly at
900-2,100 meters, but often at lower elevations, or slightly higher;
naturalized in many places in the Coban region. Native of central
China, but cultivated generally in tropical and subtropical regions
of other countries.

A small or medium-sized tree, usually 5-10 meters high; leaves almost or
quite sessile, coriaceous, obovate to oblanceolate, mostly 15-25 cm. long, acute
or acuminate, attenuate to the base, remotely and inconspicuously dentate,
glabrous and lustrous above, fulvous-tomentose beneath; flowers fragrant, 1 cm.
broad, arranged in large, dense, broad panicles, the branches densely rusty-lanate;
fruit pear-shaped, dull yellow, 3-4 cm. long; seeds 1-1.5 cm. long.

The tree is more abundant about Coban than in any other part
of Guatemala, but it is frequent also around Salama, and occasional
trees are found in most inhabited parts of the country. The fruit
is of excellent flavor and appeals particularly to the northern palate,
but is not so much esteemed by Central American people because of
its tartness. Although the tree grows at all elevations, it gives little
or no fruit in the lowlands. It does especially well about Antigua,
where the fruit is plentiful in market in October and November.

FRAGARIA L. Strawberry

Acaulescent perennial herbs with scaly rootstocks, producing long runners
that root at the nodes and produce new plants; leaves 3-foliolate, basal; hypan-
thium saucer-shaped, the bractlets, sepals, and petals each 5; petals normally
white, obovate to orbicular, obtuse; stamens about 20, in 3 series, the filaments
short, the anther cells dehiscent by longitudinal slits; receptacle hemispheric or
conic, bearing numerous pistils, in fruit becoming enlarged and juicy; styles
filiform but short, attached near the apex of the ovary, often persistent; seeds
ascending, amphitropous.

About 30 species, in the northern hemisphere, South America,
and East Indies. Probably none of the species are native in Cen-
tral America.

Leaflets thick and somewhat coriaceous, reticulate- veined; leaves usually taller
than the inflorescences; achenes mostly sunken in pits in the receptacle.

F. chiloensis.

Leaflets thin, not reticulate-veined; leaves usually shorter than the inflorescences;
achenes not sunken in pits F. vesca.

Fragaria chiloensis (L.) Duchesne, Hist. Nat. Frais. 165. 1766.
Fragaria vesca var. chiloensis L. Sp. PL 495. 1753. Fresa.

448 FIELDIANA: BOTANY, VOLUME 24

Suchitepe'quez (lower slopes of Volcan de Zunil, near Finca Las
Nubes, 500-800 meters; probably a relic of cultivation); cultivated
occasionally in the highlands for its fruit. Alaska to California;
Peru to Patagonia.

Plants with short thick rootstocks; stipules scarious, brown, 1-2 cm. long;
leaves long-petiolate, the petioles sericeous; leaflets thick and coriaceous, mostly
2-5 cm. long, glabrous and deep green above in age, densely sericeous-strigose
beneath and often tomentulose, the veins prominent and reticulate, crenate;
flowers mostly 2-3.5 cm. broad; hypanthium and calyx sericeous, the bractlets
and sepals oblong or lanceolate, acute or mucronate; petals broadly obovate, half
longer than the sepals; fruit hemispheric, red, 1.5-2 cm. in diameter or in cultivated
plants much larger; achenes set in shallow depressions.

Most of the strawberries grown in the United States are believed
to be derived from this species, although they probably are in part
of hybrid origin. The berries usually are much larger than those of
F. vesca and much more strongly flavored. This species was noted
in gardens at Tecpam and San Marcos, and doubtless is planted in
other parts of Guatemala.

Fragaria vesca L. Sp. PI. 494. 1753. Fresa; Fres (Quecchi).

Planted commonly in the highlands of Guatemala for its fruit;
thoroughly naturalized in San Marcos, Quezaltenango, and appar-
ently also in Alta Verapaz and Huehuetenango; 1,400-2,400 meters,
growing on open banks or in moist or wet meadows. Europe,
United States, and Canada.

Plants more slender and less vigorous than those of F. chiloensis, the root-
stocks short and thick; leaflets thin, sericeous on both surfaces when young but
glabrate in age, especially on the upper surface, often glaucous beneath, rhombic-
obovate, coarsely crenate-serrate; scapes several-flowered, the flowers 1-1.5 cm.
broad; sepals and bractlets ovate to lanceolate, acute, about 6 mm. long; fruit
usually subglobose or ovoid, red to whitish, 1-1.5 cm. in diameter, the achenes
superficial.

Strawberries are grown in small amounts in various regions of the
Guatemalan mountains, and often or usually can be purchased in
the markets of Guatemala and Quezaltenango. Most of the fruits
in market in Guatemala as well as in Costa Rica are of the vesca
type, which is little planted in the United States. The fruits differ
in shape from those of F. chiloensis, are less strongly flavored, and
usually are paler in color. Most of those offered for sale are small,
and in flavor somewhat suggest the wild strawberries so common in
the United States. It is said that formerly strawberries were grown
in large quantities on the slopes of Volcan de Agua for making
brandy. There is a little uncertainty about the identity of the

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 449

Guatemalan Fragarias. The plants seem to agree with F. mexicana
Schlecht. of Mexico, but they agree equally well with the ordinary
run of European specimens of F. vesca. Just how these two species,
if distinct, are to be separated is not evident. We have seen wild
plants of this type near Quezaltenango, where they grew in quantity,
but had every appearance of being escaped from cultivation. We
found them plentiful in pastures of Barranco Eminencia between
San Marcos and San Rafael Pie de la Cuesta in the Department of
San Marcos. There they were associated with various European
species, particularly grasses, and could have been either native or
introduced. Plants found in cloud forest in Sierra de los Cuchu-
matanes had every appearance of being native there, since they grew
with the normal wild plants of that region, remote from any human
habitation. It seems probable that they had escaped from some
previous plantation. Delicious strawberries are brought to Coban
by Indian peddlers, who state they are gathered from wild plants.
It seems more probable, however, that the plants have escaped from
cultivation, from some of the many German fincas of this region.
There is at present no conclusive evidence that the genus Fragaria
is native anywhere in Central America. It is worthy of note that in
the Flora of Jamaica Fawcett and Rendle consider Fragaria vesca a
native plant of that island, and under it they reduce to synonymy
F. insularis Rydb., described from Jamaica, as well asF. mexicana.

GUAMATELA Donn. Smith

Reclining shrubs; leaves opposite, simple, cordate-ovate, serrulate, palmate-
nerved; stipules free, setaceous; flowers perfect, in terminal racemes, the bracts
filiform, the flowers red or pink; hypanthium short, the 5 sepals imbricate; petals 5,
inserted in the mouth of the hypanthium; stamens 10 and 1-seriate, opposite
the petals and sepals, the filaments free, the anthers cordate-ovate, apiculate;
carpels of the ovary 3, sessile, at first connate by the stigmas, finally free, the
styles terminal, the stigmas capitellate; ovules several in each cell, biseriately
affixed to the ventral suture, ascending; seed-bearing carpel 1, membranaceous,
dehiscent by the ventral suture; seeds numerous, obovoid, without endosperm,
the testa osseous, lustrous.

The genus consists of a single species. The generic name is an
anagram of the word "Guatemala."

Guamatela Tuerckheimii Donn. Smith, Bot. Gaz. 57: 420.
1914.

Moist or wet, mixed, mountain forest, 1,750-2,400 meters,
endemic; Baja Verapaz (type collected near Purulha, Tuerckheim
3903); Zacapa (Sierra de las Minas).

450 FIELDIANA: BOTANY, VOLUME 24

I

A shrub 3 meters high, the stems branched, the branchlets, like the petioles
and nerves of the leaves, fuscous-pubescent; stipules 2-4 mm. long; leaves narrowly
long-acuminate, on petioles 5-25 mm. long, glabrous above, green, bullate, densely
white-tomentose beneath, 5-7-nerved, 4.5-9 cm. long, 2.5-5.5 cm. wide; racemes
drooping, solitary or geminate, with the peduncle 9-10 cm. long, white-tomentose,
about 10-16-flowered, the pedicels alternate, 4-5 mm. long; bracts binate, 6-8
mm. long; calyx red or rose-pink, the sepals oblong-ovate, acute, 7 mm. long,
striate; petals 5 mm. long, pink, oblong-elliptic, scarcely unguiculate, nerved,
pubescent on both surfaces; carpels lance-ovoid, the fertile one suboval, inflated,
7 mm. long, 5 mm. broad; seeds about 12, imbricate, 1.2 mm. long and thick.

This is one of the most remarkable of the plants localized in
Guatemala. The genus is a most distinct one, quite unlike any other
American member of the family. It probably is very rare, since it
has been collected but twice.

HIRTELLA L.

Shrubs or trees; stipules small, caducous; leaves alternate, simple, entire;
flowers small, perfect, in axillary and terminal panicles or racemes; calyx lobes 5,
reflexed; petals 5, deciduous; stamens 3-10 or more, the perfect ones on one side
of the receptacle, the staminodia on the other side, the filaments united at the
base, long-exserted; ovary 1-celled, inserted on one side of the receptacle, the style
almost basal; ovules 2; fruit drupaceous, slightly juicy; seed erect; cotyledons
carnose, the radicle inferior.

About 40 species in tropical America, one other in Madagascar.
One other species, H. media Standl., is known from Central America.
It occurs on the Atlantic coast of Honduras and in southern Mexico,
and is therefore to be expected in Guatemala.

Flowers in simple racemes, long-pedicellate; stamens 5; leaves sparsely appressed-

pilose or almost glabrous H. racemosa.

Flowers in thyrsiform panicles; stamens 3.

Leaf blades acute at the base; pubescence of the lower leaf surface of mostly

appressed hairs H. triandra.

Leaf blades obtuse to broadly rounded at the base; pubescence of the lower

leaf surface of spreading hairs.
Inflorescence and lower leaf surface densely pilose with very short, spreading

hairs H. americana.

Inflorescence and lower leaf surface hirsute with long spreading brown hairs,
the hairs of the lower leaf surface mostly confined to the costa.

H. paniculata.

Hirtella americana L. Sp. PI. 34. 1753. H. mollicoma HBK.
Nov. Gen. & Sp. 7: 263. 1825. H. guatemalensis Standl. Trop. Woods
11: 19. 1927 (type from Livingston, Izabal, Tuerckheim 11.1141).
Aceituno; Aceituno peludo (Pete"n).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 451

Wet forest or thickets, 500 meters or less; Pete'n; Alta Verapaz;
Izabal. Tabasco and British Honduras, along the Atlantic coast to
Panama; northern South America.

A shrub or tree, sometimes as much as 18 meters high, with a trunk to 25 cm.
or more in diameter; inner bark mulberry purple; leaves almost sessile, elliptic-
oblong, mostly 8-15 cm. long, acuminate, very obtuse at the base, thick and sub-
coriaceous, often somewhat rugose, glabrate above, densely velutinous-pilose
beneath; panicles narrowly thyrsiform or almost spike-like, chiefly 10-20 cm.
long, pedunculate, densely velutinous-pilose, the very numerous flowers sessile
or short-pedicellate; sepals 4 mm. long, oblong or oval, obtuse, densely sericeous
within; petals white; stamens purplish or deep red, 10-15 mm. long; fruit oval,
12-15 mm. long or larger, rounded at the apex, densely short-pilose.

Called "pasta" in Honduras; "pigeon plum" and "wild coco-
plum" in British Honduras. The wood in this genus is grayish or
light brown, straight-grained, hard, heavy, strong, tough, and moder-
ately durable. There is no report of any use made of it in Guate-
mala, although the bark is said to be suitable for use in tanning
and the wood for fuel, fence posts, and charcoal.

Hirtella paniculata Swartz, Prodr. Veg. Ind. Occ. 51. 1788.
Icaco de montana.

Wet mixed forest, near sea level; Izabal (Rio Frio, Cerro San
Gil, Steyermark 41615). British Honduras; West Indies and Guianas.

A tree 10 meters high or less, the trunk as much as 20 cm. in diameter, the
young branches hirsute with long spreading brownish hairs; leaves oblong or
elliptic-oblong, abruptly acuminate, obtuse to broadly rounded at the base, sub-
sessile, 11-20 cm. long, glabrate above, thinly hirsute beneath on the costa and
veins; panicles slender, narrow, short, densely hirsute, the flowers long-pedicellate;
fruit oval or subglobose, 1.5-2 cm. long, very densely brown-hirsute.

Called "achotillo" in British Honduras. The species has not
been reported previously from Central America. It is easy of recog-
nition because of the very long, brownish hairs on the leaves and
inflorescence.

Hirtella racemosa Lam. Encycl. 3: 133. 1789. Aceituno
Colorado (Pete'n); Uyamche (British Honduras, Maya); Manzanito
(fide Aguilar).

Wet or rather dry forest or thickets, sometimes in pine forest,
chiefly at 300 meters or less; Pete'n; Alta Verapaz; Izabal; Santa
Rosa; Escuintla; Retalhuleu. Southern Mexico and British Hon-
duras to Panama; West Indies; widely distributed in South America.

A shrub or small tree, sometimes 6 meters tall but usually only 2-3 meters,
the branches slender, pilose with short, mostly appressed, sometimes ascending

452 FIELDIANA: BOTANY, VOLUME 24

hairs; stipules small, subulate; leaves almost sessile, subcoriaceous, mostly oblong
or oblong-lanceolate and 5-7 cm. long, acute or short-acuminate, often with an
obtuse tip, obtuse or subacute at the base, usually strigose beneath along the
costa and nerves but elsewhere glabrate, lustrous; racemes axillary and terminal,
lax, many-flowered, often greatly elongate in fruit, the flowers on long slender
pedicels; bracts subulate, small and inconspicuous; sepals 3 mm. long or less;
petals deep pink or purplish; filaments purple; fruit oblong, rounded at the apex,
narrowed at the base, almost or quite glabrous, dark red or deep purple.

Names reported outside Guatemala are "wild coco-plum,"
"wild pigeon-plum" (British Honduras); "pasta" (Honduras);
"icaquillo" (Tabasco); "icaco months" (Salvador); "jicaquillo"
(Oaxaca). The bark is brown, the inner bark dark reddish brown;
sap wood pale yellow; heartwood reddish brown or reddish pink.
The fruit is rather sweet and edible but not attractive in flavor, and
the amount of flesh and juice is small. Some confusion has been
caused by the fact that for a time this shrub was known as H. ameri-
cana, a name now considered to belong to the species so called in the
present treatment.

Hirtella triandra Swartz, Prodr. Veg. Ind. Occ. 51. 1788.

Moist or wet, mixed forest, 350-1,260 meters; Alta Verapaz;
Suchitepe"quez. British Honduras to Panama; West Indies; northern
South America.

A large shrub or a tree, in Guatemala sometimes 15 meters tall with a trunk
15 cm. in diameter, the young branchlets strigose; leaves almost sessile, lance-
oblong to elliptic-oblong, mostly 5-15 cm. long and 2.5-5.5 cm. wide, rather
long-acuminate, acute or subobtuse at the base, strigose beneath along the costa;
panicles thyrsiform, rather dense and many-flowered, the branches often elongate;
sepals 3 mm. long, puberulent within, strigose outside; petals oval or broadly
oval, white, 5 mm. long; filaments pink or purple; fruit oblong-obovoid, as much
as 2 cm. long and 9 mm. broad, rather densely pilose, dark red or purple.

Known in British Honduras as "wild coco-plum" and "wild
pigeon-plum."

HOLODISCUS Maximowicz

Reference: Arline Ley, A taxonomic revision of the genus Holo-
discus, Bull. Torrey Club 70: 275-288. 1943.

Shrubs or small trees with sericeous pubescence; leaves alternate, simple,
dentate; stipules none; flowers small, white, perfect, racemose or paniculate;
hypanthium saucer-shaped or hemispheric; sepals 5, valvate in bud, erect in fruit,
3-nerved; disk somewhat developed, bearing about 20 stamens; petals 5, short-
unguiculate, the anthers didymous; pistils 5, alternate with the sepals, pubescent,
the styles terminal; ovules 2, collateral, pendulous; achenes enclosed in the calyx,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 453

short-stipitate, pilose, membranaceous, caducous; seeds broadly oblong; radicle
superior, the cotyledons ovate.

The species are confined to America, ranging from Colombia to
Alaska. Rydberg recognized 14, but the actual number is probably
not more than half as many and perhaps even fewer. Only one
species occurs in Central America.

Holodiscus argenteus (L. f.) Maxim. Acta Hort. Petrop. 6: 254.
1879. Spiraea argentea L. f. Suppl. 261. 1781. S. fissa Lindl. Bot.
Reg. 26: Misc. 73. 1840. Sericotheca fissa Rydb. N. Amer. Fl. 22:
265. 1908. S. velutina Rydb. loc. cit. H. argenteus var. bifrons
Focke in Bonn. Smith, Bot. Gaz. 18: 200. 1893. H. fissus Schneider,
Handb. Laubh. 1: 495. 1905. H. Loeseneri Dammer, Repert. Sp.
Nov. 15: 385. 1918 (type from Huitzan, Chiapas). H. argenteus
var. Matudai Ley, Bull. Torrey Club 70: 286. 1943 (type from
Volcan de Tacana, Chiapas, E. Matuda 2803). H. argenteus var.
alpestris Ley, op. cit. 288. 1943.

Higher mountains, in dense moist forest or on open brushy
slopes, sometimes in pine-oak or Juniperus forest, 2,100-3,500
meters; El Progreso; Sacatepe"quez; Chimaltenango; Solola; Quiche";
Huehuetenango; Totonicapan; Quezaltenango; San Marcos. South-
ern Mexico; Costa Rica; Colombia.

Usually a shrub of 1-3 meters but often somewhat larger and becoming a small
tree, the branches often slender and somewhat recurving, the bark loose, dark
gray or brownish, the young branchlets villous-tomentose; leaves small, short-
petiolate, oval to lanceolate or oblanceolate, glabrous and deep green above or
often densely whitish-sericeous, the nerves impressed, whitish-sericeous beneath,
acute or obtuse, cuneate at the base, crenate-serrate, the teeth mucronate; panicles
oblong to pyramidal, 5-15 cm. long, dense and many-flowered; sepals broadly
ovate, acuminate, 2 mm. long; petals broadly oval, 2.5 mm. long; bodies of the
carpels in fruit 2 mm. long, pilose, conspicuously rostrate.

Rydberg recognized about five species in this group that probably
are all forms of a single not very variable one. He relied upon the
pubescence (or lack of it) of the upper leaf surface for separation,
but this character seems unreliable and dependent primarily upon
the stage of development of the leaves. At best his species could be
recognized as varieties, although not well-marked ones. The shrub
is particularly characteristic of the Juniperus forests of the Cuchu-
matanes but it is plentiful enough in many other parts of Guate-
mala. As a rule, the panicles are small and not at all conspicuous,
so that in ornamental value this species is far inferior to some of
the handsome shrubs of the Rocky Mountains. The branches are

454 FIELDIANA: BOTANY, VOLUME 24

browsed by stock, and one often finds low and very dense bushes
that have developed as a result of long-continued browsing. The
plant was reported by Hemsley from Guatemala under the erroneous
name of Spiraea discolor Pursh.

LICANIA Aublet

Trees or shrubs; leaves alternate, simple and entire or nearly so, usually
coriaceous, persistent, often tomentose, commonly short-petiolate, the petiole
often 2-glandular at the apex; stipules subulate or lanceolate, deciduous or per-
sistent, free or connate; flowers small, perfect, sessile or pedicellate along the
branches of a panicle, 3-bracteolate; hypanthium urceolate to globose or hemi-
spheric, often hairy within; sepals 5, small, imbricate or subvalvate; petals 5,
sometimes none, small; stamens mostly 3-10, inserted in the throat of the hypan-
thium, included, sometimes unilateral, the filaments short, subulate or complanate,
often unequal, the anthers small; ovary immersed in the bottom of the hypanthium,
globose, 1-celled, villous or strigose, the style basilar, included in the calyx;
ovules 2, collateral, ascending from the base of the cell; fruit 1-seeded, coriaceous,
lignose, or crustaceous, often with abundant pulp, terete, obovoid to pyriform or
globose; seed erect, with membranaceous testa; cotyledons thick-fleshy, plano-
convex.

About 75 species, all American, most of them in South America.
One other species is known from Central America, in Costa Rica.

Leaves whitish beneath, covered with a closely appressed, whitish tomentum.

Leaves acute or acuminate, mostly 2-4 cm. wide L. hypoleuca.

Leaves rounded at the apex, mostly 5-10 cm. wide L. arborea.

Leaves green beneath, glabrous or nearly so, not tomentose.

Leaf blades mostly 20-30 cm. long, oblong or narrowly oblong. . . .L. platypus.

Leaf blades mostly 6-10 cm. long, lance-oblong or ovate-oblong. . .L. sparsipilis.

Licania arborea Seem. Bot. Voy. Herald 118. pi. 25. 1852-53.
L. Seleriana Loes. Verh. Bot. Ver. Brandenb. 53: 55. 1911. Encino;
Roble; Caca de nino, Zuncilla (fide Aguilar).

Chiefly in dry brushy forest, at 1,300 meters or less; Baja Vera-
paz; El Progreso; Chiquimula; Jutiapa; Santa Rosa; Escuintla;
Suchitepe"quez. Western Mexico; Salvador; Costa Rica; Panama.

A medium-sized or large tree, sometimes as much as 30 meters tall but usually
lower, the branchlets glabrate; leaves mostly oval to broadly oblong, 9-18 cm.
long, coriaceous, broadly rounded at the apex, obtuse to subcordate at the base,
glabrous and lustrous above, whitish beneath and covered with a minute close
tomentum, the lateral nerves slender, very prominent, the ultimate veins closely
reticulate; flowers small, white, in often large and much-branched panicles, these
brown-tomentose; sepals scarcely more than 1 mm. long; petals about 1.2 mm.
long and half as wide, ciliate, caducous; filaments villous; fruit obovoid or oblong-
obovoid, 2-3 cm. long or larger, containing a single large seed.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 455

Known in Salvador by the names "canilla de mula" and "jobo";
called "roble bianco" and "alcornoque" in Costa Rica; variously
known in Mexico as "cacahuananche," "frailecillo," "palo de fraile,"
and "totopostle." The tree is of some potential economic importance
because it is closely related to the oiticica tree (Lieania rigida Benth.)
of Ceara, Brazil, from whose seeds is extracted a commercially
important oil. This is exported in large quantities for use in manu-
facture of paints. There is every reason to suppose that the oil of
L. arborea may have the same properties and be of equal value if
available in sufficient quantity. Its seeds are said to contain about
30 per cent of oil and to burn readily. In western Mexico they are
sometimes strung on sticks and burned like candles. In that country
the oil has been used in substantial amounts for making candles,
soap, axle grease, and other articles. It is said to have a peculiar
odor, disagreeable flavor, and a greenish color that it imparts to
soap made from it. The wood is reported to be used at times for
heavy construction.

Lieania hypoleuca Benth. Bot. Voy. Sulph. 91. pi. 32. 1844.
Chozo.

Wet mixed forest, 450 meters or less; Pete"n; Alta Verapaz;
Izabal. Tabasco and British Honduras; Honduras; Costa Rica;
Panama (type from Veraguas).

A small to" large tree, sometimes 16 meters high with a trunk 25-40 cm. in
diameter, the branchlets slender, blackish; leaves on short petioles, mostly oblong-
ovate to elliptic-oblong and 5-9 cm. long, acute or acuminate, obtuse to narrowly
rounded at the base, rather thin, glabrous and lustrous above, covered beneath
with a very dense and fine, appressed, whitish tomentum; flowers very small, in
large or small, mostly pyramidal panicles, the individual flowers sessile or nearly
so, in small pedunculate cymules; sepals ovate, half as long as the hypanthium,
pubescent within; petals none; fertile stamens 2-3, small; fruit obovoid-pyriform,
attenuate at the base, rounded at the apex, about 2 cm. long, minutely tomentu-
lose, reddish.

The tree is plentiful near the north coast, sometimes attaining
considerable size. Frequently it is seen in pastures or in wooded
swamps. In British Honduras it is called "pigeon-plum." The
wood is brownish gray with a reddish hue, without distinctive odor
or taste, very hard and heavy (specific gravity about 1.03), of fairly
straight or somewhat roey grain, medium-textured, hard to cut, easy
to split, of dull surface when finished, apparently not resistant to
decay. It is sometimes employed for heavy construction.

456 FIELDIANA: BOTANY, VOLUME 24

Licania platypus (Hemsl.) Fritsch, Ann. Naturhist. Hofmus.
Wien 4: 53. 1889. Moquilea platypus Hemsl. Diag. PI. Mex. 1: 9.
1876. Sunza; Sunzapote; Coca de nino; Urraco (North Coast);
Sunco; Mox-pin (Quecchi); Chaute (fide Pittier, evidently an Indian
name, but the locality not indicated); Jolobob (Alta Verapaz).

Common in forests of the lowlands, especially of the North
Coast, also on the Pacific plains, chiefly at 400 meters or less; often
planted for ornament; Pete'n; Alta Verapaz; Izabal; Zacapa; Chi-
quimula; El Progreso; Jutiapa; Santa Rosa; Escuintla; Suchite-
pe"quez; Retalhuleu; Huehuetenango. Southern Mexico to British
Honduras and Panama; Colombia.

A medium-sized or large tree, sometimes attaining a height of 50 meters, the
bark pale, the branchlets glabrous; leaves somewhat distichous, on petioles 1-1.5
cm. long, oblong to narrowly oblong, mostly 6-8 cm. wide, acute or short-acumi-
nate, rounded at the base, deep green, slightly paler beneath, glabrous; inflores-
cence a large broad panicle, before anthesis provided with large deciduous purplish
bracts, 10-35 cm. long, whitish-pubescent, the flowers short-pedicellate or sub-
sessile; hypanthium and calyx whitish-puberulent, the hypanthium lanate within,
the sepals broadly ovate, acute, 2 mm. long; petals white, obovate, 2.5-3 mm.
long, acutish, ciliate; stamens 15-18, the filaments subequal, glabrous; ovary
densely pilose; fruits very large and heavy, 1-3 in each panicle, globose-obovoid,
15-20 cm. long, 10-14 cm. broad, dark brown and somewhat verrucose, covered
with white lenticels, the flesh deep yellow, juicy, sweet, somewhat fibrous; seed
usually 1, ovate-oblong, compressed, 6-8 cm. long, 4-4.5 cm. broad.

The tree has supplied the name for Sunzapote, a caserio of El
Progreso, and another in Zacapa. It is called "monkey-apple" in
British Honduras, and sometimes "sungano" in Salvador. This is
one of the finest and handsomest trees of Central America, often
towering to a great height in the forests and commonly planted in
fincas as a shade tree, a purpose that it serves admirably. The young
foliage is beautifully tinged with red or bronze, making the tree
conspicuous. The crown of the tree is normally very dense and of a
deep green. The fruit requires about a year for ripening and,
although edible, it is little esteemed, especially because there is a
popular belief that it causes fevers and other ailments. In Veracruz
the tree is sometimes called "zapote de mono." The trunk is reported
to have small buttresses sometimes; the sap wood is pale yellow or
pale yellowish brown, the heartwood reddish pink or pinkish brown;
the bark is about 2.5 cm. thick, the inner bark purplish brown.

Licania sparsipilis Blake, Contr. Gray Herb. 52: 67. 1917.

Moist or wet forest, 600 meters or less; Izabal. British Honduras,
the type from Sittee River, Peck 858.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 457

A tree 12-15 meters tall, the trunk 14-20 cm. in diameter, the branchlets
blackish or dark reddish brown, almost glabrous; leaves short-petiolate, small,
coriaceous, glabrous and lustrous, lance-oblong or ovate-oblong, mostly 2-3.5
cm. wide, acute, obtuse at the base, somewhat paler beneath; flowers fragrant,
white, in small pyramidal panicles, the stiff branches densely pilose with short
spreading whitish hairs, the flowers sessile; calyx and hypanthium densely short-
pilose, together 2.5 mm. long, the sepals broad, obtuse; stamens long-exserted ;
fruit globose, 2 cm. in diameter, broadly rounded at base and apex, green.

The wood is said to be red, hard, and close-grained.
MALUS Miller. Apple

Deciduous trees or shrubs, the branchlets sometimes spinose; leaves stipulate,
petiolate, serrate or lobate, folded or convolute in bud; flowers mostly white or
pink, fragrant, in umbelliform racemes, the petals usually suborbicular or obovate;
stamens 15-50, the anthers usually yellow; ovary inferior, 3-5-celled; styles 2-5,
connate at the base; fruit a pome, sometimes with a few grit cells, the calyx per-
sistent at its apex or deciduous.

About 25 species, in temperate regions of North America, Asia,
and Europe. Numerous species are native in the United States
but none extend so far southward as Mexico.

Malus pumila Miller, Card. Diet. ed. 8. No. 3. 1768. Pyrus
Mains L. Sp. PI. 479. 1753, in part. Manzano (tree); Manzana
(fruit); Manzdn (Quecchi).

Native of Europe and western Asia, the cultivated apples, how-
ever, of hybrid origin, their ancestry often doubtful. Apple trees
grown from seeds are planted generally in the mountains of Guate-
mala, chiefly at 1,500-2,400 meters or higher, and there are orchards
of grafted stock about Tecpam and elsewhere, but especially in the
vicinity of Cantel and Quezaltenango. We have not seen the better
apples produced at the latter places, but by competent judges they
are said to be superior to the apples imported from the United
States. The small fruit from seedling trees, usually of about the
size of northern crabapples, is offered commonly for sale in many
of the markets. As a rule the fruit is very sour and seldom is eaten
raw. The writers have noted flowers on trees of the Occidente in
January, February, and March. The lack of freezing weather in the
Guatemalan mountains upsets the normal routine of the apple tree
just as it does the peach. Some leaves remain on the trees through
most of the dry season, and blossoms and full-sized fruit often may
be seen upon the same tree, as we have noted about San Marcos.
Guatemala is the only Central American country in which apple

458 FIELDIANA: BOTANY, VOLUME 24

trees are much more than a curiosity. Pacific-coast apples from the
United States are imported in substantial quantities into Guatemala
as into other Central American countries, and are popular but
expensive. Most of these imported apples are no very favorable
advertisement for the United States, since they usually are over-ripe
when exposed for sale and hence inferior in flavor.

PHOTINIA Lindley

Trees or shrubs, glabrous or pubescent; leaves alternate, on short or long
petioles, coriaceous, persistent, simple, entire or serrate; stipules sometimes folia-
ceous; flowers small, white, perfect, in terminal panicles or corymbs; hypanthium
campanulate or turbinate; calyx lobes 5, ovate, obtuse; petals 5, spreading;
stamens 20 or fewer, inserted in the throat of the hypanthium, the filaments
subulate; ovary inferior or free at the apex, normally 2-5-celled, the styles 2-5,
free or somewhat connate below, the apices dilated and truncate; ovules 2 in each
cell, erect; fruit drupaceous or baccate, ovoid, 1-5-celled, the septa membranaceous
or chartaceous, the cells 1-2-seeded; seeds erect, with membranaceous or coria-
ceous testa; cotyledons plano-convex.

About 25 species, in Asia and North America. Only the following
species are known from Central America, but three others are found
in Mexico.

Leaves oblanceolate-oblong to obovate-oblong, broadest above the middle, incon-
spicuously crenate-serrate P. microcarpa.

Leaves lance-oblong to elliptic, broadest at or below the middle, entire or essen-
tially so P. Matudai.

Photinia Matudai Lundell, Contr. Univ. Mich. Herb. 4: 7.
1940.

Type from Chiapas, western slope of Volcan de Tacana, 2,800
meters, Matuda 2937; collected also at Siltepec and on Mount
Paxtal; doubtless extending into adjoining San Marcos; Huehue-
tenango (Cerro Canana).

A tree with stout branchlets, the young ones densely brownish-tomentose;
stipules 1 cm. long or less; leaves stiff-coriaceous, short-petiolate, 6-9.5 cm. long,
2-4.5 cm. wide, acute or subacute, acute at the base, usually entire, when young
densely brownish-tomentose but in age almost glabrous, lustrous, the lateral
nerves 8-13 pairs; inflorescence corymbose, about equaling the leaves, dense and
many-flowered, brownish-tomentose, the flowers sessile or short-pedicellate; sepals
triangular-ovate, 1.5-2 mm. long; petals suborbicular, 4 mm. long; fruit obovoid,
10-12 mm. long, becoming glabrate.

Further material is needed to determine whether this is really
distinct from the following species, which it closely resembles.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 459

Photinia microcarpa Standl. Carnegie Inst. Wash. Publ. 461:
57. 1935.

Moist or wet, mountain forest, 800-2,400 meters; Pete"n (type
collected at Camp 32 on the Guatemala-British Honduras boundary,
Schipp 1291; El Progreso(?); Zacapa (Sierra de las Minas); Hue-
huetenango (San Juan Ixcoy). British Honduras; Honduras, at
about 1,300-1,400 meters.

A small or large tree, 5-15 meters tall, the trunk up to 25-45 cm. in diameter,
the branchlets reddish brown, at first ferruginous-tomentose; stipules subulate,
minute, caducous; leaves on petioles 7-12 mm. long, narrowly oblanceolate-oblong
to obovate-oblong, mostly 5-9 cm. long and 1.5-3 cm. wide, acute or obtuse,
attenuate to the base, remotely crenate-serrate, especially toward the apex, or
subentire, glabrous above at least in age, slightly paler beneath, apparently brown-
tomentose at first but soon glabrate, the lateral nerves about 13 pairs; corymbs
shorter than the leaves, ferruginous-tomentose, short-pedunculate, dense, few-
flowered, the pedicels stout, 1.5 cm. long or less; sepals triangular-ovate, obtuse;
petals white, glabrous, twice as long as the sepals; fruit red, obovoid, 1 cm. long,
densely tomentose or glabrate.

POTENTILLA L.

Annual or perennial herbs, rarely shrubs, often with elongate, scaly, somewhat
cespitose rootstocks; leaves pinnately or digitately compound, the leaflets usually
dentate; flowers mostly cymose-paniculate, yellow, white, or purple; hypanthium
generally hemispheric; bractlets, sepals, and petals 5 each; petals deciduous,
broad, rarely unguiculate; stamens usually 20 and 3-seriate, the anthers didymous,
the filaments filiform or subulate; receptacle hemispheric or conic, bearing numer-
ous pistils, the styles short or elongate, inserted near the apex of the ovary, artic-
ulate with it and deciduous; seeds pendulous and anatropous.

A large genus of perhaps 250 species, widely distributed in both
hemispheres; 175 have been reported from North America. No
species are known in North America south of Guatemala, where they
are confined to the high mountains. The group is a temperate
rather than tropical one.

Leaves pinnate, with 3-7 leaflets P. heterosepala.

Leaves digitately compound, the leaflets all attached at the apex of the petiole.

Leaflets white-tomentose beneath P. staminea.

Leaflets green beneath, pilose but not at all tomentose P. Goldmanii.

Potentilla Goldmanii Painter ex Rydb. N. Amer. Fl. 22: 314.
1908.

In forest of pine and Juniperus, 3,700 meters; Huehuetenango
(Tojquia, Steyermark 50232). Mountains of Oaxaca, Mexico.

460 FIELDIANA: BOTANY, VOLUME 24

Perennial from an erect thick woody caudex, the stems usually several, 20-30
cm. high, erect, branched above, pilose with spreading hairs and somewhat
glandular-pubescent; basal leaves few or numerous, long-petiolate; leaflets 5,
obovate or oblong-obovate, 2-3 cm. long, rounded at the apex, broadly cuneate
at the base, rather coarsely crenate, pilose and glandular-atomiferous on the upper
surface, pilose beneath, green; flowers cream-colored or white, in rather open, few-
flowered cymes; hypanthium and calyx pilose and glandular-atomiferous, the
bractlets oblong, obtuse or subacute, about 5 mm. long, the sepals oblong-ovate,
slightly longer than the bractlets; petals 7 mm. long; stamens about 20.

Potentilla heterosepala Fritsch, Bot. Jahrb. 11: 314. 1890.
P. heterosepala var. guatemalensis Fritsch, op. cit. 315 (type from
Volcan de Agua, 3,900 meters, Scherzer). P. Donnell-Smithii Focke
in Bonn. Smith, Bot. Gaz. 16: 3. 1891 (type from Volcan de Agua,
3,600 meters, J. D. Smith 2144).

Open banks or more often in dense, coniferous or mixed forest,
mostly in the higher mountains, generally at 2,400-4,500 meters,
rarely as low as 1,500 meters, chiefly on the high peaks; collected as
a weed in cafetales near Antigua; Sacatepe"quez; Chimaltenango;
Solola; Quiche* ; Huehuetenango; Quezaltenango; San Marcos.
Southern Mexico.

Perennial, often forming dense clumps, with a thick caudex, the stems 20-50
cm. tall, leafy, decumbent or suberect, strigose and somewhat glandular; basal
leaves often very numerous, the 3-7 leaflets cuneate-obovate to broadly obovate
or oval, 3 cm. long and 2 cm. wide or smaller, obtuse or rounded at the apex, deeply
crenate or crenate-serrate, green and rather sparsely pilose on both sides, the
terminal leaflet long-stalked; inflorescence leafy, lax; hypanthium hirsute, the
bractlets 3-cleft, obtuse; sepals triangular, acute or short-acuminate, incurved in
fruit; petals bright or very pale yellow, cuneate-obovate, emarginate, twice as
long as the sepals.

The plant is a characteristic species of alpine summits of the
higher volcanoes, where it often grows in abundance, as it does also
in fir and Cupressus forests. The petals occasionally are almost
white, barely cream-colored. As compared with many species of the
genus, this is an inconspicuous and unattractive plant.

Potentilla staminea Rydb. Mem. Dept. Bot. Columbia Coll.
2: 67. 1898.

Grassy alpine meadows, Sierra de los Cuchumatanes, Huehue-
tenango, 3,250-3,700 meters. Mountains of southern Mexico.

Plants perennial from a very thick, somewhat woody taproot, the stems
stout, ascending or suberect, 30-60 cm. tall, leafy, villous-tomentose with mostly
spreading hairs; leaflets of the basal leaves 5-7, obovate-oblong, 3-10 cm. long,
broadly rounded at the apex, narrowed to the base, coarsely crenate-serrate,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 461

densely pilose but green above, densely white-tomentose and coarsely sericeous
beneath; cauline leaves with only 3-5 leaflets; cymes open and few-flowered, the
flowers long-pedicellate; hypanthium villous, the sepals ovate-lanceolate, 7-8
mm. long, acute or acuminate; petals yellow, broadly obcordate, 1 cm. long;
stamens about 40; styles filiform.

This also is an alpine plant. To this species presumably is
referable a report by Loesener of P. haematochrus Lehm. from the
Sierra de los Cuchumatanes. That is a red-flowered plant, similar
in general appearance to P. staminea, but no red-flowered Potentilla
has been found recently in Guatemala.

POTERIUM L.

Perennial herbs with rootstocks; leaves odd-pinnate, the leaflets dentate,
the stipules adnate to the petiole; flowers small, polygamo-monoecious, in very
dense, often head-like spikes; hypanthium urceolate, contracted at the mouth,
4-angulate; sepals 4, petaloid, deciduous, concave; petals none; stamens numerous
in the pistillate flowers, the filaments filiform, exserted and declined, fewer in the
perfect flowers; pistils 2, the styles terminal, the stigmas multifid and penicillate;
ovule solitary, suspended; fruit of dry achenes enclosed in the indurate, 4-angulate,
rugose or verrucose, woody hypanthium.

The genus consists of a single species, native of the Mediter-
ranean region.

Poterium Sanguisorba L. Sp. PI. 994. 1753. Pimpinela.

Grown in gardens of the Occidente, Quezaltenango and San
Marcos. Sometimes cultivated in eastern United States and spar-
ingly naturalized.

Plants glabrous or somewhat pubescent, 20-50 cm. high; leaflets 7-9, short-
petiolulate or subsessile, oval to orbicular, 1-2 cm. long, coarsely crenate-serrate,
green; spikes subglobose, 10-12 mm. in diameter; lower flowers staminate, the
upper perfect or pistillate; sepals oval, acute or apiculate, purple-tinged, 3.5-4
mm. long.

The plant is not a showy or handsome one, and it seems to be
planted in Guatemala chiefly for medicinal purposes. Bunches of
the stems and foliage are sold commonly in the Quezaltenango
market for medicinal use.

PRUNUS L.

Trees or shrubs, sometimes with thorns; leaves alternate, simple, mostly
serrulate, in bud complicate or convolute; flowers perfect, variously arranged,
white or pink; calyx deciduous or persistent, the tube obconic, urceolate, or tubular,
the 5 sepals imbricate; petals 5, inserted in the throat of the hypanthium; stamens

462 FIELDIANA: BOTANY, VOLUME 24

15-20, inserted with the petals, the filaments filiform, free; carpels solitary, the
style terminal, the stigma peltate or truncate; ovules 2, collateral; fruit drupa-
ceous, usually with juicy pulp, the stone osseous, smooth or rugose, sometimes dry
and bivalvate, 1-seeded; seed pendulous, the testa membranaceous; endosperm
scant or none; radicle superior.

Almost 200 species, in tropical and temperate regions of Europe,
Asia, and America. A few species not listed here are known from
Salvador, Costa Rica, and Panama.

Fruit and ovary pubescent; flowers sessile.

Fruit dry P. Amygdalus.

Fruit fleshy and juicy.

Stone of the fruit scarcely compressed, deeply pitted and furrowed.

P. Persica.

Stone of the fruit compressed, smooth P. Armeniaca.

Fruit and ovary glabrous.

Flowers not racemose, solitary or subumbellate.

Fruit somewhat sulcate, usually with a bloom, oval P. domestica.

Fruit not sulcate, without bloom, subglobose P. avium.

Flowers in racemes.

Racemes terminating short leafy branches; leaves finely serrulate.

P. Capuli.
Racemes axillary, leafless; leaves entire or serrulate.

Leaves serrulate P. Salasii.

Leaves entire.

Calyx persistent as a cupule beneath the fruit.

Axis of the raceme puberulent P. rhamnoides.

Axis of the raceme glabrous P. barbata.

Calyx deciduous.

Hypanthium glabrous within; rachis of the inflorescence glabrous.
Glands on the lower leaf surface 2, at the base of the blade close to

the costa P. brachybotrya.

Glands of the lower leaf surface 3-4, 2 of them near the base of the
blade close to the costa, the others near the lateral nerves.

P. Lundelliana.

Hypanthium pilose within; rachis of the inflorescence pubescent.
Leaves rather sparsely pilose beneath, the venation elevated and

conspicuously reticulate P. guatemalensis.

Leaves glabrous beneath, the venation neither elevated nor reticu-
late P. Skutchii.

Primus Amygdalus Stokes, Bot. Mat. Med. 3: 101. 1812.
Amygdalus communis L. Sp. PI. 473. 1753. P. communis Fritsch,
Sitzb. Akad. Wien 1892: 632. 1892, not Huds. Almendro. Almond.

Native of western Asia and northern Africa, cultivated since
ancient times in temperate and subtropical regions; planted and

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 463

fruiting in Guatemala City, and probably in other parts of the
country, although not of economic importance there.

A small tree, usually 8 meters high or less, with gray bark, the branchlets
glabrous; leaves on petioles 2.5 cm. long or less, ovate-lanceolate to narrowly
lanceolate, 7-12 cm. long, usually broadest slightly below the middle, long-
acuminate, broadly cuneate to almost rounded at the base, finely serrulate, gla-
brous; flowers solitary or in 2's, sessile, pink or almost white, 3-5 cm. broad;
calyx lobes oblong; fruit ellipsoid, slightly compressed, velutinous-pubescent, dry,
splitting along the margins; stone smooth but finely pitted.

Imported almond nuts are sold commonly in Guatemala. They
are probably brought from California, where the tree is cultivated
on a large scale, as it is also in the Mediterranean region. This tree
is placed by some authors in a separate genus, Amygdalus. Popenoe
states that California almonds were planted experimentally at Pana-
jachel, Solola, but after twelve years they had produced no fruit.

Primus Armeniaca L. Sp. PL 474. 1753. Armeniaca vulgaris
Lam. Encycl. 1: 2. 1780. Albaricoque. Apricot.

Planted occasionally in the highlands, but infrequently. Native
of western Asia, but in cultivation for many centuries in temperate
and subtropical regions.

A tree 10 meters high or less with rounded crown, the bark reddish, the branch-
lets brownish; leaves on petioles 2-3 cm. long, broadly ovate or orbicular-ovate,
5-10 cm. long, abruptly acuminate, subcordate or rounded at the base, closely
obtuse-serrate, glabrous or with tufts of hairs beneath in the nerve axils; flowers
solitary, white or pinkish, 2.5 cm. broad; fruit subglobose, yellowish with a reddish
cheek, pubescent at first but becoming glabrate; stone broad, compressed, almost
smooth, with a thickened edge.

The tree has been planted only experimentally in Guatemala.
It is stated that trees grown at Panajachel produced no fruit.
Dried and canned apricots from California are sold commonly in
delicatessen shops of the country.

Primus avium L. Fl. Suec. ed. 2. 165. 1755. Guinda; Cerezo.
Sweet cherry.

Native of Europe and western Asia, in cultivation since ancient
times and now grown in all temperate regions of the earth; planted
to a small extent in the highlands of Guatemala, especially about
Cantel and Quezaltenango.

Usually a tree of small or medium size with pyramidal crown ; leaves on petioles
4 cm. long or less, oblong-ovate, 6-15 cm. long, acuminate, serrate, more or less
pubescent beneath; flowers white, 2.5-3.5 cm. broad, in several-flowered umbels;
sepals usually entire; fruit subglobose, red, of rather firm texture, sweet.

464 FIELDIANA: BOTANY, VOLUME 24

The sour cherry, Prunus Cerasus L., also is probably in culti-
vation. It has usually smaller fruit of a brighter red color and sour
flavor. Cherries, like most of the other temperate fruits of the
Rosaceae, all so common in the United States, are practically un-
known in Central America except in the Guatemalan highlands.

Prunus barbata Koehne, Bot. Jahrb. 52: 284. 1915.

Type from Cumbre de Xuipach, Bernouilli & Cario 2916; Suchi-
tep^quez (lower slopes of Volcan de Zunil, southeast of Santa Maria
de Jesus, 1,300 meters); El Progreso (Sierra de las Minas, 2,500
meters) ; endemic.

A tree of 7-11 meters, glabrous throughout; petioles rather stout, 6-13 mm.
long; leaf blades oblong or oblong-lanceolate, 5-10 cm. long, 2-3.5 cm. wide, long-
acuminate, obtuse at the base and abruptly short-de cur rent, slightly paler beneath,
bearing near the base along the costa 2 small glands, the costa elevated beneath .
but the nerves and veins inconspicuous, not elevated; racemes solitary, axillary,
4-6 cm. long, rather densely many-flowered, the pedicels 2-3.5 mm. long; calyx
3 mm. broad, the sepals ovate, very obtuse; petals rounded-rhomboid, 2 mm.
long, white; stamens about 25; ovary glabrous.

Prunus brachybotrya Zucc. Abh. Akad. Muench. 2: 348.
1837. P. laurifolia Schlecht. Linnaea 13: 91. 1839; Escobo; Puc.

Moist forest, 500-2,700 meters; El Progreso(?); Quiche! (Finca
San Francisco, Cotzal); Huehuetenango. Southern Mexico.

A tree 9-12 meters high with a trunk 40 cm. in diameter, the bark rough and
furrowed, glabrous throughout; leaves on slender or stout petioles mostly 12-15
mm. long, lance-oblong or lance-oval, mostly 7-13 cm. long and 2-5 cm. wide,
acuminate or long-acuminate, rounded or very obtuse at the base, bearing 2 glands
beneath along the costa near the base of the blade, the slender costa elevated,
the nerves and veins not elevated, inconspicuous; racemes solitary in the leaf axils,
rather lax, many-flowered, 6 cm. long or less, the pedicels 4-7 mm. long; calyx 3.5
mm. wide; petals white, 1.5-2 mm. long; fruit globose, 1 cm. or more in diameter.

Prunus Capuli Cav. Anal. Hist. Nat. (Madrid) 2: 110. 1800.
P. salicifolia HBK. Nov. Gen. & Sp. 6: 190. pi. 563. 1823. Cerasus
Capollin DC. ex Seringe in DC. Prodr. 2: 539. 1825. P. Capollin
var. prophyllosa Donn. Smith, Bot. Gaz. 42: 293. 1906 (typeWrom
San Rafael, Sacatepe'quez, Maxon & Hay 3666). P. serotina var.
salicifolia Koehne, Deutsche. Dendrol. 305. 1893. Cerezo; Capulin;
Tup (Quiche").

Often planted about fincas; common at many places in the moun-
tains in pine or mixed forest, in many regions appearing as if escaped
from cultivation, in other places with the appearance of a native

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 465

tree, chiefly at 1,500-3,000 meters, rarely planted at lower elevations;
Alta Verapaz (planted about Coban); Guatemala; Sacatepe"quez ;
Chimaltenango; Solola; Quiche"; Totonicapan; Quezaltenango; San
Marcos. Mexico; naturalized in Ecuador and Peru.

A small or medium-sized tree, rarely 15 meters high with a trunk sometimes a
meter in diameter, the bark reddish brown or grayish, nearly smooth, the crown
usually broad; leaves rather thin, bright green, on slender petioles, these bearing
usually 2 glands near the apex; blades lanceolate to ovate, 6-18 cm. long, long-
acuminate, acute or obtuse at the base, closely serrate, glabrous or nearly so;
racemes usually elongate, bearing 1 or more leaves near the base, many-flowered,
lax, glabrous or sparsely pubescent, the flowers slender-pedicellate; petals white;
fruit red or almost black, 1 cm. in diameter or often larger, sweet.

The vernacular name appears in the names of two Guatemalan
caserios, El Cerezo in Quezaltenango and Los Cerezos in San Marcos.
The tree is of considerable economic importance in Guatemala
because of its fruit, great quantities of which are eaten and sold in
the markets during its rather limited season, beginning in late
April. In general appearance the fruit is strikingly like that of the
common sour cherry of the United States, although somewhat dark
in color. They are fully as large as the poorer varieties of sour
cherries, but their flavor is different, of course. However, they are
sweet and pleasant and scarcely suggest the bitter chokecherries of
the United States, with which this species has been confused by
some desk botanists without field knowledge of the Mexican and
Guatemalan tree. About Quezaltenango there is said to grow
occasionally a form with white or yellowish fruits. The fruits are
highly esteemed wherever the tree grows, and large quantities of
them are consumed in Mexico. The cherries attracted attention
from the earliest Spanish invaders, and are said to have been an
important food of Cortez' men at the time of the siege of Mexico
City in 1519. At a very early date the tree was introduced into
Peru, where it has become widely naturalized. It has even been
taken to be native there.

We do not know whether the tree is really native in Guatemala,
but it probably is native in the Occidente. Possibly seeds were
taken to Guatemala by the Mexican mercenaries who aided Pedro
de Alvarado, or they may have been transported by even -earlier
Indian traders. The seeds, like those of United States chokecherries,
are scattered by birds, which may account for the apparently wild
trees of the Occidente. Seldom if ever are trees found in what
may be assumed to be virgin forest. They are more plentiful about
Quezaltenango than elsewhere, and in the valley of that name they

466 FIELDIANA: BOTANY, VOLUME 24

are one of the most abundant of all trees. They shed all or most
of their leaves late in the dry season, the foliage turning red or yellow
before it falls. New leaves appear in late February, when their fresh
green color makes them conspicuous. The young foliage is often
tinged with pink or red. Saplings have been used successfully at
Chimaltenango as a stock upon which to graft the common Euro-
pean cherry.

The wood is said to be of good quality, and in Mexico it is used
for general carpentry and cabinetwork. The bark, leaves, and seeds
when crushed and in contact with water develop hydrocyanic acid
and under certain conditions may poison animals that eat them, as
in other species of Prunus. The bark and leaves are used in domes-
tic medicine. The bark of the closely related P. serotina Ehrh. of
the United States is official in the U. S. Pharmacopoeia, having
tonic properties and the power of calming irritation and diminishing
nervous excitability. In the United States the fruit is employed
also for flavoring spirituous liquors and non-intoxicating beverages.

Prunus domestica L. Sp. PI. 475. 1753. Ciruelo. Plum.

Native of western Asia and the Caucasus, in cultivation since
ancient times, now grown in all temperate regions; planted in some
quantity in the highlands of Guatemala, in Sacatepe"quez, Que-
zaltenango, and elsewhere.

Usually a small tree, 10 meters high or less, with narrow crown, the branchlets
glabrous or slightly pubescent; leaves on slender petioles 1.5-2.5 cm. long, elliptic
or obovate, 5-10 cm. long, coarsely crenate-serrate, pubescent beneath and reticu-
late-veined; flowers greenish white, strongly scented, 1.5-2 cm. broad; sepals
pubescent inside; fruit mostly ovoid or oblong, the stone almost free from the
flesh, nearly smooth.

The plums produced in Guatemala are said by competent judges
to be of good quality, and to be available in some quantity during their
season although they are not a common fruit. We have no data as to
varieties planted, and some of them may well be forms of the
Chinese P. salicina Lindl. Trees with dark purple foliage are planted
for ornament in the city of Quezaltenango. Plum trees in the fincas
of Alta Verapaz are said to bear well.

Prunus guatemalensis I. M. Johnston, Journ. Arnold Arb.
19: 118. 1938.

Moist or wet, mixed forest, 1,800-2,700 meters; endemic; type
collected at Chichavac, Chimaltenango, 2,400-2,700 meters, Skutch
504; Solola (Volcan de San Pedro).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 467

A tree 18 meters tall, the trunk 40 cm. in diameter, the young branchlets
sparsely puberulent; leaves on petioles 1-2 cm. long, oblong or lance-oblong, 10-19
cm. long, 4-8 cm. wide, entire, acute or acuminate, obtuse or rounded at the base,
coriaceous, glabrous above, pilose beneath, the veins conspicuous and reticulate;
racemes arising from defoliate nodes, solitary, 5-8 cm. long, puberulent, the
pedicels 3-5 mm. long, puberulent; hypanthium hemispheric, 2.5-4 mm. broad,
puberulent outside, villosulous within; petals white, broadly ovate, 2-3 mm.
long; sepals triangular, 1.3 mm. long; stamens 30-40; ovary sparsely villosulous,
soon glabrous.

Primus Lundelliana Standl. Field Mus. Bot. 22: 77. 1940.

Moist mixed mountain forest, 500-2,000 meters; Alta Verapaz
(?; sterile); Guatemala; Sacatepe"quez; Suchitepe"quez; Quezal-
tenango; San Marcos. Chiapas, the type from Hacienda Siltepec.

A glabrous tree; leaves on slender petioles 1-1.5 cm. long, lance-oblong to
elliptic-ovate, 7-15 cm. long, 2.5-7 cm. wide, long-acuminate, acute to rounded
at the base, rather thin and bright green, entire, brownish beneath when dry,
bearing 3-4 small glands remote from the costa, the venation inconspicuous, not
elevated; racemes axillary or from defoliate nodes, 3-5 cm. long, short-peduncu-
late, few-many-flowered, the slender pedicels 4-9 mm. long; calyx broadly cam-
panulate, 3 mm. broad and high, the sepals very short, broadly rounded; petals
white, 2 mm. long; ovary glabrous.

Primus Persica (L.) Stokes, Bot. Mat. Med. 3: 100. 1812.
Amygdalus Persica L. Sp. PI. 677. 1753. Durazno; Duraznal (the
tree); Doraz (Quecchi). Peach.

Native of China and cultivated since ancient times; grown in all
temperate regions; commonly planted in almost all mountain
regions of Guatemala, chiefly at 1,400-2,700 meters, but sometimes
at higher or lower elevations; abundantly naturalized in some regions.

A small tree, seldom more than 8 meters high, the branchlets glabrous; leaves
on petioles 1-1.5 cm. long, elliptic-lanceolate or oblong-lanceolate, broadest near
or slightly above the middle, 8-15 cm. long, long-acuminate, broadly cuneate at
the base, serrulate, glabrous, the petioles glandular; flowers usually solitary, pink,
2.5-3.5 cm. broad, almost sessile; sepals pubescent outside; fruit subglobose,
tomentose, the stone very hard and thick, not compressed, deeply pitted and
furrowed.

Most of the trees of Guatemala are seedlings, of inferior cling-
stone varieties, but especially in the orchards of Cantel and Que-
zaltenango the better budded varieties have been planted. The name
"prisco" is used for the freestone peach. Most of the dwellings of
the highlands have at least one or two peach trees, to provide fruit
for home use or more probably for market. Only in the case of the
intelligently managed orchards like those about Cantel is the fruit

468 FIELDIANA: BOTANY, VOLUME 24

ever allowed to ripen on the trees. In fact, although most people
of Guatemala and Costa Rica probably have seen peaches, few of
them ever have seen or. tasted a ripe one. It is said that if the fruit
is left on the trees, it is ruined by birds, mammals, or insects. The
green fruit, of course, is eaten only after having been cooked, and
while it is a welcome relief after the usual desserts, concocted from
such things as sweet potatoes and squash, it is very inferior in
quality and certainly would not be eaten in the United States, where
peaches never are eaten until fully ripe. Although in the Guate-
malan highlands the climate in its cycles resembles that of the North,
the winter weather is not severe enough to stop growth, with the
result that peach and other temperate trees seem bewildered and
sometimes behave in a manner that would be strange indeed in
the North. In the coolest regions, such as Quezaltenango and San
Marcos, peach trees lose all or most of their leaves during the winter
months, perhaps more because of dryness than cold, and everywhere
in Guatemala they shed their leaves some time before spring. While
most of the flowers open in January and February, peach blossoms
can be found at almost any time of the year, often hidden among
the green leaves and associated with nearly ripe fruit. It may be
remarked here that in the North the flowers always appear in earliest
spring when the trees have no leaves at all. The abundant peach
blossoms in January in such places as Quezaltenango and Totoni-
capan give a pleasing variation to the landscape, and recall to one
the pleasant spring months of the United States.

In the valley of Quezaltenango there are huge trees, apparently
very old. They are of interest because in the North peach trees are
short-lived and worthless after only ten years or so. One of the
lowest regions at which peaches are grown in Guatemala is that of
Coban, where the trees were noted in abundant flower in late March.
The fruit from the highlands is carried to most of the lowland
markets for sale, and fresh stewed peaches are served on the table
at such a remote place as Puerto Barrios. Substantial quantities of
canned peaches from the United States are sold in Guatemala, but
such canned fruit is so expensive that it is available only to the rich.
Canned fruit of almost all kinds may be purchased, however, and
one of the authors remembers to have eaten in one of the smaller
hotels cherry pie made from cherries canned in the United States, a
rare dish indeed in Central America. Because of the manner in
which peach trees occur in Guatemala, it would be impossible to
make an accurate census of them; but according to a report of the
Direction de Agricultura, the number of peach trees in the country

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 469

was estimated for 1938-39 at 81,249. The principal departments
producing them, in order of their importance, were Sacatep^quez
(13,796 trees), San Marcos, Huehuetenango, Totonicapan, Chimal-
tenango, Solola, and Quezaltenango.

Primus rhamnoides Koehne, Bot. Jahrb. 52: 283. 1915.

Usually in dense, mixed or Cupressus forest, 2,000-3,000 meters;
endemic; Chimaltenango; Quiche" (type from San Miguel Uspantan,
Heyde & Lux 3090); Huehuetenango (?; sterile); San Marcos.

A tree 9-15 meters tall with broad crown, the trunk sometimes 60 cm. in
diameter, the branchlets blackish or ferruginous, glabrous; leaves small, on petioles
4-10 mm. long, sometimes with 2 small glands beneath at the base near the costa,
lance-oblong or narrowly elliptic-oblong, mostly 5-8 cm. long and 1.5-2.5 cm.
wide, very long-acuminate, rounded or obtuse at the base, barbate beneath in the
nerve axils, otherwise glabrous, paler beneath, brownish when dry, entire, rather
thick, the lateral nerves conspicuous beneath and somewhat elevated; racemes
axillary, solitary, lax and few-flowered, the rachis very minutely puberulent, the
pedicels 2.5-5 mm. long, minutely puberulent; calyx 2-2.5 mm. broad, glabrous, the
sepals triangular; petals rounded, 2 mm. long, white; stamens 20; ovary glabrous;
fruit ovoid, dull reddish, 1 cm. long or slightly larger.

The species has been reported from Guatemala under the name
P. sphaerocarpa Swartz. Skutch reports it as a pollarded tree
growing in hedgerows along the trail between Nebaj and Aguacatan.

Primus Salasii Standl. Trop. Woods 32: 14. 1932. Carreto;
Carretero.

Moist mixed forest, 1,400-2,800 meters; endemic; often planted
as a shade tree; Jalapa (Volcan de Jumay); Guatemala; Sacatep£-
quez; Chimaltenango; Huehuetenango; Quezaltenango; San Marcos.

A glabrous tree 9-15 meters high or larger, the branchlets blackish or dark
reddish brown; leaves large, rather thick, on petioles 13-18 mm. long, the petiole
bearing 2 large glands near the apex; blades oblong-lanceolate to oblong-ovate,
mostly 8-17 cm. long and 3-8 cm. wide, long-acuminate, rounded at the base,
acutely appressed-serrate; racemes arising from defoliate nodes, 15-18 cm. long
or longer, laxly many-flowered, the pedicels 3-4.5 mm. long; hypanthium 3.5 mm.
broad, glabrous within, the sepals broadly ovate-triangular, obtuse, 1.5 mm. long;
petals white, broadly rounded, 4-4.5 mm. long; fruit subglobose or ovoid-
globose, usually about 1.5 cm. long and broad, becoming dark red at maturity,
with scant pulp and juice.

The tree is a well-known one in the central mountains, especially
about Antigua, where it is much planted in some of the coffee fincas.
It makes a handsome ornamental or shade tree and is grown for
this purpose in many localities. There are large trees in Central

470 FIELDIANA: BOTANY, VOLUME 24

Park of Guatemala City, and it was observed in the park of Chiantla
(Huehuetenango) as well as elsewhere. The wood is said to be of
good quality for cart construction. The fruits, although large and
handsome, unfortunately are useless, for their flavor is intensely
bitter.

Primus Skutchii I. M. Johnston, Journ. Arnold Arb. 19: 117.
1938.

Known only from the type, collected at Finca Moca, Suchite-
pe"quez, 1,140 meters, in forest on ridge, Skutch 2077.

A tree 36 meters high, the trunk 1.5 meters in diameter, covered with rough,
dark brown bark; leaves on petioles 1.5-2 cm. long, subcoriaceous, oblong or
elliptic-oblong, 12-15 cm. long, 6-9 cm. wide, acute or short-acuminate, rounded
or obtuse at the base, entire, paler beneath, bearing 2 glands close to the costa
at the base of the blade; racemes solitary from defoliate nodes, 5-9 cm. long,
sparsely puberulent, laxly many-flowered, the pedicels 8-12 mm. long, puberulent;
hypanthium 5-6 mm. broad, puberulent outside, pilose within, the sepals deltoid,
1.5 mm. long; petals white, 3 mm. long and wide; stamens about 30.

Pyracantha crenulata Roemer, native of eastern Asia, is in culti-
vation in the Jardin Botanico of Guatemala City, and may be found
elsewhere. It is a shrub with small racemes of white flowers, the
fruit a small pome, the leaves about 2 cm. long, coriaceous, glabrous
or nearly so, and finely serrulate.

PYRUS L. Pear

Deciduous trees or shrubs, sometimes thorny; leaves petiolate, serrate or
entire, involute in bud, stipulate; flowers appearing with or before the leaves,
in umbelliform racemes, white or rarely pinkish; sepals commonly reflexed or
spreading; petals unguiculate, orbicular to oblong; stamens 20-30, the anthers
usually red; styles 2-5, green; ovules 2 in each cell; fruit normally a pyriform
pome, the flesh with numerous grit cells, the cell walls cartilaginous; seeds black
or nearly so.

About 20 species, all natives of the Old World.

Pyrus communis L. Sp. PL 470. 1753. Pera; Peral. Pear.

Cultivated in the mountains at 1,500 meters or higher, especially
in Sacatepe"quez and Quezaltenango, also occasionally in the moun-
tains of Alta Verapaz. Native of Europe and western Asia, in
cultivation since ancient times.

A small or medium-sized tree with pyramidal or narrow crown, sometimes
thorny, the young branchlets glabrous or sparsely pubescent; leaves on slender

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 471

petioles 1.5-5 cm. long, orbicular-ovate to elliptic, 2-8 cm. long, acute or short-
acuminate, subcordate to broadly cuneate at the base, crenate-serrulate, glabrous
or villous when young; inflorescence villous or almost glabrous, the pedicels 1.5-3
cm. long, the flowers about 3 cm. broad.

It is only in Guatemala that the pear is cultivated to any impor-
tant extent in Central America, and even here the fruit is not at
all common although it is offered frequently in the markets of Guate-
mala and Quezaltenango. The fruit was noted as plentiful in the
Guatemala market in late April, but it was hard and green. Trees
near Coban were in bloom in early April. Handsome and delicious
pears are said to be produced about San Bartolo and Santa Lucia
Milpas Altas.

ROSA L. Rose

Shrubs, deciduous or evergreen, sometimes scandent or trailing, usually
prickly; leaves alternate, stipulate, mostly odd-pinnate; flowers solitary or corym-
bose at the ends of short branchlets; sepals and petals each 5, variously colored;
stamens numerous; pistils numerous, enclosed in a usually urceolate receptacle,
this becoming fleshy and berry-like at maturity and enclosing several or many
osseous achenes.

Perhaps 200 species, almost all in the northern hemisphere. In
America the genus reaches its southern limit of distribution near
Mexico City.

Styles united to form a column, usually about as long as the stamens.

R. multi flora.
Styles free, about half as long as the stamens R. chinensis.

Rosa chinensis Jacq. Obs. Bot. 3: 7. 1768. R. Montezumae
Bertol. Fl. Guat. 423. 1840 (described from Volcan de Agua, Velas-
quez}. R. indica Auct., not L. Rosa.

Native of China. To this species probably belongs the majority
of the garden roses of Guatemala, of which there are many varieties,
some of them doubtless in cultivation in the country from early
colonial days. Roses are one of the favorite flowers of Guatemala
and they thrive in almost all parts of the country, from sea level
high into the mountains. Finest of all are those of Coban, where
the climate, cool and moist, seems to be exactly right for their best
growth; but there are handsome displays of roses in many other parts
of the country, in gardens of rich and poor. Bushes probably refer-
able to R. chinensis have run wild in some parts of the Pacific foot-
hills and may be found established in hedges in other parts of
Guatemala.

472 FIELDIANA: BOTANY, VOLUME 24

In Guatemala, Coban, and elsewhere there have been introduced
some of the finer varieties of roses from Europe and the United
States, and large quantities of the blossoms are on sale in the princi-
pal markets. Many of the varieties have well-established local
names. Among the handsomest of all is the rose called "la reina"
or "bola de nieve," with huge, double, pure-white blossoms. It is
seen almost everywhere, but it thrives best where there is abundant
moisture, as at Coban and in the foothills near Mazatenango and
Retalhuleu. A striking and handsome rose is a large vine with
clusters of small, double, bright-yellow flowers. It is not abundant
but is found occasionally in the higher mountains, especially in
Totonicapan and Huehuetenango and westward. Vines of the roses
known in the United States as "ramblers" are of frequent occurrence
in Guatemala. Tea roses are but little grown.

Rosa multiflora Thunb. Fl. Japon. 214. 1784. Rosa.

Native of Japan and Korea. Probably brought to Guatemala in
early colonial days from Spain; little cultivated at present but
thoroughly naturalized at many places in the higher mountains,
from Chimaltenango westward to Huehuetenango and Quezalte-
nango; Alta Verapaz; Chimaltenango; Totonicapan; Huehuetenango;
Quezaltenango; San Marcos; growing in thickets, especially in
hedges.

A stout shrub, abundantly armed with stout prickles, suberect or often
scandent over shrubs and low trees; leaflets usually 9, obovate to oblong, 1.5-3
cm. long, acute or obtuse, serrate, pubescent; flowers usually corymbose, about
3 cm. broad, deep pink to almost white.

This is apparently the same rose that has become so thoroughly
naturalized in the mountains of Costa Rica, where it is called "rosa
de Castilla." Although not a plant that is much to be admired in
cultivation, it is rather attractive when seen along the hedges of the
mountain roads, where often it occurs in great abundance. It is
particularly plentiful in Huehuetenango, and also about Tactic in
Alta Verapaz.

RUBUS L.

Reference: Wilhelm Olbers Focke, Species Ruborum. Mono-
graphiae generis Rubi Prodromus, Bibl. Bot., Hefte 72, 73. 1910-14.

Shrubs or rarely herbs, erect or often scandent or trailing, usually armed
with prickles; leaves alternate, simple, 3-foliolate, or pinnately or pedately com-
pound, with stipules; flowers perfect, white or pink, in racemes, corymbs, or

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 473

panicles or solitary, chiefly terminal; sepals 5, persistent; petals 5, sometimes none;
pistils few to many, borne on a convex torus, the styles subterminal; mature
carpels normally drupelets, juicy, occasionally dry.

More than 400 species, chiefly in temperate and cold regions
of the northern hemisphere, but numerous species present in tropical
mountains in both North and South America. Besides the species
enumerated below, a European raspberry with pale yellow fruits is
planted occasionally about Coban, where it produces well, and also
in other parts of Guatemala. Bushes seen at Coban in April were
loaded with fruit. It is probably a form of R. idaeus L. A European
blackberry also is planted and thriving in the Coban region and
doubtless in other parts of the country, and it may well be that some
of the varieties cultivated in the United States, originating from
native American species, have been introduced into Guatemala.

Plants unarmed; leaves simple R. trilobus.

Plants armed with prickles; leaves compound.

Leaves pinnate, with 5-15 leaflets; flowers double R. rosaefolius.

Leaves 3-foliolate or pedately compound; flowers not double.

Drupelets united to form a thimble-shaped aggregate fruit, this falling entire
from the dry receptacle; leaflets white-tomentose beneath.

Fruit hemispheric; sepals enclosing the fruit or spreading R. Pringlei.

Fruit oval or oblong; sepals reflexed in fruit.

Leaflets glabrous or nearly so on the upper surface R. glaucus.

Leaflets puberulent on the upper surface R. eriocarpus.

Drupelets remaining on the fleshy receptacle at maturity, or falling off
together with the receptacle, or falling off separately.

Stems and petioles densely hispid with long glandless hairs . . R. urticaefolius.

Stems and petioles not hispid, or hispid with gland-tipped hairs.
Stems bearing gland-tipped hairs.

Leaflets simply serrate, the teeth very short and salient; inflorescence
racemose R. miser.

Leaflets duplicate-serrate, the teeth lanceolate, directed forward;
inflorescence paniculate.

Sepals 1.5-2 cm. long or more R. leptosepalus.

Sepals much shorter.

Hairs of the stem mostly 2-5 mm. long; drupelets glabrous.

R. adenotrichus.

Hairs of the stem short, rarely more than 1 mm. long; drupelets
sparsely or densely pubescent at the apex.

Mature fruit 1-1.5 cm. long; inflorescence elongated, loosely
much branched, pyramidal, 15-20 cm. long, 6-15 cm. broad;
secondary nerves on lower surface of leaflets not prominent.

R. irasuensis.

Mature fruit 1.5-2.2 cm. long; inflorescence short, dense, con-
tracted, narrowly oblong, 3.5-6 cm. long, 3-5 cm. broad,
secondary nerves on lower surface of leaflets prominent.

R. hadrocarpus f. adenophorus.

474 FIELDIANA: BOTANY, VOLUME 24

Stems without gland-tipped hairs, the inflorescence sometimes glandular-
pubescent.
Inflorescence usually densely prickly; stems often scandent.

Leaflets simply serrate, very lustrous on the upper surface, glabrate

beneath R. fagifolius.

Leaflets duplicate-serrate, dull on the upper surface, densely pubes-
cent beneath.

Pedicels 1.5-2 cm. long; mature fruit subglobose, 1-1.2 cm. long.

R. sapidus.

Pedicels 0.2-1 cm. long; mature fruit oblong, 1.5-2.2 cm. long.

R. hadrocarpus.
Inflorescence usually without prickles; stems not scandent.

Leaflets glabrous beneath or nearly so, pubescent only along the

veins, if at all R. alpinus.

Leaflets densely pubescent beneath.

Leaflets simply serrate with small short teeth.

Inflorescence glandular as well as tomentose; leaflets more or
less pilose with simple hairs on the upper surface.

R. macrogongylus.

Inflorescence not glandular; leaflets finely pubescent on the
upper surface with branched hairs R. Smithii.

Leaflets duplicate-serrate with lanceolate teeth conspicuously
directed forward.

Drupelets glabrous; inflorescence elongated, .much branched,
12-20 cm. long, 6-10 cm. broad; secondary nerves on lower
surface of leaflets not prominent R. coriifolius.

Drupelets villous; inflorescence short, dense, contracted, narrow,
3.5-6 cm. long, 3-5 cm. broad; secondary nerves on lower
surface of leaflets prominent R. hadrocarpus.

Rubus adenotrichus Schlecht. Linnaea 13: 267. 1839. Zarza-
mora; Mora; Tocdn (Quecchi and Poconchi).

Moist or wet thickets or hillsides, rarely in rather dry places,
often in oak or pine forest, 1,200-2,500 meters; Alta Verapaz; Baja
Verapaz; El Progreso; Zacapa; Jalapa; Escuintla; Guatemala;
Chimaltenango; Solola; Quiche"; Huehuetenango; Totonicapan;
Quezaltenango; San Marcos. Central and southern Mexico; Hon-
duras; Costa Rica; Panama; Colombia and Ecuador.

Stems mostly 1-3 meters high, arching, densely covered with long stiff gland-
tipped hairs, also densely short-pilose, armed with curved prickles; leaves digi-
tately 5-foliolate, the upper ones 3-foliolate, the petioles glandular-setose and
aculeate; leaflets ovate to elliptic or obovate, mostly 5-10 cm. long and 3-5 cm.
wide, acuminate, rounded or subcordate at the base, rather thin, acutely and
rather finely duplicate-serrate, sparsely pilose above, paler and densely pilose
beneath, often glandular-hispid and aculeate on the costa; flowers in pyramidal
panicles, the branches densely glandular-hispid and pilose; sepals ovate, subulate-
acuminate, about 1 cm. long, white-pilose and glandular; petals white or pink,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 475

1 cm. long; fruit ovoid, red or at maturity black, about 1 cm. thick, the drupelets
numerous, glabrous.

The species may be recognized easily by the abundant, long,
spreading, gland-tipped hairs. The fruit is usually sour but some-
times rather sweet. The fruit of this and other blackberries is much
gathered in Guatemala and often is sold in quantity in the markets.
Usually it is served stewed. In some of the species the seeds are
large and troublesome when the fruit is eaten, but in other species
the seeds are small and relatively inconspicuous. At Coban the
juice is used to make a wine-colored fresco or beverage, called there
"srub," presumably a corruption of the English "shrub." The raw
fruit is eaten in large amounts from the bushes by the country
people. A decoction of the root of various species of Rubus is
employed in Guatemala as a household remedy for dysentery.

Rubus alpinus Macfad. Fl. Jam. 2: 7. 1850. R. superbus
Focke in Donn. Smith, Bot. Gaz. 18: 210. 1893 (type from San
Miguel Uspantan, Quiche", Heyde & Lux 3326). Mora.

Damp or wet, mountain thickets or in open fields, 750-3,000
meters; Alta Verapaz; Baja Verapaz; Quiche"; Suchitepe"quez;
Quezaltenango; San Marcos; Huehuetenango. Costa Rica and
Panama; Jamaica; Colombia and Guianas.

A suberect or arching shrub or often a large vine over shrubs or trees, the stems
terete, usually purplish, glabrate, armed with recurved prickles; leaves pedately
3-5-foliolate, the petioles sparsely aculeate; leaflets rather thin, or coriaceous,
bright green, elliptic, mostly 6-10 cm. long, abruptly acuminate, rounded or
subcordate at the base, unequally and acutely serrate, glabrous on both sides or
sparsely pubescent beneath, the lateral nerves salient beneath, 10-13 on each side;
flowers paniculate, the panicles small or large and pyramidal, often densely pubes-
cent; sepals lanceolate, cuspidate-acuminate, tomentose, usually appressed to the
fruit; petals pure white, obovate, 1 cm. long or shorter; drupelets numerous,
glabrous, falling off separately.

Rydberg separates R. superbus from R. alpinus mainly by the
larger petals. In this genus, at least among Central American
species, size of petals scarcely can be considered a good specific
character.

Rubus coriifolius Liebm. Vid. Medd. 1852: 157. 1853. R.
floribundus f. laxiflora Focke in Donn. Smith, Enum. PI. Guat. 4: 54.
1895, nomen. R. floribundus f. pauciflora Focke, loc. cit., nomen.
R. laxus Rydb. N. Amer. Fl. 22: 454. 1913 (type from Zamorora,
Santa Rosa, Heyde &Lux 4474). Mora.

476 FIELDIANA: BOTANY, VOLUME 24

Moist or dry thickets, often in pine-oak forest, 1,600-2,400
meters; Santa Rosa; Guatemala; Chimaltenango; Solola; Huehue-
tenango; Quezaltenango; San Marcos. Southern Mexico.

Plants suberect or arching, or often scandent or sprawling over other shrubs,
the stems subterete, densely pubescent, rather sparsely aculeate; leaves coria-
ceous, pedately 3-5-f oliolate, the petioles densely pubescent, aculeate or unarmed ;
leaflets elongate-ovate or oblong-elliptic, mostly 6-10 cm. long, abruptly acuminate
or caudate-acuminate, obtuse or rounded at the base, acutely duplicate-serrate,
puberulent above, densely soft-pilose beneath, the nerves very prominent; flowers
in terminal and axillary panicles, these small or often large, densely tomentose,
often glandular-pilose; sepals ovate, acute or acuminate, tomentose, reflexed in
age; petals white or pale pink, longer than the sepals; fruit small, black when ripe,
glabrous, the drupelets 8-30, usually falling apart separately.

This has been reported from Guatemala as R. floribundus HBK.
(R. abundus Rydb.), which Focke considers to be confined to South
America.

Rubus eriocarpus Liebm. Vid. Medd. 1852: 162. 1853. Mora.

Moist thickets or pine-oak forest, 2,000-4,000 meters; Zacapa;
Sacatepe*quez; Chimaltenango; Huehuetenango; Quezaltenango.
Central and southern Mexico; reported from Panama.

An erect or subscandent shrub, the stems usually glabrous, subterete, glaucous-
pruinose, armed with numerous small prickles; leaves mostly 3-f oliolate, the
petioles glabrous, aculeate; leaflets ovate or lanceolate, 6-10 cm. long or smaller,
acuminate, rounded or cordate at the base, puberulent above, densely white-
tomentose beneath, finely duplicate-serrate; corymbs terminal, few-flowered,
tomentose and sparsely aculeate; sepals lanceolate, long-acuminate, 5-6 mm. long,
reflexed in fruit; petals elliptic, white, shorter than the sepals; fruit oblong or
subglobose, 10-12 mm. long, 6-8 mm. thick, the drupelets numerous, villous-
tomentose.

This is too closely related to R. glaucus, of which it may be only a
form, at least as far as Guatemalan material is concerned.

Rubus fagifolius Schlecht. & Cham. Linnaea 5: 571. 1830.

Moist or wet forest, 250-800 meters; Pete"n (British Honduras
boundary); Alta Verapaz (west of Cubilgiiitz). Southern Mexico,
the type from Papantla, Veracruz; British Honduras.

Scandent over trees, the stems sometimes 18 meters long, the young branches
sulcate, puberulent, armed with compressed recurved prickles; leaves coriaceous,
pedately 3-5-f oliolate, the petioles puberulent, retrorse-aculeate; leaflets elliptic
or elliptic-oblong, mostly 8-12 cm. long, caudate-acuminate, obtuse or rounded
at the base, deep green and lustrous above, glabrous, dull beneath and pubescent
on the veins, acutely serrulate, the lateral nerves conspicuous beneath, 12-15
on each side; panicles terminal and axillary, densely pubescent, unarmed or

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 477

aculeate; sepals ovate, pilose or glabrate, reflexed in age; petals white; fruit small,
red, the drupelets usually only 4-6, falling apart separately, pilose at first but
glabrate.

Rubus glaucus Benth. PL Hartweg. 173. 1845. Mora; Tocdn
uuc (Quecchi).

Moist or wet thickets or open fields, 1,200-3,000 meters; Alta
Verapaz; El Progreso; Zacapa; Huehuetenango; Quezaltenango;
San Marcos. Costa Rica; Panama; southward to Ecuador.

Plants usually erect or arching and 1-2.5 meters tall, the stems glabrous,
glaucous-pruinose, armed with rather small, compressed prickles; leaves usually
all 3-foliolate, the petioles and often the midnerves of the leaflets (beneath)
aculeate; leaflets thin, ovate or ovate-lanceolate, 6-15 cm. long, acuminate,
rounded or subcordate at the base, finely duplicate-serrate, bright green and
glabrous above, densely and closely white-tomentose beneath; inflorescences few-
flowered, terminal or in the upper leaf axils, the branches tomentose and sometimes
glandular; sepals lanceolate, 6-7 mm. long, gradually acuminate, densely tomen-
tose, reflexed in fruit; petals white, equaling the sepals; fruit red-purple or dark
purple, 12-20 mm. long, 8-15 mm. thick or larger, the drupelets numerous, very
juicy, tomentose when young.

This shrub produces one of the best fruits of the whole earth,
and it is unfortunate that it has not been introduced into cultivation
in frost-free regions where it might thrive. The fruit is quite differ-
ent in flavor from blackberries and is not too much like raspberries,
to which it is related. It suggests more the loganberry of the United
States, but we agree with Wilson Popenoe who states that in flavor
it is superior to that fruit. The seeds are surprisingly small and
unobtrusive. The "mora blanca," as it is called in Costa Rica, is
more abundant there than in Guatemala, and its fruit is highly
esteemed locally. Wherever they grow, the bushes are conspicuous
because of the pale canes and the white under-surface of the leaves.

Rubus hadrocarpus Standl. & Steyerm., sp. nov. Mora.

Wet thickets and damp forested slopes, 2,100-3,000 meters;
endemic; Solola; Quezaltenango; San Marcos; Huehuetenango.

A mostly subscandent shrub, 1-2 meters high, the stems subterete, finely and
usually densely pubescent, aculeate; leaves subcoriaceous, pedately 3-5-foliolate,
the petioles finely and densely pubescent, prominently aculeate; leaflets broadly
ovate or oblong-elliptic, 6-12 cm. long, 3.5-6 cm. wide, abruptly acuminate or
caudate-acuminate, obtuse or rounded at the base, acutely duplicate-serrate,
puberulous above, rather densely pilose on the main and secondary nerves beneath,
the lower surface with prominent nerves; flowers in terminal and axillary panicles,
these usually contracted, small and narrow, 3.5-6 cm. long, 3-5 cm. broad, densely
tomentose, sometimes aculeate near the base, the pedicels very short, 2-10 mm.

478 FIELDIANA: BOTANY, VOLUME 24

long; sepals ovate, acuminate or caudate, 5-11 mm. long, tomentose, reflexed in
age; petals white, 7-8 mm. long, about equaling the sepals in anthesis; fruit large,
in compact clusters, black when ripe, sour, broadly oblong, the mature fruit
1.5-2.2 cm. long, 1-1.2 cm. broad; drupelets numerous, 50-75, villous.

Frutex subscandens, ramis plerumque densissime pilosulis, aculeatis; folia
3-5-foliolata; foliola late ovata vel oblongo-elliptica, 6-12 cm. longa, 3.5-6 cm.
lata, acute duplicato-serrata, supra puberula, subtus prominente nervata, venis
dense pilosulis; paniculae terminales et axillares contractae breves angustae,
3.5-6 cm. longae, 3-5 cm. latae, pedicellis 2-10 mm. longis; fructus oblongus,
1.5-2.2 cm. longus, 1-1.2 cm. latus, carpellis numerosis, 50-75, apice pubescenti-
bus.

GUATEMALA: Dept. San Marcos: Barranco Eminencia, road
between San Marcos and San Rafael Pie de la Cuesta, in upper part
of the barranco between Finca La Lucha and Buena Vista, alt.
2,500-2,700 meters, February 6, 1941, Paul C. Standley 86270
(type in Herb. Chicago Nat. Hist. Mus.).

This is well marked among Mexican and Central American
species of Rubus by the relatively short, narrow, contracted inflores-
cences together with the short pedicels and large villous fruits. The
prominent secondary nerves on the lower leaf surface are also very
characteristic. It is most closely related to R. sapidus and to R.
coriifolius. From the former it may be distinguished by its larger
fruits and shorter pedicels, while from the latter it may be separated
by its shorter and narrower inflorescences and villous drupelets.

Rubus hadrocarpus, forma adenophorus Standl. & Steyerm.,
f. nov.

Known only from the type, in wet cloud forest at Cruz de Limon,
between San Mateo Ixtatan and Nuca, Sierra de los Cuchumatanes,
Huehuetenango, 2,600-3,000 meters, Steyermark 49859.

A subscandent shrub about 2 meters high, the stems densely ferruginous-
pubescent with long gland-tipped hairs 1-2 mm. long, aculeate; petioles densely
ferruginous-glandular-setose and aculeate; leaflets sparsely pilose above, paler
and more densely pilose-glandular on the main and secondary nerves beneath,
the costa usually aculeate; panicles densely ferruginous-glandular-pilose; in all
other respects similar to the species.

A forma typica speciei differt caulibus, petiolis, paniculisque glanduloso-
pubescentibus; foliola subtus venis dense glanduloso-pilosa.

GUATEMALA: Dept. Huehuetenango: Wet cloud forest at Cruz de
Limon, between San Mateo Ixtatan and Nuca, Sierra de los Cuchu-
matanes, alt. 2,600-3,000 meters, July 31, 1942, Julian A. Steyer-
mark 49859 (type in Herb. Chicago Nat. Hist. Mus.).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 479

This was found growing with the species (Steyermark 49860),
from which it differs only in the glandular pubescence.

Rubus irasuensis Liebm. Vid. Medd. 1852: 160. 1853. Mora;
Zarzamora.

Damp or wet, mountain thickets, 1,700-2,700 meters, some-
times in open forest, frequently in oak forest; Jalapa; Guatemala;
Sacatepe"quez ; Chimaltenango; Solola; Huehuetenango; Quezal-
nango; San Marcos. Costa Rica.

An erect or arching shrub about 1.5 meters high, the stems subterete, densely
sordid-pubescent with short gland-tipped hairs, sparsely aculeate with compressed
recurved prickles; leaves pedately 3-5-foliolate, the petioles shortly glandular-
setose and aculeate; leaflets oval to elliptic-oblong, mostly 6-11 cm. long, acumi-
nate, obtuse or rounded at the base, acutely duplicate-serrate, deep green and
puberulent above, densely sordid-pilose beneath, the costa often aculeate; panicles
few- many-flowered, terminal and lateral, mostly unarmed; sepals ovate, acute,
reflexed in fruit; petals obovate, white or pinkish, longer than the sepals; fruit
1-1.5 cm. long, 8-9 mm. thick, the small drupelets numerous, pubescent at the
apex.

The name "mora" is applied commonly in Central America to
blackberry fruit. The proper name for the plant should be "zarza-
mora" but that term is used but little in Guatemala.

Rubus leptosepalus Bonn. Smith, Bot. Gaz. 57: 421. 1914.
Mora.

Known only from the vicinity of Coban, Alta Verapaz, about
1,300-1,400 meters, where collected several times, growing in wet
thickets or brushy pastures; type Tuerckheim 2452.

An arching shrub about 1.5 meters high, the stems subangulate, densely pilose
and more or less glandular-setose, armed with numerous recurved prickles; leaves
pedately 3-5-foliolate, the petioles glandular-pilose and aculeate; leaflets oval to
oblong-elliptic, mostly 7-15 cm. long, caudate-acuminate, obtuse or rounded at
the base, sharply duplicate-serrate and often almost laciniate, green and dull
above, glabrate, densely and softly pilose beneath, often aculeate on the costa;
panicles usually large and many-flowered, sparsely aculeate, glandular-setulose,
the bracts often large and foliaceous; sepals linear-lanceolate, setaceous-append-
aged, 1.5-2 cm. long or often considerably larger, conspicuously nerved; petals
pink, 1.5 cm. long or less; drupelets numerous, glabrous.

This shrub has the appearance of being a teratological form,
perhaps of R. adenotrichos, the greatly elongate and somewhat
foliaceous sepals having an abnormal appearance. It is, however,
rather frequent in pastures about Coban. It certainly is easy of
recognition among the several Rubus species of the region and
apparently is a normal specific unit.

480 FIELDIANA: BOTANY, VOLUME 24

Rubus macrogongylus Focke, Repert. Sp. Nov. 9: 236. 1911.
Mora.

Moist mountain thickets, 1,300-2,000 meters; Chiquimula;
Jalapa; Quezaltenango. Central and southern Mexico.

An arching shrub 1.5-2.5 meters tall, the stems closely grayish-tomentose,
armed with recurved prickles; leaves pedately 3-5-foliolate, rather thin, the slender
petioles sparsely recurved-aculeolate; leaflets oblong or ovate-oblong, mostly 6-11
cm. long, long-acuminate, obtuse or rounded at the base, regularly and closely
serrulate, puberulent above, somewhat paler beneath and appressed-tomentose,
subsericeous on the nerves, the lateral nerves about 10 pairs; inflorescence terminal,
small and few-flowered, unarmed, sordid-tomentose and sparsely stipitate-glandu-
lar; sepals ovate, mucronate, grayish-tomentose outside, white-tomentose within,
reflexed in fruit; petals slightly longer than the sepals; fruit black at maturity,
oblong or cylindric-oblong, the drupelets numerous, glabrous.

Rubus miser Liebm. Vid. Medd. 1852: 156. 1853. Mora;
Zarzamora; Cakitocan, Tocan (Coban, Quecchi).

Pine-oak forest or moist or dry thickets, often in brushy fields,
1,100-2,000 meters; Alta Verapaz; Baja Verapaz; Jalapa; Guate-
mala; Chimaltenango; Quiche"; Huehuetenango; Totonicapan. Hon-
duras; Costa Rica.

An arching shrub or a small vine, the stems subterete, densely fulvous-
tomentulose and bearing numerous short gland-tipped setae, armed with short
compressed recurved prickles; leaves pedately 3-5-foliolate, thick and firm, the
petioles recurved-aculeate; leaflets ovate or ovate-oblong, mostly 6-12 cm. long,
acuminate, obtuse or rounded at the base, sharply serrate, dull and puberulent
above, densely and softly sordid-pilose beneath; flowers racemose, the racemes
terminal, few-flowered; sepals ovate-lanceolate, acuminate, densely grayish-
tomentose and glandular-setulose, reflexed in fruit; petals shorter than the sepals;
fruit almost black at maturity, very sour, the drupelets small, numerous, glabrous.

Rubus Pringlei Rydb. N. Amer. Fl. 22: 443. 1913. R. occi-
dentalis var. grandiflorus Focke in Donn. Smith, Bot. Gaz. 16: 3.
1891 (type from Volcan de Agua, 2,550 meters, J. D. Smith 2168).
R. occidentalis var. mexicanus Focke, Bibl. Bot. 17, pt. 72: 210. 1911.

Moist mountain thickets, 2,500-3,000 meters; Sacatepe"quez
(Volcan de Fuego); Solola (Volcan de Atitlan). Mexico.

Stems subterete, 1-2 meters tall, glabrous, armed with small compressed
recurved prickles; leaves all 3-foliolate, the petioles aculeate, glabrous; leaflets
lanceolate to lance-ovate, mostly 5-10 cm. long, narrowly long-acuminate, sub-
acute to rounded at the base, acutely duplicate-serrate, dark green and sparsely
puberulent above or almost glabrous, closely white-tomentose beneath; flowers
solitary or in clusters of 2-3, the pedicels tomentose, setose or weakly aculeate,
recurved in fruit; sepals ovate, caudate-acuminate, 6-7 mm. long, tomentose on
both sides, suberect and enclosing the fruit; petals elliptic, about equaling the

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 481

sepals; fruit ovoid, 2 cm. long, 1.5 cm. thick, red or at last deep purple, with a
bloom, the drupelets numerous, tomentose.

This is probably the plant reported from Volcan de Agua by
Hemsley as R. occidentalis L., a species of the United States.

Rubus rosaefolius J. E. Smith, PL Icon. ined. pi. 60. 1791.

Native of southern and eastern Asia, often cultivated for orna-
ment; sometimes planted in Guatemala, and perhaps at least
partially naturalized in the mountains of San Marcos.

A shrub 1-1.5 meters tall, the stems erect or recurved, pilose or glabrate,
aculeate; leaves pinnately 5-15-foliolate, the petiole and rachis pilose and aculeate;
leaflets lanceolate or lance-oblong, 4-8 cm. long, acuminate, obtuse or rounded
at the base, incised-serrate or duplicate-serrate, sparsely pilose or glabrate, the
lateral nerves 10-15 pairs, slender but prominent; flowers solitary or in small
cymes, usually double; sepals lanceolate, caudate-acuminate, often with foliaceous
tips; petals white, 1-2 cm. long; fruit thimble-shaped, 2-3.5 cm. long, bright red
or orange, the drupelets very numerous, small, glabrous.

The double-flowered form found in Guatemala is var. coronarius
Sims.

Rubus sapidus Schlecht. Linnaea 13: 269. 1839. R. sapidus
var. grandifolius Focke in Donn. Smith, Enum. PI. Guat. 2: 19.
1891, nomen. R. amplior Rydb. N. Amer. Fl. 22: 456. 1913 (type
from Santa Rosa, Baja Verapaz, Tuerckheim 1424). R. Tuerck-
heimii Rydb. op. cit. 457. 1913 (type from Coban, Alta Verapaz,
Tuerckheim 8387). Mora; Sakitocan (Coban, Quecchi).

Moist or rather dry thickets or open forest, often in open fields,
1,100-2,800 meters; Alta Verapaz; Baja Verapaz; El Progreso;
Zacapa; Jalapa; Guatemala; Sacatepe'quez; Solola; Huehuetenango;
Quezaltenango ; San Marcos. Southern Mexico.

Stems mostly 1.5-2.5 meters tall, erect or arching, sometimes subscandent,
subterete, sparsely pilose or in age glabrate, often purplish, armed with stout
retrorse prickles; leaves pedately 3-5-foliolate, the petioles usually rather densely
retrorse-aculeate; leaflets mostly subcoriaceous, broadly ovate to ovate-oblong,
6-10 cm. long, acuminate, obtuse to subcordate at the base, closely and acutely
serrate, dull and sparsely pilose above but soon glabrate, densely soft-pilose
beneath; inflorescence corymbiform or broadly paniculate, usually many-flowered,
densely armed with recurved prickles; sepals about 6 mm. long, mucronate,
whitish-tomentose on both sides; petals white, 1 cm. long; fruit subglobose,
black at maturity, the drupelets numerous, pubescent at the apex.

Rubus Smithii Rydb. N. Amer. Fl. 22: 453. 1913. R. polio-
phyllus Focke in Donn. Smith, Bot. Gaz. 18: 202. 1893, not Kuntze,
1891.

482 FIELDIANA: BOTANY, VOLUME 24

Sacatepe"quez, at about 1,800 meters; type from San Rafael,
J. D. Smith 2141; also on Volcan de Fuego. Southern Mexico.

Stems subterete, puberulent-tomentose, armed with compressed recurved
prickles; leaves pedately 3-5-foliolate, the petioles aculeate and tomentulose;
leaflets subcoriaceous, oval to elliptic, mostly 5-10 cm. long, abruptly acuminate,
obtuse or rounded at the base, acutely serrate, puberulent above with branched
hairs, fulvous-tomentose beneath, the lateral nerves 8-10 pairs; panicles terminal
and axillary, tomentulose, with occasional glands or bristles; sepals ovate, short-
acuminate, tomentose on both sides; petals scarcely longer than the sepals, white;
fruit hemispheric, black, the drupelets about 20, glabrous.

Rubus trilobus Seringe in DC. Prodr. 2: 566. 1825. R. trilobus
var. guatemalensis Focke in Bonn. Smith, Bot. Gaz. 18: 201. 1893
(type from Volcan de Agua, Sacatepe"quez, W. C. Shannon 3631).
Oreobatus trilobus Rydb. N. Amer. Fl. 22: 428. 1913. Morita (fide
Aguilar).

Moist or wet, mixed or coniferous, mountain forest, frequently
in forest of oak, Cupressus, or Abies, sometimes on white-sand
slopes, 2,000-4,200 meters; Sacatepe"quez; Chimaltenango; Solola;
Quich^ ; Huehuetenango ; Totonicapan; Quezaltenango ; San Marcos.
Southern Mexico.

Plants slender, suberect, unarmed, the stems often straggling and supported
upon other shrubbery, sometimes 5 meters high or even more, sparsely branched,
the bark brown or purplish, deciduous, the branchlets puberulent or pilose; leaves
long-petiolate, triangular-cordate, thin, somewhat 3-lobate, deep green above,
paler beneath, pilose on both surfaces, finely serrate; flowers mostly solitary;
sepals ovate, caudate-acuminate, 1.5 cm. long, pilose outside, tomentose within,
usually appressed to the fruit and enclosing it; petals white, 2 cm. long; fruit
hemispheric, black-purple, 1.5 cm. broad, the drupelets large and distinct.

The shrub is a typical one of the high forests, occurring mostly at
2,700 meters or more. It seldom is plentiful in any locality, occurring
as isolated individuals, although in the Sierra de los Cuchumatanes
it is abundant. The fruit is sweet but with a distinctly acidulous
flavor.

Rubus urticaefolius Poir. in Lam. Encycl. 6: 246. 1804.
R. trichomallus Schlecht. Linnaea 13: 268. 1839. (?)#. adenotrichus
subsp. leptaleos Focke, Bibl. Bot. 18, Heft 83: 70. 1914 (type from
Dept. Santa Rosa, Heyde &Lux 4473, 4474). Mora; Tocan, Cakito-
cdn (Coban, Quecchi).

Moist or wet thickets or forest, sometimes in brushy fields,
600-1,500 meters; Alta Verapaz; Chiquimula; Santa Rosa; Guate-
mala; Suchitepe"quez; Quezaltenango; San Marcos. Southern

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 483

Mexico; Honduras; Costa Rica; Panama; southward to Peru and
Brazil.

Usually a shrub of 1.5-2.5 meters with arching stems, these obtusely angulate,
densely pubescent and densely covered with long spreading setae, sparsely re-
curved-aculeate; leaves pedately 3-5-foliolate, the petioles densely pubescent and
setose, densely aculeate; leaflets ovate to lance-oblong, mostly 7-15 cm. long,
short-acuminate, obtuse to subcordate at the base, acutely and finely duplicate-
serrate, dark green and densely pubescent above, grayish-tomentose beneath or
whitish; panicles terminal or axillary, often large and pyramidal, many-flowered,
the branches densely reddish-setose; sepals lanceolate, about 5 mm. long, subulate-
acuminate, spreading in age, tomentose and setulose; petals mostly white, scarcely
longer than the sepals; fruit rather small, black or dark purple at maturity, sour,
the drupelets glabrous.

This is one of the commonest of Guatemalan blackberries, easy
of recognition because of the very abundant, glandless bristles
covering the stems and other parts.

SPIRAEA L.

Deciduous shrubs; leaves alternate, simple, dentate or serrate, sometimes
lobate, usually short-petiolate, without stipules, commonly penninerved; flowers
normally perfect, in umbelliform racemes, corymbs, or panicles; hypanthium
campanulate or cup-shaped, the sepals 5, small; petals 5, commonly rounded and
longer than the sepals; stamens 15-60, inserted between the disk and the sepals;
pistils generally 5, distinct; fruit of follicles, these dehiscent along the inner suture,
containing several minute oblong seeds.

Species 80 or more, in temperate regions of both hemispheres,
many of the species well known in cultivation because of their
handsome flowers. In America the genus reaches its southern limit
in southern Mexico, where one species is native.

Spiraea cantoniensis Lour. var. lanceata Zabel, Gartenzeit.
41. 1893. Buquet de novia.

Native of China and Japan but widely cultivated, and frequent
in Central America; planted for ornament in gardens of the Guate-
malan mountains and more or less naturalized about Coban and in
San Marcos, as well as probably elsewhere.

A glabrous shrub 1-1.5 meters high with slender branches; leaves rhombic-
oblong or rhombic-lanceolate, obtuse or subacute, cuneate at the base, incised-
serrate, deep green above, pale bluish green beneath, short-petiolate; flowers
white, double, about 1 cm. broad, in small, rather dense umbels.

The leaves remain on the shrub all or most of the year and flowers
may be found at almost any season. The blossoms are not abundant,
and the shrub is not an attractive one for the genus but it is much

484 FIELDIANA: BOTANY, VOLUME 24

planted in the Guatemalan mountains, and sometimes even at
low elevations.

CONNARACEAE

Reference: Gustav Schellenberg, Connaraceae, Pflanzenreich IV.
127. 1938.

Shrubs or small trees, often woody vines; leaves alternate, odd-pinnate or
1-foliolate, without stipules; flowers small, perfect, regular, in terminal or lateral
panicles, these lax or dense, often arising from defoliate nodes; sepals 5, imbricate
or subvalvate, free or rarely connate; petals 5, free or coherent above the base;
stamens 10, the inner 5 epipetalous, the outer 5 episepalous or sometimes reduced
to staminodia, the filaments united at the base; carpels of the ovary 5, free, some-
times only 1; ovules 2 in each cell, erect, collateral, anatropous; several or only 1
of the carpels fertile, follicular, irregularly dehiscent, or indehiscent; seed 1 in
each follicle, rarely 2, subtended by a basal aril; endosperm abundant or none.

Genera about 24, widely dispersed in the tropics of both hemi-
spheres. Only the following are known in North America.

Calyx lobes valvate; capsule densely tomentose at maturity, sessile; leaflets

densely pubescent beneath Cnestidium.

Calyx lobes imbricate; capsule glabrous or glabrate at maturity; leaflets glabrous
or glabrate beneath.

Capsule sessile; calyx accrescent after an thesis Rourea.

Capsule stipitate; calyx not accrescent Connarus.

CNESTIDIUM Planchon

Woody vines; leaves odd-pinnate; inflorescences paniculate, pseudoterminal,
the pedicels almost obsolete; sepals 5, very narrowly imbricate or valvate, in fruit
erect, not accrescent, tomentose on both surfaces; petals 5, only slightly longer
than the sepals, glabrous; filaments glabrous, the anthers dorsifixed, dehiscent by
longitudinal introrse slits, the connective broad; carpels of the ovary 5, free,
hispidulous, the styles free, glabrous, the stigmas capitate; ovules 2 in each carpel,
erect; usually a single follicle maturing, velutinous-pilose outside, glabrous within;
seed 1, the testa coriaceous, black, lustrous, subtended at the base by a cupular
aril; endosperm rudimentary.

One other species is known, in the Guianas.

Cnestidium rufescens Planch. Linnaea 23: 440. 1850. Rourea
hondurensis Donn. Smith, Bot. Gaz. 40: 2. 1905 (type from Tela,
Honduras). Bejuco Colorado (Pete"n); Uayaumac (Pete"n, Maya,
fide Lundell).

Moist or wet thickets or forest, sometimes on limestone, often
in pine forest, 600 meters or less; Pete"n; Alta Verapaz; Iza-
bal; Retalhuleu. Tabasco; British Honduras to Panama; Cuba;
Colombia.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 485

A small or often large vine, climbing to a height of 12 meters, the stems as
much as 7 cm. in diameter, the young branchlets densely rufous-pubescent; leaves
large, 7-9-foliolate; leaflets oblong or obovate-oblong, mostly 3-8 cm. long and
1.5-4 cm. wide, abruptly short-acuminate, rounded or obtuse at the base, coria-
ceous, entire, the margins often revolute, glabrous and lustrous above, densely
rufous-tomentose beneath; inflorescences mostly axillary and forming a pseudo-
terminal panicle, the branches densely rufous-tomentose; sepals 3 mm. long,
oblong, tomentose; petals white, 4 mm. long; fruit about 1.5 cm. long, obtuse,
somewhat arcuate, densely rufous-tomentose or reddish; seed 12 mm. long.

CONNARUS L.

Usually woody vines with scant pubescence, often glabrous; leaves odd-pin-
nate, 3-foliolate, or rarely 1-foliolate, the leaflets entire, opposite or subalternate;
flowers perfect, generally white, the inflorescences mostly terminal and paniculate;
sepals 5, broadly or narrowly imbricate, more or less punctate; petals 5, generally
longer than the sepals, glabrous or pubescent, sometimes glandular; filaments
more or less connate below into a tube, pilosulous; anthers oblong, introrsely
dehiscent, the cells mostly glandular at the base, the connective glandular at the
apex; carpel of the ovary 1, ovoid, tomentose outside, glabrous or pilose within;
style villous at the base, usually glandular above; stigma oblique-reniform, the
margin lobulate; ovules 2, collateral, erect; fruit follicular, dehiscent by the ventral
suture, or sometimes also by the dorsal suture, oblique-pyriform or somewhat
cylindric, fusiform, or clavate, often mucronate or rostrate, narrowed at the base
into a long or short stipe, the pericarp ligneous or coriaceous; seed 1, the testa
usually dark purple or almost black, lustrous, arillate at the base; endosperm
none; cotyledons thick.

Species about 120, in the tropics of both hemispheres. Two or
three additional ones are known from southern Central America.

Leaflets 5 C. lentiginosus.

Leaflets 3 C. Lambertii.

Connarus Lambertii (DC.) Sagot, Ann. Sci. Nat. IV. 13: 295.
1882. Omphalobium Lambertii DC. Me*m. Soc. Hist. Nat. Paris 2:
389. 1825. C. Pottsii Wats. Proc. Amer. Acad. 21: 463. 1886 (type
from shores of Lago de Izabal, S. Watson). C. brachybotryosus Donn.
Smith, Bot. Gaz. 57: 417. 1914 (type from Cubilgiiitz, Alta Verapaz,
Tuerckheim 4027). C. lonchotus Blake, Contr. Gray Herb. 52: 69.
1917 (type from Moho River, British Honduras, M. E. Peck 727).

Moist or wet forest or thickets, 350 meters or less; Pete"n; Alta
Verapaz; Izabal. British Honduras; Honduras; West Indies; north-
ern South America.

A rather large, woody vine, or often an erect shrub or small tree, sometimes
9 meters high, the trunk as much as 8 cm. in diameter, the branchlets glabrous or
nearly so; leaves 3-foliolate, glabrous; leaflets elliptic to obovate-elliptic or oblong-

486 FIELDIANA: BOTANY, VOLUME 24

elliptic, 6-15 cm. long or somewhat longer, 8 cm. wide or less, coriaceous, acumi-
nate, rounded or obtuse at the base, lustrous, glabrous or when young somewhat
pubescent beneath, especially on the costa, the lateral nerves usually 5-8; inflores-
cences axillary or terminal, paniculate or composed of fascicled racemes or panicles,
much shorter than the leaves, rufous-tomentose; sepals 3 mm. long, acute, tomen-
tose outside, glabrous within, densely punctate; petals white, 4.5 mm. long,
subacute, glabrous, densely punctate; follicle obliquely obovoid, about 2 cm.
long and 12-14 mm. broad, turgid or somewhat compressed, obliquely apiculate,
short-stipitate, glabrous or glabrate; seed black, lustrous, subtended by a small
aril.

Schellenberg recognized as distinct species all four here treated
as one. He gives a key for their separation, but apparently had no
clear idea of the characters by which they were to be separated,
for the characters he uses are apparently of little significance, if
they exist at all. C. Pottsii, of which we have seen no authentic
material, he placed in a different subgenus from the other three, but
the characters on which he bases subgenera and sections are quite
as confused as those used for separating species. There is now avail-
able from Guatemala and British Honduras a substantial amount
of material of this group and one would expect that all the local
species would be represented by one or more specimens. All the
material actually seems to represent a single species that exhibits
little variation.

Connarus lentiginosus Brandegee, Univ. Calif. Publ. Bot. 6:
186. 1915. Trompillo.

Moist or rather dry thickets or mixed forest of the Pacific low-
lands, 2,500 meters or less; Solola; Suchitepe"quez; Retalhuleu; San
Marcos. Chiapas, the type from Huitla.

A small or large, woody vine, the branches glabrous; leaves large, long-petio-
late, usually 5-foliolate; leaflets elliptic-oblong or elliptic, 6-16 cm. long, 3-7 cm.
wide, rather abruptly short-acuminate, obtuse or cuneate at the base, subcoria-
ceous, glabrous, conspicuously punctate beneath, somewhat lustrous, the lateral
nerves about 8 pairs; inflorescences paniculate, terminal or axillary, about 15 cm.
long, usually much branched, densely rufous-tomentulose, the flowers very
numerous, pale yellowish; sepals 2.5 mm. long, ovate-lanceolate, tomentulose;
petals 3.5 mm. long, oblong, acute, punctate; filaments glabrous; follicle about
2 cm. long, oblique-ellipsoid, short-stipitate, glabrous or glabrate, apiculate
laterally near the rounded apex.

ROUREA Aublet

Usually woody vines, sometimes more or less erect shrubs or trees; leaves
odd-pinnate, rarely 1-foliolate; flowers small, 5-parted, paniculate, terminal,
pseudoterminal, or axillary; sepals imbricate, sometimes glandular; petals usually

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 487

longer than the sepals, glabrous; filaments glabrous, connate at the base into a tube,
the anthers dorsifixed, dehiscent by longitudinal introrse slits; carpels of the
ovary 5, free, pubescent outside, glabrous within; styles free, glabrous, the stigmas
capitate; ovules 2 in each carpel, collateral, erect; only 1 follicle maturing, sub-
tended at the base by the somewhat accrescent sepals, obovoid, rounded at the
apex, mucronulate, longitudinally striate, dehiscent by a ventral suture; seed 1,
lustrous, black, subtended at the base by a cupular aril; endosperm none; cotyle-
dons thick, the radicle superior.

About 30 species, in tropical America. Three other species occur
in southern Central America.

Sepals glabrous R. glabra.

Sepals densely appressed-pilosulous R. Schippii.

Rourea glabra HBK. Nov. Gen. & Sp. 7: 41. 1825. Canjuro;
Uayumac (Pete"n, Maya, fide Lundell).

Moist or dry thickets or mixed lowland forest, often in second
growth, 300 meters or le§s; Pete"n; Izabal; Santa Rosa; Escuintla;
Retalhuleu; probably in all Pacific coast departments. Mexico;
British Honduras to Salvador and Panama; West Indies; South
America.

A small or large, woody vine, glabrous throughout or nearly so, the branches
terete; leaves rather large, mostly 5-foliolate, sometimes 3-foliolate, the petiolules
3-3.5 mm. long; leaflets oblong-elliptic, mostly 3-10 cm. long and 1-3.5 cm. wide,
acuminate or long-acuminate with an obtuse tip, rounded or obtuse at the base,
chartaceous, with 6-7 pairs of lateral nerves; inflorescences axillary or forming
terminal panicles, usually 5 cm. long or less but sometimes larger, lax, many-
flowered, the pedicels as much as 5 mm. long, slender, articulate near the base;
sepals 2 mm. long, triangular-elliptic, ciliate, thickened in age and 5 mm. long or
more; petals white or yellowish white, 6 mm. long, glabrous; follicle 14-16 mm.
long, 5-7 mm. thick, oblong, subterete, slightly curved; seed 10-12 mm. long,
6 mm. thick, black and shining, arillate at the base.

Called "tietie" in British Honduras, doubtless because the pliable
stems are employed as a substitute for cordage; "chilillo," "mata-
perros" (Yucatan). The roots are employed in Mexico for imparting
a bright red dye to skins. The plant is well known in Central
America and Mexico because of the poisonous properties of its seeds,
which have been employed at times, it is said, for criminal poisoning
of people, and commonly for killing rats and other noxious animals.
The seeds or fruits are eaten commonly by some birds, particularly
those called chachas, and it is stated that if dogs eat birds that have
eaten the fruit, they die. Further, if people eat the flesh of such
birds, they are poisoned and die. (For a detailed account of the
poisonous properties of Rourea glabra, see Standley & Calderon,
Lista Prel. PI. El Salvador 91-94. 1925.)

488 FIELDIANA: BOTANY, VOLUME 24

Rourea Schippii Standl. Carnegie Inst. Wash. Publ. 461: 58.
1935.

Known only from the type, in forest, Rio Grande, British Hon-
duras, 75 meters, W. A. Schipp 1168.

A woody vine 12 meters long, the trunk 5 cm. in diameter, the branchlets
minutely strigillose or almost glabrous; leaflets 7, on petiolules 3-4 mm. long,
ovate or oblong-elliptic, 7-11 cm. long, 3.5-5.5 cm. wide, shortly obtuse-acuminate,
rounded at the base, subchartaceous, glabrous, densely puncticulate above, the
lateral nerves about 5 pairs; panicles axillary, branched, half as long as the leaves,
the branches pilose with short ascending hairs, the pedicels 11 mm. long or less;
sepals 2 mm. long, orbicular, rounded or apiculate at the apex, ciliate and ap-
pressed-pilosulous; petals white, spatulate-obovate, 6-7 mm. long, glabrous,
broadly rounded or truncate at the apex.

KRAMERIACEAE

Reference: N. L. Britton, Krameriaceae, N. Amer. Fl. 23: 195-
200. 1930.

Shrubs or perennial herbs, usually pubescent, the pubescence commonly
sericeous; leaves alternate, simple and entire, rarely 3-foliolate; stipules none;
flowers rather large, irregular, axillary or in terminal racemes, the peduncles
usually bearing 2 opposite foliaceous bracts; sepals 4-5, unequal; petals 5, the
3 upper ones long-unguiculate, distinct or partly united, the 2 others commonly
much smaller, broad, thick, and sessile; stamens 4 in North American species,
free or borne on the united claws of the upper petals; anthers 2-celled, the cells
dehiscent by a pore; ovary 1-celled, the ovules 2, collateral, pendulous, anatropous;
style cylindric, acute; fruit globose, indehiscent, 1-seeded, covered with sharp
slender spines; cotyledons thick; endosperm none.

The family consists of only the following genus. By many
authors it has been united with the Leguminosae, but Bentham
and Hooker placed it in the Polygalaceae. The dried roots of some
of the South American species are known in commerce as rhatany
roots, and are official in the U. S. Pharmacopoeia. They are, or
were, used as a tonic and powerful astringent in treating chronic
diarrhea. Roots of the Mexican species have been exported for
the same purpose. The plants yield a yellow or brownish red dye.
The roots of some species have been employed in Europe for making
ink, coloring wine, and in manufacture of dentifrices.

KRAMERIA L.

Perhaps 25 species, ranging from southwestern United States to
Chile. Only the following are known from Central America, but
there are numerous species in Mexico.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 489

Leaves conspicuously petiolate, oblong or lance-oblong K. cuspidata.

Leaves sessile, linear or nearly so K. rewluta.

Krameria cuspidata Presl, Rel. Haenk. 2: 103. 1835.

Dry rocky open slopes, 1,200-1,500 meters; Chiquimula (south-
east of Quezaltepeque, Steyermark 31343). Western Mexico; Hon-
duras; Costa Rica.

An erect or ascending shrub, a meter high or less, sometimes forming loose,
much-branched bushes a meter broad, the branches densely tomentose with white
or grayish hairs; petioles 4-6 mm. long; leaf blades oblong or lance-oblong, 2 cm.
long and 7 mm. wide or smaller, cuspidate with a spinulose tip, acute at the base,
densely tomentose; peduncles mostly shorter than the subtending leaves, bracteate
near the middle; sepals oblong, obtuse, 6-7 mm. long, densely strigose; upper
petals united to near the middle, brown-red; fruit globose, hard, densely white-
villous, 5-6 mm. in diameter, covered with acicular spines 3-4 mm. long, these
retrorse-barbate near the apex.

Krameria revoluta Berg, Bot. Zeit. Regensb. 14: 751. 1856.
K. dichrosepala Bonn. Smith, Bot. Gaz. 49: 453. 1910 (type from
Gualan, Zacapa, C. C. Deam 6273).

Dry rocky open hillsides, sometimes in pine-oak forest, 200-1,000
meters; Zacapa; Jalapa; Jutiapa. Southern Mexico.

A low shrub, usually 40 cm. high or less, erect or prostrate, often densely
branched, the branches densely white-strigose; leaves sessile, linear or nearly so,
8-12 mm. long, acute and spinulose-tipped, densely strigose; peduncles shorter
or longer than the leaves, the 2 bracts similar to the leaves; sepals linear to oblong-
lanceolate, 7-9 mm. long; upper petals united to much above the middle, brown-
red; fruit densely white-sericeous, 5-6 mm. in diameter, densely covered with
slender brown spines 3-4 mm. long, these sparsely barbate above.

INDEX

Abuta, 259
Acaena, 433
Achatocarpus, 193
Achyranthes, 144
Agdestis, 193 •
Agonandra, 87
Agrimonia, 435
Aizoaceae, 203
Alchemilla, 436
Allionia, 175
Alternanthera, 146
Amaranthaceae, 143
Amaranthus, 152
Amelanchier, 440
Ampelocera, 2
Anaxagorea, 271
Anemone, 244
Annona, 272
Annonaceae, 270
Anredera, 215
Antidaphne, 63
Antigonon, 105
Apodanthes, 102
Arceuthobium, 63
Arenaria, 218
Argemone, 348
Aristolochia, 93
Aristolochiaceae, 93
Artocarpus, 11

Balanophoraceae, 92
Basellaceae, 214
Beilschmiedea, 307
Berberidaceae, 256
Beta, 138
Bocconia, 349
Boerhaavia, 176
Boldoa, 178
Bougainvillea, 179
Boussingaultia, 216
Brasenia, 239
Brass! ca, 356
Brosimum, 13
Brunellia, 424
Brunelliaceae, 423
Bryophyllum, 404

Cabomba, 240
Cakile, 362
Calandrinia, 208
Cananga, 280
Capparidaceae, 380

Capparis, 381
Capsella, 363
Cardamine, 363
Caryophyllaceae, 217
Cassytha, 308
Castilla, 17
Cecropia, 20
Celosia, 157
Celtis, 3
Cerastium, 223
Ceratophyllaceae, 242
Ceratophyllum, 243
Chaetpptelea, 5
Chamissoa, 159
Chenopodiaceae, 137
Chenopodium, 139
Chlorophora, 23
Chrysobalanus, 441
Cinnamomum, 309
Cissampelos, 259
Clarisia, 25
Clematis, 245
Cleome, 388
Cnestidium, 484
Coccoloba, 108
Commicarpus, 181
Compsoneura, 295
Connaraceae, 484
Connarus, 485
Couepia, 442
Coussapoa, 25
Crassulaceae, 404
Crataegus, 444
Crataeva, 391
Cruciferae, 354
Cunoniaceae, 424
Cyathula, 160
Cydonia, 445
Cymbopetalum, 281

Delphinium, 248
Dendrophthora, 64
Descurainia, 367
Desmopsis, 283
Dianthus, 226
Disciphania, 262
Distylium, 426
Dorstenia, 27
Draba, 368
Drimys, 269
Drosera, 400
Droseraceae, 399

491

492

FIELDIANA: BOTANY, VOLUME 24

Drymaria, 227
Duchesnea, 446

Echeveria, 406
Eriobotrya, 446
Erysimum, 369
Eschscholtzia, 352

Ficus, 30

Forchhammeria, 393
Fragaria, 447
Froelichia, 162

Glinus, 203
Gomphrena, 163
Grevillea, 59
Guamatela, 449
Guatteria, 286
Gymnopodium, 119
Gynandropsis, 394
Gypsophila, 232
Gyrocarpus, 345

Hamamelidaceae, 426
Heisteria, 88
Helosis, 92
Hernandia, 346
Hernandiaceae, 344
Hirtella, 450
Holodiscus, 452
Hydrangea, 417
Hyperbaena, 263

Iresine, 166

Krameria, 488
Krameriaceae, 488

Lamprophragma, 370
Lauraceae, 302
Ledenbergia, 194
Lepidium, 370
Licania, 454
Licaria, 311
Liquidambar, 427
Litsea, 314
Lobularia, 373
Loranthaceae, 62
Lozanella, 6

Magnolia, 266
Magnoliaceae, 266
Mahonia, 256
Malmea, 287
Malus, 457
Marathrum, 401
Matthiola, 374
Matudaea, 429
Menispermaceae, 258
Mesembryanthemum, 204
Microtea, 195
Mirabilis, 181

Mitrastemon, 102
Mollinedia, 299
Mollugo, 205
Monimiaceae, 299
Montia, 209
Moraceae, 10
Moringa, 398
Moringaceae, 398
Morisonia, 395
Morus, 48
Muehlenbeckia, 120
Myristica, 295
Myristicaceae, 294

Nasturtium, 374
Nectandra, 317
Neea, 185

Neomillspaughia, 122
Nigelia, 249
Nyctaginaceae, 174
Nymphaea, 240
Nymphaeaceae, 239

Ocotea, 322
Olacaceae, 88
Opiliaceae, 86
Oreobroma, 210
Oryctanthus, 65

Papaver, 353
Papaveraceae, 347
Persea, 330
Petiveria, 195
Pfaffia, 172
Philadelphus, 418
Philoxerus, 173
Phoebe, 336
Phoradendron, 66
Photinia, 458
Phthirusa, 76
Phyllonoma, 420
Phyllostylon, 7
Phytolacca, 196
Phytolaccaceae, 192
Pilostyles, 103
Pisonia, 188
Platanaceae, 430
Platanus, 431
Pleuropetalum, 174
Podopterus, 123
Podostemonaceae, 401
Polanisia, 395
Polygonaceae, 104
Polygonum, 124
Portulaca, 211
Portulacaceae, 207
Potentilla, 459
Poterium, 461
Poulsenia, 51
Pourouma, 52
Proteaceae, 58
Prunus, 461

INDEX 493

Pseudolmedia, 53 Sparattanthelium, 346

Psittacanthus, 78 Spergula, 235

Pyrus, 470 Spiraea, 483

Stegnosperma, 201

Rafflesiaceae, 101 Stellaria, 236

Ranunculaceae, 243 Steriphoma, 396

Ranunculus, 249 Struthanthus, 80
Raphanus, 376

Reseda, 397 Talauma, 268

Resedaceae, 397 Talinum, 213

Rheum, 130 Thalictrum, 253

Ribes, 421 Tillaea, 414

Rivina, 200 Torrubia, 191

Rollinia, 288 Tovaria, 380

Romanschulzia, 377 Tovariaceae, 380

Rorippa, 379 Trema, 8

Rosa, 471 Trianthema, 206

Rosaceae, 432 Trichostigma, 202

Roupala, 60 Triplaris, 136

Rourea, 486 Tristicha, 403

Rubus, 472 Trophis, 56
Rumex, 131

Ruprechtia, 134 Ulmaceae, 1

Unonopsis, 292
Sagina, 233

Sapranthus, 290 Villadia, 415

Saxifragaceae, 416 Virola, 296
Schoepfia, 89

Sedum, 410 Weinmannia, 425

Sesuvium, 205 Winteraceae, 269
Silene, 234

Siparuna, 300 Ximenia, 90

Sorocea, 55 Xylopia, 293

Publication 577