LI B RAR.Y

OF THE

UNIVERSITY
OF ILLINOIS

v. 24-
pt.7

BIOLOGY

v. 24

no- *7
t. 3

FLORA OF GUATEMALA

ROGERS McVAUGH

FIELDIANA: BOTANY
VOLUME 24, PART VII, NUMBER 3

Published by

CHICAGO NATURAL HISTORY MUSEUM
OCTOBER 25, 1963

3IOLOGV LiBRAnV
101 BURRIIL HALL

FLORA OF GUATEMALA

PART VII

ROGERS McVAUGH

Curator of Vascular Plants, University Herbarium
The University of Michigan

FIELDIANA: BOTANY

VOLUME 24, PART VII, NUMBER 3

Published by

CHICAGO NATURAL HISTORY MUSEUM
OCTOBER 25, 1963

Assisted by National Science Foundation grant No. 19071

Library of Congress Catalog Card Number: 63-21 765

PRINTED IN THE UNITED STATES OF AMERICA
BY CHICAGO NATURAL HISTORY MUSEUM PRESS

1

3

CONTENTS

Family MYRTACEAE
Genera Included in Part VII, Number 3

PAGE PAGE

Eucalyptus 285 Myrcianthes 377

Calyptranthes 290 Myrciaria 379

Eugenia 309 Pimenta 382

Myrcia 374 Psidium 385

Ugni.. ... 403

LIST OF ILLUSTRATIONS

TEXT FIGURES

PAGE

49. Flowers and inflorescence in Calyptranthes and Myrcia 295

50. Inflorescences in Calyptranthes and Eugenia 329

51. Eugenia Oerstedeana 355

52. Dichasium and embryo in Myrcianthes 378

53. Myrciaria floribunda 380

54. Floral morphology in Psidium 388

55. Ugni montana 404

Flora of Guatemala

MYRTAGEAE. Myrtle Family

Reference: Otto Berg, Revisio Myrtacearum Americae, Linnaea
27: 1-472. 1855-1856.

Shrubs or trees or rarely subherbaceous. Leaves simple, opposite (except
in some introduced genera), exstipulate, entire or rarely crenate, punctate with
resinous or pellucid glands, usually pinnately veined. Midvein usually elevated
and prominent on the lower surface. Principal lateral veins usually uniting
distally into a "marginal vein" which extends nearly the length of the blade
and more or less parallel to the margin but somewhat separated from it. Flowers
borne on axillary (or rarely terminal) branches, solitary or in specialized brac-
teate inflorescences with opposite branching, these modified in various ways,
e.g., by elongation of the axis and reduction of the lateral axes to one flower
each ("racemes"); by suppression of the axis and reduction of the lateral axes
to one flower each (flowers in "glomerules" or "umbelliform clusters"); by re-
duction of lateral axes to one pair, these arising just below the flower which
terminates the central axis ("dichasium"); by potentially indefinite elongation
of both central and lateral axes, this resulting in a "panicle" with proximal branches
elongate and a transition from these to short simple branches and terminal triads
of flowers. Flowers regular or essentially so, hermaphrodite or rarely by abor-
tion unisexual. Ovary inferior, the hypanthium adnate to the ovary its whole
length or prolonged beyond it so that the stamens, petals and calyx-lobes appear
to arise from the distal margin of a short tube surrounding the summit of the
ovary. Calyx-lobes usually 4 or 5, distinct and imbricate, or the calyx calyp-
trate and circumscissile, or rupturing irregularly in anthesis. Petals usually
4 or 5 (sometimes reduced in number or size, or wanting). Stamens usually
indefinitely many, in one-many series about the margin of the usually thickened
calycine disk, usually inflexed in the bud. Filaments usually filiform and distinct
in Guatemalan species. Anthers usually short, versatile or basifixed, bilocular,
opening (at least in Guatemalan genera) by longitudinal slits. Style simple,
elongate, with small capitate or peltate stigma. Ovary 2- to many-locular,
the placentae affixed to the axis or parietal and coalesced into a central axis,
the ovules 2 or more. Fruit fleshy or capsular. Embryo various.

The American Myrtaceae, with one exception noted below, are
all members of the tribe Myrteae, comprising 40 or more genera.
With few exceptions the native genera are exclusively American.
The total number of species recognized by Berg a century ago
was more than 1600.

283

284 FIELDIANA: BOTANY, VOLUME 24

In Guatemala there are a few introduced species belonging to
the tribe Leptospermeae, subfamily Leptospermoideae, a group that
is developed to the greatest extent in the Australian region. The
total number of species approaches 1000, including, according to
some authors, nearly 500 species of the vast Australian genus Euca-
lyptus. The only native American member of this subfamily is
the endemic Chilean species Tepualia stipularis (Barn.) Griseb.

One genus not a member of the Guatemalan flora, Calycolpus,
is represented in eastern Central America (Panama).

KEY TO THE TRIBES

(Guatemalan representatives only)

Fruit dry, capsular, consisting of the capsule immersed in the hardened hypan-
thium, the valves sometimes projecting beyond the rim of the hypanthium;
adult leaves all or nearly all alternate, the juvenile ones often differently
formed and more or less opposite Tribe I. Leptospermeae.

Fruit fleshy (a few- or many-seeded "berry"); leaves opposite, rarely a few sub-
opposite Tribe II. Myrteae.

Tribe I. LEPTOSPERMEAE DC.

Flowers in pedunculate axillary umbels; petals and calyx-lobes united into an
operculum which is deciduous at anthesis; large trees with elongate, long-

petiolate and often vertically hanging leaves Eucalyptus.

Flowers closely sessile in spikes at or near the tips of the branches; petals and
calyx-lobes distinct, soon deciduous; shrubs or usually small trees with narrow
and often short-petiolate leaves.
Leaves 1-2 cm. wide, with several conspicuous longitudinal nerves; stamens

creamy white, united in 5 bundles opposite the petals Melaleuca.

Leaves mostly 1 cm. wide or less, finely pinnately veined; stamens red to pink
or pale yellow, not united Callistemon.

A species of Callistemon, tentatively identified as C. salignus
DC., is planted frequently in parks and in grounds about houses,
for ornament, especially around Guatemala. It is a tall shrub or
a low tree, the branches usually more or less recurved and with
pendent tips; leaves alternate, lance-linear, about 4-5 cm. long,
long-attenuate, glabrous or nearly so, with thick margins; flowers
spicate, the apex of the spike usually continuing to grow and forming
a leafy branch; stamens long and conspicuous, pale yellow or pinkish;
fruit a small hard woody capsule.

The cajeput tree, Melaleuca Leucadendron L., is planted rather
frequently in Guatemalan parks, principally at middle or low eleva-
tions. The alternate leaves are narrowly lanceolate, acute or acum-
inate, pale green, with numerous conspicuous longitudinal nerves.
The creamy white flowers are borne in terminal spikes, whose tips
after anthesis grow into leafy branches. The fruit is a small woody

McVAUGH: FLORA OF GUATEMALA 285

capsule. The pale bark is thick and cork-like, and very resistant
to fire.

EUCALYPTUS L'Heritier. Eucalipto

Glabrous trees or shrubs, the leaves alternate, leathery, usually elongate,
lanceolate and 8-12 cm. long or more, markedly petiolate and often hanging
vertically; juvenile foliage (of seedlings or shoots from felled trees) often broad,
subsessile and more or less opposite, rarely hairy; flowers usually in pedunculate
axillary umbels, sometimes forming panicles; bracts and bracteoles deciduous
so early as to be seldom seen; ovary usually 3- to 4-locular, immersed in and
surrounded by the fleshy hypanthium which hardens in fruit and is prolonged
beyond the the summit of the ovary into a rim which bears the numerous stamens;
petals and calyx-lobes united into an operculum which is continuous with the
rim of the hypanthium in bud and circumscissile at an thesis; stamens widely
spreading in anthesis and forming the showy part of the flower; style about as
long as the operculum; ovules and seeds numerous, but only a few in each locule
fertile.

A large and almost exclusively Australian genus, at one time
supposed to include nearly 500 species; some recent authors have
suggested that the actual number is somewhat smaller. A recent
account of the Northern Australian species, by S. T. Blake (Austral.
Jour. Bot. 1: 185-352, t. 1-36. 1953), includes 50 species in this
part of the continent; J. M. Black (Fl. South Austral, ed. 2, 612-632.
1952) lists 52 species.

Numerous species have been introduced into the tropical and
warm-temperate regions of America for ornament, for purposes of
reforestation, for wood and for lumber; only the following seems
to have been entirely successful.

Eucalyptus globulus Labill. Voy. 1: 153, t. 13. 1799. Euca-
lipto; Ocalipto.

Planted abundantly in Guatemala, from the highlands down
almost to sea level; most common, according to Standley (mss.)
in Los Altos, at 1500 meters and higher. Native originally of Vic-
toria, New South Wales, and Tasmania, this species is now ex-
tensively planted and naturalized from California to Argentina and
Chile, especially in semi-arid regions.

A large tree, said to reach a height of 75-90 meters, with pale deciduous bark
and yellowish-green angled branchlets; adult leaves alternate, lanceolate and
often falcate to narrowly ovate, 2-3 (-6) cm. wide at base, 12-25 cm. long, (3-)
5-8 times as long as wide, attenuate from the base to the slenderly pointed apex,
the base often obliquely unequal-sided, abruptly rounded to the flexuous petiole
2-4 cm. long; midvein pale, flat or concave above, convex beneath; leaf -margins

286 FIELDIANA: BOTANY, VOLUME 24

bordered by heavy cartilaginous veins about equal to the midvein but compressed
at right angles to the plane of the leaf and often standing somewhat above and
below it; lateral veins delicate and inconspicuous, joining an equally slender
and nearly straight submarginal vein just within the cartilaginous border; foliage
with numerous small dark glands on both surfaces; juvenile shoots and leaves
conspicuously whitened and waxy-glaucous, the leaves opposite, sessile, ovate
to oblong, cordate, abruptly short-acuminate at tip, 4-5 cm. wide, 7-15 cm.
long; flowers large, subsessile, solitary or rarely 3 in an axil, on a massive, broadly
2-angled peduncle up to 5 mm. long, buds 1.7-2.5 cm. long, conspicuously whitened
by a heavy waxy-farinose coating; hypanthium about 1 cm. long, truncate at
base, strongly 4-angled, obpyramidal, irregularly and coarsely warty-roughened
especially on the angles and the thickened margin; calyptra dome-like, roughened
like the hypanthium, usually with a broad knoblike or acute central beak; sta-
mens 1.5 cm. long (the flower when expanded 3-4 cm. across), borne on the inner
edge of the disk which projects about 3 mm. beyond the thickened margin of
the hypanthium; style 8-10 mm. long; fruit 2-2.5 cm. broad and high, flat-topped
or the surface convex, the 4-5 valves not exserted but nearly plane with the
surface; seeds 1-3 mm. long, very numerous, prismatic, irregularly several-angled.

The following notes are by Mr. Standley: These are trees of
rapid growth, sometimes said to attain (in Australia) a height of
45 meters and a trunk diameter of more than a meter when no
more than 30 years of age. In North America they grow vigorously
but have found little favor because native woods are superior to
them. In Central America the trees fortunately are little planted
at the present time. It is presumably this species [E. globulus]
that is most planted in Guatemala, but several others are in culti-
vation there. To judge from the size of some of the existing trees,
the eucalyptus must have been introduced into the Guatemalan
highlands many years ago, just why we do not know; perhaps on
account of its rapid growth. Now that the trees are grown, no
use is made of them except as a source of shade, of which sufficient
and more may be obtained more satisfactorily from other trees.
The trees have a wide mass of roots, within whose circumference
no other plants can grow. Little use is made of the wood; when
large trees are felled it is all but impossible to get rid of the trunks.
They do not burn readily, and the wood is so hard it is unsatis-
factory for fuel. There are some very fine and giant trees about
Chimaltenango, Quezaltenango, and other places in the highlands,
sometimes in handsome avenues along the roads. When the trees
are old the bark peels off in great strips, covering the ground or
hanging untidily from the trunks. The leaves, too, often fall in
great quantities.

Eucalyptus trees are well known as the source of eucalyptus
oil, which has a very distinctive odor and is used in medicine,

McVAUGH: FLORA OF GUATEMALA 287

especially in treating affections of the respiratory system. Between
Guatemala and Amatitlan there is a large grove where oil is ex-
tracted from the leaves. Small bunches of the branches and leaves
are sold commonly in the Guatemalan markets for use in domestic
medicine. The dry leaves are often burned in houses to "disinfect"
them.

Because of their peculiar glaucous coloring it is generally possible
to recognize eucalyptus trees from a great distance. They appear
altogether alien in the Central American landscape. On the dry
mountains above Quezaltenango they are the most conspicuous
of all trees, towering high above all the rest, and dwarfing even the
famous cipreses romanos of that region.

Tribe II. MYRTEAE DC.

KEY TO FLOWERING MATERIAL

Inflorescence a compound pedunculate panicle, usually many-flowered, the lower
branches of the panicle opposite and themselves compound [panicles reduced
to spikes or 3-flowered dichasia in a few species of Calyptranthes with calyp-
trate calyx]; bracts and bracteoles usually deciduous at or before anthesis.

Buds calyptrate, circumscissile, the calyx entirely closed in the bud or else trun-
cate, the lobes then scarcely discernible, soon deciduous, the tube (hypan-
thium) after anthesis naked, truncate at the mouth.

Calyx closed in bud, the calyptra consisting of a part of the calyx; petals
minute or often wanting, mostly adherent to the inside of the calyptra
and falling with it; inflorescences axillary (sometimes falsely terminal),
usually at leafy nodes; dibrachiate hairs often present and conspicuous,
forming the greater part of the pubescence if any; ovules 2 in each of

two locules Calyptranthes.

Calyx open and truncate in the bud, the calyptra consisting of the united
petals; inflorescences terminal or (usually) lateral at leafless nodes on
old wood; plants glabrous; ovules numerous Eugenia Cumini.

Buds not calyptrate, the calyx open in the bud, its lobes evident and persistent
in flower and in fruit; petals free, conspicuous, falling individually; dibrachi-
ate hairs none or inconspicuous; inflorescences axillary; ovules 1 or 2 in
each of two locules.

Calyx-lobes 4; style prominently capitate, the stigma twice as thick as the
style; vigorous branchlets often 4-angled; petioles 1.5-2 (-3) cm. long;

herbage strongly fragrant Pimenta.

Calyx-lobes 5; style not at all or scarcely capitate; branchlets terete or com-
pressed; petioles less than 1 cm. long; herbage not or but slightly fragrant.

Myrcia.

Inflorescences usually not compound, or if so dichasially forked with a sessile ter-
minal flower in each fork; flowers sometimes solitary and axillary, mostly in
racemes with few opposite and decussate pairs of flowers; axis of the raceme
often so short that the flowers appear fasciculate or glomerate; bracts and/or
bracteoles often persistent; ovules usually more than 2 in each locule; locules
usually 2, sometimes several.

288 FIELDIANA: BOTANY, VOLUME 24

Calyx in bud closed or essentially so, without definite lobes, sometimes with an
apical porelike aperture, opening by irregular longitudinal fissures, or cir-
cumscissile.
Calyx calyptrate, circumscissile, the calyptra often remaining attached at

one side.

Petals minute, often adhering to the calyptra and falling with it; hypan-
thium persisting in a ring on the fruit, usually not splitting longitudi-
nally; ovules 2 in each locule; dibrachiate hairs usually present and
conspicuous; flowers 3 or more in a dichasium-like or irregularly spike-
like inflorescence Calyptranthes.

Petals evident, sometimes showy, falling independently; hypanthium usu-
ally splitting longitudinally into several irregular lobes; ovules numer-
ous; dibrachiate hairs none; flowers solitary and axillary, or sometimes
in 3-flowered dichasia with the central flower sessile Psidium.

Calyx opening by irregular longitudinal fissures, the 3-5 resulting lobes irreg-
ularly shaped, one sometimes larger than the others Psidium.

Calyx-lobes normally developed even in the bud, the calyx at anthesis some-
times splitting beyond the bases of the lobes.
Calyx-lobes 5; flowers axillary and solitary, or in a 3-flowered dichasium.

Bracteoles foliaceous and persistent even in the fruit, 3-8 mm. long; anthers
sagittate, the connective dilated; flowers solitary; shrubs of high moun-
tains, with leaves 3 cm. long or less Ugni.

Bracteoles deciduous at or about the time of anthesis; anthers not sagittate,
the filaments filiform; flowers solitary or occasionally 3 in a dichasium;
small shrubs of savannahs at middle and low elevations Psidium.

Calyx-lobes 4.

Inflorescence a raceme, the flowers in opposite decussate bracteate pairs;
axis of the raceme often so much abbreviated that the flowers super-
ficially appear to be glomerate or umbellate; terminal flower of the
axis usually wanting.

Hypanthium much prolonged beyond the summit of the ovary, forming
a funnelform, campanulate or bowl-shaped "calyx-tube" bearing the
stamens about its margin.

Inflorescence terminal, or lateral at leafless nodes on old wood; hypan-
thium persistent, not circumscissile, attenuate at base to a narrow
stipelike pseudostalk above the place of attachment of the decid-
uous bracteoles; flowers large, the buds 6 mm. long or more, the
style 8 mm. long or more; ovules numerous, 6-8 or more in each
locule; ["Syzygium"] Eugenia.

Inflorescences borne in the axils of foliage leaves; hypanthium circum-
scissile at base after anthesis, not attenuate, usually partly covered
by the conspicuous paired persistent involucre-like bracteoles;
ovules 2 (-4) in each locule; flowers usually 4, nearly sessile, small,
the buds mostly 2.5-3 mm. long, the style 4-6 mm. long.

Myrdaria.

Hypanthium little or not at all prolonged, not circumscissile, persistent
with the calyx-lobes in flower and fruit; inflorescence various, never
terminal; ovules usually numerous, rarely fewer than 5 in each locule.

Calyx-lobes partly or almost wholly united in the bud, only their tips
free, the cracks between the lobes extending at anthesis farther
than the length of the free tips, and reaching the edge of the

McVAUGH: FLORA OF GUATEMALA 289

staminal disk or beyond; flowers in small clusters at usually leaf-
less nodes, apparently without definite arrangement on a racemose
axis; leaves often subsessile, mostly rounded to subcordate at base,
the petioles 5 mm. long or less and very short in proportion to the
blade; lowlands near the Atlantic Coast Psidium.

Calyx-lobes distinct in bud and in anthesis, not at all or scarcely sep-
arating in anthesis and in fruit; inflorescence racemose, the axis
of the raceme usually evident even if very short, and the flowers
in pairs on the axis Eugenia.

Flowers solitary and axillary, or in simple or compound dichasia, the pri-
mary axis of the inflorescence if once-forked with a sessile or nearly
sessile flower in the fork.

Flowers mostly 3 or 7 in a dichasium, the sessile terminal flowers in the
forks developing earlier than the ones at the tips; flowers, if occa-
sionally solitary, arising from axils of ordinary leaves; bracts and
bracteoles inconspicuous, falling at anthesis or before; calyx-lobes
large in proportion to the hypanthium, 2.5-3.5 mm. wide.

Myrcianthes.

Flowers solitary or occasionally 3 in a dichasium, if solitary then mostly
confined to the 1-2 lowermost nodes of a new leafy branch, and then
subtended by small bracts rather than by foliage leaves; flowers if
more than 3 in elongated or much abbreviated racemes.

Calyx-lobes minute, or united except at the tips, the calyx at anthesis
opening by irregular longitudinal cracks extending usually to the
summit of the ovary Psidium.

Calyx-lobes distinct in bud and in anthesis, not at all or scarcely sep-
arating in anthesis and in fruit Eugenia.

KEY TO FRUITING MATERIAL

Cotyledons foliaceous, folded and contorted; radicle elongate; testa mostly mem-

branaceous, fragile; calyx-lobes, if present, 5; inflorescence compound, often

many-flowered, with branches opposite near base and irregularly ternate or

solitary near tips; bracts and bracteoles not present at this time.

Calyx-lobes normally developed and present on the fruit Myrcia.

Calyx wanting (the tip of the fruit umbilicate) or represented by a shrunken
calyptra attached at one side Calyptranthes.

Embryo not as above, but undivided and homogeneous, or elongate and curved
or coiled, or the cotyledons large and plano-convex; calyx-lobes usually 4 and
distinct if the flowers are more than 7; bracteoles often persistent.

Inflorescence compound, many-flowered, with branches opposite near the base
of the panicle, and irregularly ternate or solitary near the tips.

Calyx-lobes 4, persistent; inflorescences in the axils of the upper leaves; seeds
usually 2, small, the embryo elongate, curved into a double spiral.

Pimenta.

Calyx-lobes deciduous, the fruit surmounted by the collar-like remains of the
hypanthium; seed usually 1, large, the cotyledons plano-convex, unequal;
inflorescences sometimes terminal, usually lateral at leafless nodes on

old wood Eugenia Cumini.

Inflorescence a simple raceme or if compound a dichasium, or the flowers soli-
tary or apparently glomerate or fasciculate.

290 FIELD IANA: BOTANY, VOLUME 24

Inflorescence a raceme, the flowers in opposite decussate bracteate pairs; axis
often so short that the flowers appear glomerate or fasciculate; terminal
flower usually not developing; calyx-lobes 4; embryo massive, undivided,
or the cotyledons imperfectly separated.

Hypanthium prolonged above the ovary, circumscissile at base after an-
thesis, leaving a circular scar on the fruit; fruit edible, soft-fleshy and
juicy, nearly sessile Myrciaria.

Hypanthium not circumscissile, scarcely or not at all prolonged; calyx-
lobes mostly persistent on the fruit; fruit scarcely edible, with meager
flesh, variously disposed Eugenia.

Flowers solitary or in simple or compound dichasia, the primary axis of the
inflorescence if once-forked with a sessile or nearly sessile flower in the
fork; calyx-lobes 4 or 5, or the calyx irregularly, longitudinally dehiscent.

Seeds one or two; embryo undivided or the cotyledons large and plano-
convex; testa thin; calyx-lobes 4, usually distinct and persistent.

Embryo undivided; flowers solitary, or racemose in some axils . .Eugenia.

Cotyledons fleshy, large, distinct, plano-convex; flowers 3 or 7 in a dicha-
sium, or a few solitary Myrcianthes.

Seeds several or many; embryo arcuate ("C-shaped"), with long radicle,
short inconspicuous cotyledons and usually bony testa; calyx-lobes
4 or 5, or the calyx splitting irregularly.

Bracteoles foliaceous, persistent, 3-8 mm. long; flowers solitary; calyx-
lobes 5, persistent, not separating at the base in fruit; shrubs of high
mountains, with small coriaceous leaves 3 cm. long or less .... Ugni.

Bracteoles usually deciduous at or before anthesis, if occasionally per-
sistent small and scarious; flowers solitary or in a 3-flowered dicha-
sium, or several together at a leafless node; hypanthium somewhat
irregularly split down to the summit of the fruit, the remnants of
the calyx 3-5, often persisting, the broken tissue along their margins
usually evident Psidium.

CALYPTRANTHES Swartz

References: Berg, Linnaea 27: 18-33. 1855; and in Mart. Fl.
Bras. 14, pt. 1: 38-55. 1857; McVaugh, Fieldiana, Bot. 29: 397-
412. 1963.

Calyx completely closed in the bud, circumscissile, the operculum usually
attached at one side in anthesis, finally completely dehiscent. Petals none,
or 2-3 (-5), small and inconspicuous and often attached to the operculum (ca-
lyptra) and falling with it. Paired panicles arising from opposite axils at the
lowest node of an abortive axis, or the axis prolonged and leafy and the paired
panicles from the lowest node, subtended by small or foliaceous bracts. Pubescence
usually at least in part of dibrachiate hairs.

A distinctive American genus of perhaps 50-75 species, ranging
from Florida through the West Indies and eastern South America
to Uruguay. A few species not treated below occur in Costa Rica
and Panama, and a few additional species in Mexico.

McVAUGH: FLORA OF GUATEMALA 291

Members of this genus may often be recognized, even in the
sterile condition, by the characteristically forking branchlets and
the consequently somewhat zigzag branches. This habit results
from the abortion of the terminal buds, and the concomitant develop-
ment of axillary branchlets, at the growing tips. Potentially florif-
erous axillary branchlets often abort in the same manner, at the
first node above the axil; subsequent development of the opposite
floriferous secondary branches from the same node produces the
so-called "paired panicles," which often at first glance seem to
arise directly from the axil. A more careful examination usually
reveals the fact that these two panicles seem to arise from the tip
of a very short axillary branch.

In some species of Calyptranthes (e.g. C. Bartlettii, C. Karlingii)
the inflorescences ordinarily develop at the terminal node of an
old leafy shoot. If, as often happens, an axillary (floriferous)
branchlet arises from only one side of the terminal node, then in its
growth it pushes aside the undeveloped terminal vegetative bud
and becomes aligned with the older part of the leafy branchlet
so as to appear terminal rather than axillary. The lowermost
secondary branches of this axillary (but apparently terminal) shoot
are floriferous (the "paired panicles"). Interpretation of the mor-
phology of this inflorescence may be even more difficult because
the central shoot between the paired panicles (which ordinarily
aborts in axillary inflorescences) now often elongates and becomes
leafy above the panicles, ultimately simulating a new terminal
leafy branch, this bearing at its lowermost node a pair of opposite
panicles subtended by bracts (not normal leaves).

Leaf-bases broadly cordate or subcordate, the blades nearly sessile or subclasping,
on stout short petioles 3 mm. long or less; blades usually acute or obtuse,
sometimes obscurely acuminate, never caudate; young leafy branchlets 2- or
4-angled or -winged except in C. chiapensis with leaves 12-23 cm. long.

Young branchlets and inflorescence densely ferruginous-tomentose with con-
torted and mostly 1- or 2-branched but nearly erect hairs 0.4-0.7 mm. long.

Leaves lanceolate, 2-5 cm. wide, 6-12 cm. long; branchlets bicarinate; British
Honduras C. Bartlettii.

Leaves oblong, 6.5-7.5 cm. wide, 12-23 cm. long; branchlets terete or com-
pressed, not carinate; Chiapas C. chiapensis.

Young branchlets and inflorescence glabrous or thinly pubescent with appressed
flat dibrachiate hairs 0.1-0.2 mm. long; branchlets 2- or 4-angled or -winged.
Branchlets biconvex in cross section, bicarinate, the winglike angles termi-
nating distally in the midlines of the leaf-bases C. Contrerasii.

Branchlets 4-angled, two of the angles terminating distally in the midlines
of the leaf-bases, the other angles winged, terminating distally between
the leaf-bases C. Karlingii.

292 FIELDIANA: BOTANY, VOLUME 24

Leaf -bases acute or cuneate or sometimes abruptly rounded, never cordate; blades
usually definitely petiolate, often acuminate, sometimes caudate; branchlets
various.

Flowers 9 or fewer in each panicle, the two lateral panicle-branchlets and the
terminal one each bearing a single pedicellate flower or a cluster of 2-3
sessile flowers; inflorescence 3-5 cm. long, the very slender peduncle usually
much longer than the rest of the panicle; leaves 3-7.5 cm. long, long-
acuminate.

Leafy branchlets 4-angled or -winged, glabrous or essentially so; panicles gla-
brous, 3- or 5-flowered, the peduncle filiform C. Hylobates.

Leafy branchlets terete, at least when young with abundant long straight
appressed dibrachiate hairs; panicles pubescent, about 9-flowered, the
peduncle up to about 1 mm. wide C. paxillata.

Flowers more numerous, usually 20-30 or more in each panicle, if only 9 or fewer
the leaves 10-16 cm. long, the buds 7-8 mm. long; panicles usually 5 cm.
long or more, twice or more compound, the flowers often sessile or sub-
sessile in small clusters near the tips.

Plants glabrous or sparingly pubescent; pubescence, if any, usually consisting
of minute flat appressed dibrachiate hairs 0.1-0.3 mm. long, these often
nearly confined to the base of the hypanthium, the nodes of the inflores-
cence, and the undeveloped vegetative buds.

Panicles 15-20 cm. long; peduncles compressed, 10-12 cm. long, 3-4.5 mm.
wide; leaves, as far as known, 11-12 cm. wide, up to 35 cm. long;
inflorescence and buds thinly reddish-pubescent; Pet6n-British Hon-
duras boundary C. megistophylla.

Panicles mostly 10 cm. long or less (infrequently up to 15 cm. long);
peduncles 3-5 cm. long, less than 2 mm. wide; leaves mostly 12-15 cm.
long or less.

Leaves prevailingly obovate to elliptic, obtuse or rounded at tip, 3.5-
6.5 cm. long; small connecting veins evident on the lower surface
in dried leaves as a raised reticulum; branchlets not wing-angled;
pubescence appressed, rather sparse, grayish- or yellowish- white;
British Honduras C. cuneifolia.

Leaves prevailingly elliptic to lanceolate or ovate, acute or acuminate
(if small and obtuse then the branchlets winged, the small veins
obscure, the pubescence if any reddish).

Leaves small and obtuse, mostly 2.5-4.5 cm. long, usually narrow,
often varying on the same plant from 2-5 (-7) times as long as
wide; branchlets persistently and prominently wing-angled; pubes-
cence sparse, evanescent, of rather loosely appressed yellow- or
reddish-brown hairs; Honduras to British Honduras.

C. hondurensis.

Leaves usually larger and acute or acuminate, usually 2-3 times as
long as wide; branchlets wingless or somewhat obscurely winged.

Flower-buds 3-4.5 mm. long, glabrous; branchlets not winged; whole
plant essentially glabrous; calyptra conic, thickened, 4 mm.
wide; persistent collar-like base of the calyx 3-4 mm. wide in
fruit; leaves 10-14 cm. long, prominently acuminate; Chiapas.

C. perlaevigata.

Flower-buds 2-3 mm. long; plants usually sparingly pubescent;
calyptra 1.5-2.5 mm. wide; persistent collar-like base of the
calyx 1.5-3 mm. wide in fruit.

McVAUGH: FLORA OF GUATEMALA 293

Plants glabrous except for a few hairs on the youngest vegetative
parts; leaves elliptic-lanceolate, often 3-4 times as long as
wide; petioles 7-10 mm. long; branchlets not winged; El Sal-
vador and British Honduras C. Calderonii.

Flower-buds and distal nodes of the panicle sparingly pubescent

with appressed reddish or pale hairs; leaves mostly elliptic,

2-3 times as long as wide; petioles 2.5-5 mm. long.

Leafy branchlets (at least some of them) wing-angled in most

plants; inflorescence rather slender, the principal secondary

branches of the panicle less than 1 mm. wide; mountains

of Chiapas C. pollens var. mexicana.

Branchlets not wing-angled; inflorescence stout, the principal

secondary branches of the panicle much flattened, their

lowest internodes often 6-8 mm. long, 1.5 mm. wide;

Veracruz; British Honduras to Yucatan. . .C. Millspaughii.

Plants rather generally pubescent on the young growth and the inflorescence,

often softly and loosely so (at least some of the hairs then branched and

spreading or erect) ; hairs often up to 0.5 mm. long, at least to some extent

persistent on the branchlets, the inflorescence and the fruits.

Leaves prevailingly obovate to elliptic, obtuse or rounded at tip, 3.5-
6.5 cm. long; branchlets not winged; small connecting veins evident
on the lower surface in dried leaves as a raised reticulum; pubescence
appressed, rather sparse, grayish- or yellowish- white; British Hon-
duras C. cuneifolia,

Leaves prevailingly elliptic to lanceolate or ovate, acute or acuminate (if
small and obtuse the branchlets winged, the small veins obscure, the
pubescence reddish).

Leaves small and obtuse, mostly 2.5-4.5 cm. long, usually narrow, often
varying on the same plant from 2-5 (-7) times as long as wide;
branchlets persistently and prominently wing-angled; pubescence
sparse, evanescent, of rather loosely appressed yellow- or reddish-
brown hairs; Honduras to British Honduras C. hondurensis.

Leaves usually larger and acute or acuminate, if narrow usually consist-
ently so; branchlets usually wingless; pubescence usually copious.
Young leafy branchlets softly brown- to rufous-tomentose with hairs
of two types: Straight and nearly erect, simple or short-branched
hairs up to 2 mm. long, these standing above the tomentum of
contorted and somewhat appressed hairs with two equal or
strongly unequal branches; hairs of the panicles of the contorted,
appressed type; buds 3 mm. long or less, obovoid, mostly short-
apiculate; branchlets not winged.

Leaves narrowly elliptic-lanceolate, 1-2.3 cm. wide, 4-6 times as
long as wide C. Lindeniana.

Leaves elliptic-ovate, 4-6 cm. wide, 2-3 times as long as wide.

C. Chytraculia var. americana.
Pubescence of leafy branchlets not as above, not noticeably different

from that of the panicle branches, or if so more appressed.
Hairs coarse, brown to yellowish brown, 0.3-1.5 mm. long, more or
less erect and the branchlets tomentose; buds as far as known
4.5 mm. long or more; branchlets not wing-angled.
Leaves broadly rounded at base; lateral veins impressed above;
petioles 8-11 mm. long; buds ovoid, short-apiculate, 4.5-

294 FIELDIANA: BOTANY, VOLUME 24

6 mm. long; tomentum dense, the very numerous erect hairs
simple or forked, up to 1.3-1.5 mm. long C. macrantha.

Leaves acute at base; lateral veins not impressed above; petioles
5-7 mm. long; buds unknown, the calyptra shallow, short-
apiculate; tomentum bristly, the very coarse hollow erect
forked hairs up to 0.3-0.7 mm. long C. zanquinensis.

Hairs minute, dibrachiate, often appressed, sometimes crisped,
mostly 0.1-0.3 mm. long, the pubescence of the young branch-
lets and inflorescence rusty brown, yellowish brown or, espe-
cially on the leaves, nearly colorless; buds 3 mm. long or less.

Branchlets wing-angled; leaves cuneate at base, often 4-7 cm. long,
usually glandular-punctate above; inflorescence copiously
soft-pubescent with reddish or sometimes pale hairs; West
Indies; southern Florida; Isthmus of Tehuantepec; Yucatan.

C. pollens var. pollens.

Branchlets terete or compressed, not wing-angled or occasionally
so; leaves acute at base, varying to somewhat rounded or
occasionally to cuneate; branches of the inflorescence usually
sparingly pubescent except about the distal nodes and the
hypanthium; Mexico and Central America.

Inflorescence stout, the principal secondary branches of the
panicle much flattened, their lowest internodes often 6-
8 mm. long, 1.5 mm. wide; pubescence of very small flat
closely appressed longitudinally oriented hairs; British
Honduras to Yucatan C. Millspaughii.

Inflorescence more slender, the internodes of the secondary
panicle-branches terete or slightly compressed, 1 mm. wide
or less; pubescence at least in part of crisped or soft hairs;
mountains of Mexico and Guatemala.

Leaf -blades often 7-12 cm. long, eglandular above; petioles
8-10 mm. long; branchlets not wing-angled; panicles
often 50-60-flowered C. pendula.

Leaf-blades often 3-7 cm. long, often glandular-punctate
above but sometimes inconspicuously so; petioles 2.5-
5 mm. long; branchlets (at least some of them) wing-
angled in most plants; panicles 30-35-flowered.

C. pattens var. mexicana.

Calyptranthes Bartlettii Standl. Field Mus. Bot. 11: 136. 1932.

River banks, rocky ledges along streams, wet forest; known only
from El Cayo District, British Honduras (the type from Mountain
Pine Ridge, H. H. Bartlett 11837).

A shrub or small tree up to 3.5 meters high; branchlets glabrate, bicarinate,
the keels terminating distally in the median lines of the leaf -bases; branchlets
when young, and the inflorescence, densely ferruginous-tomentose with more
or less erect, flaccid, often contorted and 1- or 2-branched hairs 0.4-0.7 mm.
long; leaves coriaceous, glabrate, lanceolate, nearly sessile, (1-) 2-5 cm. wide,
6-12 (-18) cm. long, 3-6 times as long as wide, acute at tip, cordate at base, the
margins abruptly decurrent on the inner angles of the very stout petiole 1-2 mm.
long and up to 2.5 mm. thick; midvein concave or sulcate above; lateral veins

FIG. 49. Flowers and inflorescence in Calyptranthes and Myrcia. A, Slightly
immature fruit of C. Bartlettii (Lundell 6686); X 5. B, Flower of C. Calderonii,
slightly past anthesis (Lundell 6857); X 5. C, Flower, and buds of two ages, of
M.fallax (Gabriel s.n. in 1802); X 10. The flower is seen from above, the summit
of the ovary surrounded by the staminal ring and the five calyx-lobes; the petals,
stamens and style have fallen. D, Seed of C. Chytraculia var. americana, the outer
coats removed to show the contorted cotyledons (Gentle 1363); X 2.5. E, The
myrcioid panicle, in C. pattens var. mexicana; shown is one of the paired panicles
at the base of a leafy shoot (Wilbur & Wilbur 1415); X 1.5.

295

296 FIELD IANA: BOTANY, VOLUME 24

15-20 pairs in addition to numerous intermediate ones; marginal vein equaling
or stronger than the laterals and but little arched between them, 1-2 mm. from
the margin, usually with a smaller submarginal vein beyond it; leaves dark green
and smooth above, paler beneath; glands often obscure; panicles 4-10 cm. long,
about 30-60-flowered, 3 (-4) times compound, usually apparently single (not
paired), each arising at the lowest node of a new leafy shoot and subtended by
a leafy bract 8-25 mm. long; peduncles (2-) 4-6 cm. long; branches of the panicle
subtended by foliaceous boat-shaped bracts, the smallest of these (those sub-
tending the terminal clusters of 3 or more sessile flowers) 2-3 mm. long; buds
at maturity 4.5-5 mm. long, obovoid, broadly apiculate; calyptra bowl-shaped,
about 3 mm. broad; hypanthium at anthesis widely flaring and bowl-shaped;
style 9 mm. long; stamens on the margin of the hypanthium, about 100, up to
8-9 mm. long; petals 2 (-3), tiny and strap-shaped, adhering to the inner surface
of the calyptra and falling with it; fruit dark red to purple, globose, about 1 cm.
in diameter, tipped by a flaring collar 1-2 mm. high and 4-5 mm. wide, the ca-
lyptra often persistent as well; seeds lustrous, compressed-orbicular, 5 mm. wide.

Calyptranthes Calderonii Standl. Field Mus. Bot. 4: 318. 1929.

Along a stream, Vaquero, El Cayo District, British Honduras,
C. L. Dandell 6857; El Salvador (Cerro de Apaneca, S. Calderdn 2423,
type).

A shrub or treelet to 3 meters high, the branchlets terete or compressed;
plants glabrous except for a few flat reddish-brown dibrachiate hairs (0.1-0.2) -0.5
mm. long, on the youngest vegetative parts; leaves coriaceous, elliptic or elliptic-
lanceolate, 1.5-3 cm. wide, 5-10 cm. long, about 3 (2.5-4) times as long as wide,
broadly or gradually acuminate at tip, the very tip obtuse; base acute, the mar-
gins decurrent on the inner angles of the channeled petiole 7-10 mm. long, 1-1.5
mm. thick; midvein impressed above at least near base; lateral veins 10-12 pairs,
obscure in mature leaves; marginal vein about equaling the lateral ones, 0.5-1 mm.
from the margin; blades dark green above, paler beneath; glands obscure, the
upper surface somewhat impressed-punctate; panicles 4-6 cm. long, 3 times
compound, paired at the first node of an abortive axillary branch, or opposite
at the lowest (leafless) node of a leafy shoot; peduncle 2-3.5 cm. long, somewhat
compressed, 1-1.3 mm. wide at the summit; flowers about 40 or fewer, in 2's
and 3's near the tips; buds 2.5-2.7 mm. long, obovoid, apiculate; calyptra domed,
1.5-1.7 mm. wide; hypanthium funnelform, 1.5 mm. long and wide; style 5-6 mm.
long; stamens about 50, up to about 5 mm. long; petals ?0, or 1-2, narrow, attached
to the calyptra and falling with it; fruit unknown.

Calyptranthes chiapensis Lundell, Contr. Univ. Mich. Herb.
7: 28. 1942.

Subdeciduous forest, often along streams, 1000 meters or less.
To be expected in Guatemala. The type, Matuda 4488, came from
Chicomuselo, Chiapas, less than 50 km. from the Guatemala border.

A tree up to 10 meters high, the inflorescence, branchlets and young leaves
velutinous with compact purplish-brown tomentum, the mostly upright simple

McVAUGH: FLORA OF GUATEMALA 297

or forked hairs up to about 1 mm. long; leaves oblong or oblong-ovate, cordate
and subamplexicaul, 3.5-7.5 cm. wide, 12-23 cm. long, 3-4 times as long as wide,
the tip acute or obscurely acuminate, the base broadly rounded, divided by a
narrow sinus into two rounded auricles 5-12 mm. long; petiole scarcely apparent
from above in dried specimens, 7-12 mm. long as seen from below including the
enlarged basal portion of the costa, 2.5-4 mm. thick including the hairs; midvein
as seen from above with central concave channel separating parallel convex
ridges; lateral veins 15-25 including some weak intermediate ones, in dried leaves
convex but inconspicuous above, well differentiated beneath; marginal vein about
equaling the lateral ones and somewhat arched between them, 2-7 mm. from the
margin, often with a series of submarginal arches, connected by short straight
veins, beyond it; blade lustrous above, paler and dull beneath, obscurely glandular;
inflorescence stout, many-flowered, the paired panicles up to 15 cm. long, about
3 times compound, arising from an abortive axillary axis or from the lowermost
(bracteate) node of a leafy branchlet, the bracts then lanceolate, foliaceous, up
to 2 cm. long; branches of the panicle proliferous, the flowers in small groups
near the tips; peduncle 4.5-7 cm. long, up to 4 mm. wide; buds subglobose to
obovoid (actually short-apiculate but appearing rounded because of the dense
covering of hairs), 3.5-4 mm. long; calyptra shallowly convex, membranous,
soon explanate, 3-3.5 mm. wide; style 7 mm. long; stamens about 125, up to
5.5 mm. long, the anthers 0.4-0.5 mm. long; petals 2, minute, narrow, 2 mm.
long, attached to the calyptra and falling with it; fruit apparently black, globose
to oblate or didymous, 7-10 mm. high, 7-12 mm. wide, surmounted by the per-
sistent collar-like neck of the hypanthium 2.7-3 mm. wide.

Calyptranthes Chytraculia (L.) Sw. Prodr. 79. 1788. Myrtus
Chytraculia L. Syst. Nat. ed. 10. 1056. 1759.

The typical variety of the species is known only from Cuba and
Jamaica; the species is represented in Guatemala by the following
variety, differing from the var. Chytraculia in its somewhat larger
leaves and in the paler pubescence (cf. Fieldiana, Bot. 29: 403. 1963).

Calyptranthes Chytraculia var. americana McVaugh, Fieldi-
ana, Bot. 29: 404. 1963. Pimiento; Guayabillo.

Lowlands, sometimes on limestone, often along rivers or on lake
shores, 200 meters or less; Pete"n (the type from Uaxactun, Bartlett
12447); Alta Verapaz; Izabal. British Honduras; Nicaragua; Pan-
ama; northern Colombia.

A tree to 10 meters high, the branchlets terete; inflorescence and often the
young branchlets shortly tomentulose with appressed or loosely spreading lustrous
pale brown to reddish brown dibrachiate hairs; hairs of the panicles mostly much
contorted, loosely appressed and with two equal or strongly unequal branches;
hairs of the young leafy branchlets partly as above, partly of nearly straight,
erect, simple or short-branched hairs up to 2 mm. long; foliage glabrate; leaves
elliptic to ovate or sometimes obovate, (2.5-) 4-6 cm. wide, (5.5-) 7-13 cm.
long, 2-2.5 (-3) times as long as wide; blades acuminate at tip, acute to rounded

298 FIELDIANA: BOTANY, VOLUME 24

at base, the margins decurrent on the channeled petiole 7-10 mm. long; midvein
impressed above at least near base; lateral veins 12-15 pairs, stronger than the
intermediate ones, more prominent on the lower surface; marginal vein about
equaling the lateral ones and slightly arched between them, 1-1.5 (-3) mm. from
the margin; leaves dark green and probably lustrous above, paler and usually
gland-dotted beneath; inflorescence 7-14 cm. long, the principal (paired) panicles
mostly 3 times compound, in the axils of terminal nodes of an old branchlet, or
sometimes basal and opposite on a leafy shoot from the same node, the peduncles
2-4 (-7) cm. long; flowers often 80-100, mostly in clusters of 3 or more near the
tips; buds obovoid, 2-3 mm. long, scarcely apiculate; calyptra dome-shaped,
1.5-1.8 mm. wide, remaining attached at one side (sometimes even in the mature
fruit); style 4-5 mm. long; hypanthium hairy inside around the base of the style;
stamens about 60, up to 3.5 mm. long; anthers 0.3 mm. wide, broader than long;
petals about 2, oblong or irregularly shaped, less than 1 mm. long, adherent
to the calyptra and falling with it; fruit globose, red (or yellow?), finally black,
about 7 mm. in diameter, glabrous at maturity but usually with some tomentum
about the persistent neck of the calyx.

Specimens from Guatemala have been confused with Calyp-
tranthes Millspaughii, which is quite a different species, lacking
the shaggy pubescence that is characteristic of C. Chytraculia.

Calyptranthes Contrerasii Lundell, Wrightia 2: 205. 1961.

In moist or wet forest, at low elevations, 200 meters or less;
Pete"n (Dolores, in high forest, E. Contreras 2360, the type); Alta
Verapaz. Endemic.

A tree 6-8 meters high, nearly glabrous; branchlets bicarinate, the wings
terminating distally in the median line of the leaf -base; vegetative buds appressed-
pubescent with flat rusty dibrachiate hairs about 0.1 mm. long; leaves subsessile,
coriaceous, ovate to oblong or elliptic, 1.5-4 cm. wide, 4.5-10 cm. long, 2-2.5 (-3)
times as long as wide; tips bluntly pointed or rounded, the base cordate and
subclasping, the margins abruptly decurrent on the very stout petiolar base 1.5-2
mm. long and wide; midvein sulcate above; lateral veins 15-20 pairs including
some intermediate ones; marginal vein about equaling the lateral ones and some-
what arched between them, 1.5-3 mm. from the margin; leaves "dark green and
shining above, pale green beneath" (Steyermark), thickly glandular-punctate
above when dry, obscurely glandular beneath; inflorescence a panicle, bracteate
at base, simulating a terminal leafy shoot with floriferous branches at its lowest
node (as in C. Bartlettii and C. Karlingii) ; bracts at the base of the panicle branches
6-7 mm. long, 3 mm. wide; paired panicle branches 2.5 cm. long, 5-flowered,
the peduncle 1.5 cm. long, more than 1 mm. wide; buds turbinate, glabrous,
3.5 mm. long, attenuate-stipitate at base, prominently apiculate.

Known only from the type collection, which is in bud, and from
a sterile specimen, Steyermark 45628, collected on Cerro Chinaja,
between Finca Yalpemech and Chinaja, Alta Verapaz.

Calyptranthes cuneifolia Lundell, Am. Midi. Nat. 29: 478.
1943.

McVAUGH: FLORA OF GUATEMALA 299

British Honduras, the type from Toledo District, Monkey River,
in pine ridge along Jenkins Creek, P. H. Gentle 4096.

A tree up to 6 meters high, 15 cm. in diameter, glabrate, the branchlets terete
or compressed, the inflorescence and young growth thinly appressed-pubescent
with grayish- or yellowish-white dibrachiate hairs 0.1-0.3 mm. long (on vege-
tative buds up to 0.5 mm. long); leaves coriaceous, obovate, less often elliptic
or elliptic-ovate, 1.5-3 cm. wide, (2.5-) 3.5-6.5 cm. long, (1.5-) 2-2.5 (-3) times
as long as wide, narrowed to a blunt and often rounded tip (in the more broadly
obovate leaves broadly rounded from about the middle); base cuneate, the thick
revolute margins decurrent on the inner angles of the channeled petiole 2-3 mm.
long, 1 mm. thick; midvein nearly flat above, somewhat sulcate near the base;
lateral veins 8-12 pairs, more evident on the lower surface where little stronger
than the intermediate and connecting veins, the whole forming an irregular reticu-
lum; marginal vein about equaling the lateral ones, 1-2 mm. from the margins,
connecting the lateral veins by ill-defined and irregular arches; blades drying
pale gray-green, nearly concolorous, with small dark glands on both sides; in-
florescences appearing solitary and axillary, or at the lowest (bracteate) nodes
of leafy shoots, 2-5 cm. long; flowers 20-30 in each panicle, short-pedicellate
in small groups near the tips; peduncle 1.5-2.7 cm. long, 1-1.5 mm. thick; lateral
branches of the panicle 1-2 pairs, 1.5 cm. long or less; buds 2.5-3 mm. long, pyri-
form, substipitate, densely pale-strigose on the stipe and at the base, glabrous
above the middle, rounded or obscurely apiculate at the apex; calyptra 2-2.5
mm. wide; hypanthium after anthesis broadly campanulate, 2.5 mm. wide at the
mouth; style 3 mm. long, capitate; stamens 50-60; petals cuneate-obovate, 1-1.5
mm. long, mostly adherent to the calyptra and falling with it; immature fruit
globose, up to 8 mm. in diameter, surmounted by a collar 2 mm. wide, up to
1 mm. high.

Calyptranthes hondurensis Standl. Journ. Arnold Arb. 11: 36.
1930. Eugenia belizensis Standl. Field Mus. Bot. 11: 137. 1932.
C. belizensis (Standl.) Lundell, Phytologia 1: 218. 1937. C. Aguilarii
Standl. & Steyerm. Field Mus. Bot. 23: 128. 1944.

Forested ravines, especially in and along streams, often in pine
or oak forest, sometimes in dry situations, 1500 meters or less; Za-
capa; Chiquimula; Guatemala; Quiche" (without definite locality,
Aguilar 810, type of C. Aguilarii} ; El Progreso. British Honduras
(Mountain Pine Ridge, El Cayo District, H. H. Bartlett 11756, type
of E. belizensis) ; Honduras (the type from El Achote, near Siguate-
peque, Dept. of Comayagua, Standley 56164).

A shrub or small tree up to 4-5 meters high, at maturity glabrous or essen-
tially so, the young growth and the inflorescence rather loosely and sparsely
appressed-pubescent with flat longitudinally oriented yellow-brown or reddish-
brown dibrachiate hairs 0.2-0.3 mm. long, these sometimes densely aggregated
below the nodes of the panicle-branches, and about the base of the hypanthium;
branchlets 2-winged, the wings up to 0.5 mm. high, terminating distally between
the leaf-bases; leaves coriaceous, mostly narrow and blunt-tipped, elliptic-ovate

300 FIELDIANA: BOTANY, VOLUME 24

or -obovate, (0.5-) 1-1.8 (-2.6) cm. wide, 2.5-4.5 (-7.5) cm. long, variable in
width even on the same plant, on most plants 2-5 (-7) times as long as wide, on
others 1.5-3 times, on some 3.5-7 times; blades usually rounded at tip, some-
times long-tapering to a rounded point; base acute or somewhat rounded, the
margins rather abruptly decurrent on the inner angles of the broadly channeled
petiole 2-5 mm. long, 1-1.5 mm. thick; midvein impressed above at least in the
proximal half of the blade; lateral veins about 10-15 pairs, like the marginal
vein obscure and hardly apparent in mature leaves; leaves drying pale gray-
green, when fresh probably dark green above, paler beneath; glands inconspicuous
on both surfaces, the blades sometimes impressed-punctate above; inflorescences
axillary, the paired floriferous panicles 5-7.5 cm. long, 2 or 3 times compound,
each subtended by a deciduous bract 3-5 mm. long, an elongate leafy branchlet
sometimes developed beyond (between) them; peduncle up to 3-4.5 cm. long,
somewhat compressed, about 1 mm. wide, usually more than half as long as the
whole inflorescence; flowers (10-) 15-25 in each panicle, mostly in 3's at the tips
of the branches; principal branches of the panicle one pair, up to 1.5-2.5 cm.
long; buds plump, sometimes quite glabrous, 2.5-3.5 mm. long, broadly fusiform
to obovoid, distally rather broadly and abruptly rounded to the short broad
apiculum, acute or somewhat rounded at the base; calyptra firm-textured, 2.5-3
mm. wide; hypanthium after anthesis broadly campanulate with flaring margins,
3-3.5 mm. wide; style 5-7 mm. long; stamens about 100, up to 5 mm. long; petals
about 2, 1-1.5 mm. long, narrow, adhering to the inner surface of the calyptra
and falling with it; fruit first red, black at maturity, probably globose, 4-6 mm.
in diameter, surmounted by a prominent collar.

The plant described as Eugenia belizensis was distinguished as
a species on the basis of its "small and remarkably narrow leaves."
The leaves in the type collection are indeed narrow (4-7 times
as long as wide) but are not smaller than other specimens of C.
hondurensis. In the type of E. belizensis the blades are also more
acutely pointed and more prominently glandular-punctate above
than is usual in C. hondurensis. In view of the very considerable
variability in the foliage of individual plants of the latter species,
it does not seem that E. belizensis should be accorded specific
rank on the basis of foliage-characters alone. The type of C.
Aguilarii is almost entirely glabrous, including the flower-buds, but
does not seem to differ otherwise from C. hondurensis.

Calyptranthes Hylobates Standl. ex Amsh. Ann. Missouri Bot.
Gard. 45: 169. 1958.

Rain-forests in lowlands; to be expected in Guatemala. Mexico
(Chiapas, Escuintla, Matuda 16873); Nicaragua; Costa Rica; Pan-
ama (Prov. Bocas del Toro, Almirante, Cooper 366, type).

Tree 4-7 meters high, up to 10-12 cm. in diameter, glabrous except for a
few tiny reddish dibrachiate hairs 0.1 mm. long at the base of the hypanthium
and on new growth; branchlets 4-angled, corky-winged, 2 low wings or angles

McVAUGH: FLORA OF GUATEMALA 301

terminating distally in the median lines of the leaf-bases, the other pair more
distinctly winglike, terminating between the leaf -bases in straight or falcate linear
auricles 1.5-2.5 mm. long; leaves ovate or lance-ovate, 1.5-2.5 cm. wide, 4-8
cm. long, (2-) 2.5-3 times as long as wide, acuminate or caudate-acuminate,
rounded or somewhat cuneate at base, the margins decurrent on the inner angles
of the petioles 1.5-2.5 mm. long, 0.7 mm. thick; midvein impressed above in a
narrow concave channel near the base of the blade; lateral veins obscure, rather
close and parallel, about 10-20 pairs; marginal vein about 1 mm. from the margin;
blades darker and obscurely glandular above, somewhat or scarcely glandular
beneath; inflorescences paired from an abortive compressed axis 1-1.5 mm. long,
the filiform panicles 2-4.5 cm. long, the axis 0.25-0.3 mm. thick; flowers 3 (-5?),
on filiform pedicels 2-6 (-15) mm. long, one pair opposite and lateral at a node
about 2-3 cm. above the base of the branch, the third one terminal; buds obovoid,
2.5-3.5 mm. long including the narrow apiculum 0.5-0.7 mm. long and a conic
pseudostalk about the same length; calyptra deeply bowl-shaped, about 1.5 mm.
across the mouth and 0.8 mm. high exclusive of the apiculum; hypanthium about
1 mm. long, broadly campanulate with flaring and revolute margin; style 4 mm.
long; stamens 60-70, up to 4 mm. long; anthers about 0.3 mm. long; petals appar-
ently none; fruit globose, probably about 5 mm. in diameter, with prominent con-
vex glands.

Calyptranthes Karlingii Standl. Field Mus. Bot. 8: 29. 1930.

Moist forests, river-bank forests, near sea level; Pete"n; Izabal.
British Honduras, the type from Tower Hill Estate, J. S. Karling 39;
Mexico (Tabasco, Chiapas).

A small tree, up to 5 meters high, 20 cm. in diameter; branchlets somewhat
quadrangular, carinately angled below the median lines of the leaf-bases, and
with thin, definite winglike ridges up to 0.5 mm. high, terminating distally be-
tween the leaf-bases; inflorescence, and the young branchlets and foliage, thinly
pubescent with flat appressed pale rusty longitudinally oriented dibrachiate hairs
0.1-0.2 mm. long; leaves coriaceous, lanceolate or occasionally elliptic-lanceolate,
nearly sessile, 1.5-3.5 cm. wide, 5-9 (-12) cm. long, 2.5-3.5 (-5) times as long as
wide, acute at tip (varying to obtuse or to obscurely acuminate), broadly rounded
and subcordate at base, the margins abruptly decurrent on the stout adaxially
concave petioles 1-3 mm. long, 1-2.5 mm. thick; midvein shallowly or narrowly
sulcate above; lateral veins obscure above and somewhat more evident beneath,
often 20 or more pairs including some smaller ones; marginal vein about equaling
the lateral ones and slightly arched between them, 1-1.5 mm. from the margin;
glands usually apparent as minute dark dots above, or the upper leaf-surface
punctate, the lower surface sometimes glandular also; blades dark green and
lustrous above, paler beneath; panicles 3-7 cm. long, with about 30 flowers or
fewer, 2-3 times compound, usually appearing single (not paired), each arising
at the lowest node of a new leafy shoot and subtended by a leafy bract 10-12 mm.
long; peduncles 1.5-4 cm. long, like the panicle branches somewhat angled and
compressed; bracts of the panicle deciduous before full anthesis, glabrous, oblong-
boat-shaped, the smaller ones (those subtending the terminal clusters of 3 flowers
or the bracteole-like ones at the bases of individual flowers) 1.5-2 mm. long;
buds at maturity 3-4 mm. long, prominently gland-dotted, obovoid, or ellipsoid

302 FIELDIANA: BOTANY, VOLUME 24

and more or less equally contracted to the substipitate base and the prominently
apiculate tip, the base or the entire bud often pale scurfy and appressed-pubescent
as well; calyptra bowl-shaped, 2 mm. broad; hypanthium at anthesis turbinate
with somewhat revolute margins, not lobed or divided; style 5 mm. long; sta-
mens on the margin of the hypanthium, 60-75, up to 4 mm. long; petals 1 or 2,
narrow, 2 mm. long, adhering to the inner surface of the calyptra and falling with
it; fruit globose or oblate, often somewhat didymous or tridymous, 6-9 mm. in
diameter, tipped by a flaring collar about 1 mm. high, 2.5 mm. wide, the calyptra
sometimes present as well.

Calyptranthes Lindeniana Berg, Linnaea 29: 213. 1858. C.
fluviatilis Lundell, Bull. Torrey Club 64: 553. 1937.

Forests along streams, river banks, 500 meters or less; Alta Vera-
paz (Pittier 231, 299). British Honduras (El Cayo, H. H. Barllett
12949, type of C. fluviatilis) ; southern Mexico (Tabasco, Linden 620,
type).

A shrub or treelet 2-3 meters high, the branchlets terete or somewhat com-
pressed, soon glabrate; young branchlets and inflorescence softly tomentose with
pale reddish brown hairs, those of the panicles mostly much contorted, loosely
appressed and with two equal or strongly unequal branches; hairs of the young
leafy branchlets partly as above, partly of nearly straight, erect, simple or short-
branched hairs up to 2 mm. long; leaves narrowly elliptic, lanceolate or oblance-
olate, 1-2.3 cm. wide, (5.5-) 7-8 (-11) cm. long, 4.5-6 times as long as wide,
about equally narrowed to both ends, the tip slenderly acuminate (the narrow
acumen 1-2 cm. long), the base acute, the margins decurrent on the inner angles
of the channeled petiole 6-8 mm. long, 1.2 mm. thick; midvein sulcate above
at least near base; veins except the midvein obscure above, the lateral ones 15-20
pairs in addition to some intermediates, ascending, straight; marginal vein about
equaling the laterals and little arched between them, 0.5-1 mm. from the margin;
blades probably dark green above, nearly featureless when dry, the lower surface
paler with indistinct glands; inflorescence 3-7 cm. long, 3 times compound, the
panicles paired in the axils of terminal nodes of an old branchlet, or sometimes
basal and opposite on a leafy shoot from the same node and then subtended by
bracts up to 8 mm. long; peduncle 2-3.5 cm. long; flowers up to about 40, in twos
and threes at the tips; buds obovoid, 2.3-2.7 mm. long, rounded at the tip, usually
weakly apiculate; calyptra dome-shaped, 1.7-2 mm. wide; hypanthium funnel-
form or campanulate with revolute margins, hairy within at the base of the
style; style about 7 mm. long; stamens about 75; petals 1-2, narrow, 1 mm.
long, adherent to the calyptra and falling with it; fruit probably globose, ?black,
less than 1 cm. in diameter, surmounted by a collar.

A species not very distinct from Calyptranthes Chytraculia, and
resembling the latter closely in all morphological details, including
the distribution and quality of the pubescence. It is probable that
the two species are closely related. The very narrow leaves of
C. Lindeniana, however, are distinctive, and its flowers are some-
what larger than those of C. Chytraculia.

McVAUGH: FLORA OF GUATEMALA 303

Calyptranthes macrantha Standl. & Steyerm. Field Mus. Bot.
23: 127. 1944.

Known only from the type collection, Alta Verapaz, wet mixed
forest along banks of Rio Frio, about 8 km. below Tactic, 1400 me-
ters, Standley 90532.

A shrub with terete or compressed branchlets; branchlets, inflorescence,
and petioles brown-tomentose with coarse erect or matted simple or forked hairs
up to 1.3-1.5 mm. long; leaves elliptic-ovate, 10-14 cm. long, 5-6.5 cm. wide,
2-2.3 times as long as wide, obscurely acuminate (the very tip obtuse), rounded
at base, the margins somewhat cuneately decurrent on the inner angles of the
deeply channeled petiole 8-11 mm. long, 1.7-2.5 mm. thick; midvein deeply
and narrowly impressed above; lateral veins 15-20 on each side, slightly impressed
above when dry, straight and parallel, more prominent beneath; marginal vein
about equaling the lateral ones and arching between them, forming a well-defined
border 1.5-3 mm. from the margin; blades dark green, smooth and somewhat
impressed-punctate above, paler and sparingly gland-dotted beneath; inflores-
cence of paired panicles from an abortive axillary axis, the panicles 7-8 cm.
long, up to 4 cm. across, about twice compound, about 15-20-flowered, the
peduncle 3.5-4.5 cm. long, 3 mm. wide including the hairs; buds ovoid, sessile,
in groups of 2-4 at the tips of the panicle-branchlets, obtuse or apiculate, 4.5-6
mm. long, 3-4 mm. in diameter including the hairs, the conic apiculum 1 mm.
long when denuded; hypanthium prolonged 2.5 mm. beyond the ovary; style
not seen expanded; stamens about 150, the anthers up to more than 0.5 mm.
long; petals 1 (-2), minute, attached to the calyptra and falling with it; fruit
unknown.

Calyptranthes megistophylla Standl. Carnegie Inst. Wash.
Publ. 461:75. 1935.

Known only from the type collection, British Honduras, in forest,
Camp 32, British Honduras-Guatemala boundary, 810 meters,
W. A. Schipp 1265.

A tree 15 meters high, 30 cm. in diameter, the inflorescence and young herb-
age sparingly appressed-pubescent with flat minute reddish dibrachiate hairs 0.1-
0.2 mm. long; leaves in age glabrous or nearly so, very large for the genus, ovate-
lanceolate, up to 12 cm. wide and 35 cm. long, obscurely short-acuminate, rounded
at base, the margins cuneately decurrent on the petioles 8-13 mm. long, 3-4
mm. thick; midvein on the upper surface with a central concave channel separating
two low parallel convex ridges; lateral veins close and parallel, 30-40 or more
including some intermediate ones; marginal vein about equaling the lateral ones
and slightly arched between them, forming a prominent bordering line 2-5 mm.
from the margin; blades probably lustrous above, paler and dull beneath, some-
what gland-dotted beneath; inflorescence very large, the paired panicles in the
type probably arising from an abortive axillary axis; panicles 15-22 cm. long,
3 or 4 times compound, many-flowered, trichotomously branched in one plane
at the principal nodes, the internodes above these nodes strongly flattened at
right angles to the plane of the branchlets; peduncle up to 12.5 cm. long and
5 mm. wide; flowers in dense clusters of 6-12 at the tips of the branchlets; buds

304 FIELDIANA: BOTANY, VOLUME 24

glabrous except at the base, obovoid or almost globose, scarcely apiculate, 3-3.5
mm. long; calyptra about 3 mm. wide, membranous, gland-dotted; hypanthium
funnelform after anthesis, 2.5 mm. long; style about 5 mm. long; stamens about
100, up to 5 mm. long, the anthers 0.3 mm. long and wide; petals 1 or more,
elliptic, half as long as the width of the calyptra and falling with the calyptra;
fruit unknown.

Calyptranthes Millspaughii Urb. Symb. Antill. 7: 294. 1912.
C. euryphylla Standl. Contr. U. S. Nat. Herb. 23: 1034. 1924. Walk
Naked.

Along streams, near sea level. British Honduras, Stann Creek
Dist., Freshwater Creek, J. Kelly no. 1. Yucatan Peninsula of
Mexico (Cozumel Island, Millspaugh 1537, the type) ; Veracruz ( Nel-
son 421, the type of C. euryphylla).

A tree or shrub, the branchlets and inflorescences rather thinly pubescent
with longitudinally oriented closely appressed flat coppery dibrachiate hairs
0.2-0.4 mm. long; branchlets compressed, not 2-edged; leaves elliptic to elliptic-
ovate or -obovate, 2.5-4 cm. wide, 6-12 cm. long, 2.25-3 times as long as wide,
gradually acuminate at the tip, acute at base, the margins decurrent on the
inner angles of the concave-channeled petiole 4-7 mm. long; midvein concave
above on the nearly plane surface of the rather thick and opaque blade; lateral
veins 10-12 on each side in addition to some intermediate ones, inconspicuous,
scarcely convex in dried leaves even on the lower surface; marginal vein about as
strong as the lateral ones and arched between them, 2-3.5 mm. from the margin;
blades dark green and often impressed-punctate above, paler and drying brown
beneath and finely dark-dotted; inflorescence 4-6 cm. long, the paired panicles
mostly 3 times compound, 30-40-flowered, in the axils of terminal nodes of old
leafy branchlets, or occasionally basal and opposite on a leafy shoot from the
same node, the peduncles 1.5-3 cm. long; principal secondary branches of the
panicle much flattened, their lowest internodes often 6-8 mm. long, 1.5 mm.
wide; flowers mostly in clusters of 3-5 near the tips; buds obovoid, apiculate,
2.5-2.8 mm. long; calyptra dome-shaped, membranous, 2 mm. wide; funnel-
form hypanthium after opening 1.5 mm. long, 2 mm. wide at the mouth, glabrous
within; style 4-5 mm. long; stamens about 50, up to 4 mm. long, the anthers
0.4 mm. long; petals minute, adherent to the calyptra and falling with it.

This species is known only from the collections cited above.
Two are flowering specimens, that from British Honduras collected
in early March, the type in mid-February. The fruit is unknown.
Probably the whole plant is glabrous or nearly so by the time the
fruit matures.

Formerly C. Millspaughii was confused with C. Chytraculia, from
which it may be distinguished most readily by the distinctively
different pubescence.

Calyptranthes pallens Griseb. Veg. Kar. 67. 1857. C. Chytra-
culia 8 pauciflora Berg, Linnaea 27: 27. 1855.

McVAUGH: FLORA OF GUATEMALA 305

A tree or tree-like shrub 5-8 meters high; leaves coriacous, rather broadly
elliptic, or sometimes broadest a little above or below the middle, narrowed
about equally to both ends, 2-4 (-6) cm. wide, 3-7 (-10) cm. long, 2-3 (-3.5)
times as long as wide; margins of the blade decurrent at the base on the inner
angles of the channeled petiole; mid vein impressed above at least near the base;
lateral veins close and parallel, obscure on both surfaces, 10-20 pairs including
some poorly defined intermediate ones; marginal vein about equaling the lateral
ones and slightly arched between them, 0.5-1.5 mm. from the margin; leaves
smooth and dark green above; panicles usually bracteate and paired at the base
of an otherwise leafy shoot, sometimes axillary on ordinary leafy branches, the
individual panicles pyramidal, 3-4 times compound, the flowers in small clusters
near the tips, not appearing closely sessile; peduncles 2-5 cm. long; buds 2.5-3
mm. long including the narrow base, obovoid, rounded and often slightly apiculate;
calyptra domelike; hypanthium after opening campanulate, 2 mm. long; style
4-5 mm. long; stamens about 50-60, up to 3.5 mm. long; petals tiny, adhering
to the calyptra and falling with it; ripe fruit dark red, globose or oblate, 5-8
mm. in diameter.

A species primarily of the Caribbean basin, the nomenclaturally
typical variety occurring in the West Indies, southern Florida, the
Yucatan Peninsula, and the Isthmus of Tehuantepec, Mexico. The
two following varieties may be expected to occur in Guatemala.
For notes on the taxonomy of this species see Fieldiana, Bot. 29:
406-409. 1963.

Calyptranthes pallens Griseb., var. pallens.

Lowlands, to be expected in Pete*n and British Honduras. Mexico
(Yucatan; Isthmus of Tehuantepec); southern Florida; West Indies.

Vegetative branches prominently 2-winged, the wings terminating distally
between the leaf -bases; inflorescence, branchlets, and to some extent the foliage,
rather persistently appressed-pubescent with pale rusty-brown, yellowish brown
or, especially on the leaves, nearly colorless dibrachiate hairs 0.1-0.3 mm. long;
leaves prominently acuminate at tip, broadly cuneate at base, usually glandular-
punctate above; petioles 6-8 mm. long; panicles 10-12 cm. long or less, usually
with 50-60 or more flowers, the lowest branches often as long as the peduncles,
the latter 2-4 cm. long; calyptra 1.3-1.7 (-2) mm. across.

Calyptranthes pallens Griseb., var. mexicana (Lundell)
McVaugh, Fieldiana, Bot. 29: 409. 1963. C. mexicana Lundell,
Wrightia 2: 166. 1961.

Arroyos, brushy slopes, stream-sides, mountain forests, to be ex-
pected in Guatemala. Mexico (Chiapas to Sinaloa).

Vegetative branches winged or wingless, the wings sometimes apparent only
on a few of the young branchlets of individual plants; inflorescence and branchlets
very sparingly pubescent, the appressed and usually coppery hairs mostly 0.1-0.2

306 FIELDIANA: BOTANY, VOLUME 24

mm. long, and aggregated chiefly about the base of the hypanthium and the
distal nodes of the panicle; leaves acuminate at tip (sometimes obscurely so or
some leaves merely obtusely rounded), acute or sometimes cuneate at base, often
glandular-punctate above; petioles 2.5-5 mm. long; panicles 3-7.5 cm. long, 30-35-
flowered, the lowest branches shorter than the peduncles, the latter 2-4.5 cm. long;
calyptra 2-2.5 mm. wide.

Calyptranthes paxillata McVaugh, Fieldiana, Bot. 29: 410.
1963.

Known only from the type collection, Guatemala, Alta Verapaz,
wet forest above Tactic, above the bridge across Rio Frio, about
1400-1500 meters, Slandley 90457.

A small tree with nearly terete or compressed branchlets; branchlets, in-
florescence and young foliage covered with appressed nearly sessile, dibrachiate
and longitudinally oriented coppery hairs 0.5-1 mm. long; leaves glabrate, elliptic
or elliptic-lanceolate, 1.5-3.5 cm. wide, 4-7 cm. long, 2.3-3 times as long as wide,
caudate-acuminate, the slender tapering tip 1-1.5 cm. long, obtuse; base of blade
acute or cuneate, the margins decurrent on the inner angles of the petiole 3-6
mm. long, 0.8-1.2 mm. thick; midvein channeled above near base; lateral veins
obscure, close and parallel, about 15 on each side including some intermediate
ones; marginal vein about equaling the lateral ones, nearly straight between them,
1 mm. from the margin; inflorescence of paired panicles from an abortive axillary
axis or from the lowermost (bracteate) node of a leafy shoot; panicles 3-4.5 cm.
long, twice compound, bearing 9 flowers or fewer; peduncle up to 3.5 cm. long,
compressed, 1 mm. wide below the lowest branches; buds obo void-fusiform, 4.5-5
mm. long, 2-2.5 mm. in diameter, broadest above the middle, tapering to the
base, rather abruptly contracted to a blunt peglike apiculum 1-1.3 mm. long;
hypanthium after anthesis funnelform, 2.5-3 mm. high, the rim prolonged 2
mm. beyond the summit of the ovary; calyptra about 3 mm. across, 2.5 mm.
long including the apiculum; style 7-8 mm. long; stamens about 100, up to 4 mm.
long; petals 1-2, minute, remaining attached to the calyptra and falling with it;
fruit globose, glabrous, about 8 mm. in diameter, surmounted by the collar-
like base of the hypanthium 2.5 mm. wide.

Calyptranthes pendula Berg, Linnaea 27: 21. 1855.

Mountain slopes in wet forest, or on dry rocky hillsides, 900-
1900 meters; Escuintla; San Marcos; Quezaltenango; Sacatepe"quez.
Mexico (Oaxaca, Galeotti 2870, the type; Chiapas); El Salvador.

A tree or tree-like shrub 5-8 meters high, the vegetative branchlets slightly
compressed, not 2-edged; inflorescence and young branchlets, and to some extent
the foliage, rather sparingly appressed-pubescent with crisped pale rusty-brown
or nearly colorless dibrachiate hairs 0.1-0.3 mm. long, the hairs often densely
aggregated and coppery about the base of the hypanthium, paler and often
evanescent elsewhere in the inflorescence; leaves coriaceous, rather broadly elliptic
or elliptic-ovate, 3-4 cm. wide, 7-12 cm. long, 2-3 times as long as wide, gradually
but prominently acuminate at tip, rounded or sometimes merely acute at the base,

McVAUGH: FLORA OF GUATEMALA 307

the margins decurrent on the inner angles of the petiole 8-10 mm. long; mid-
vein impressed-concave above at least near base; lateral veins close and parallel,
10-20 on each side including some poorly defined intermediate ones, more evi-
dent beneath; marginal vein about equaling the lateral ones and arched between
them, 1.5-2.5 mm. from the margin; mature blades smooth and dark green and
without apparent glands above, paler beneath; panicles usually bracteate and
paired at the base of an otherwise leafy shoot, sometimes axillary on ordinary
leafy branches, the individual panicles pyramidal, 3-4 times compound, 5-10
cm. long, often 50-60-flowered, the flowers in small clusters near the tips; pe-
duncles 3-5 (-8) cm. long, the lowest secondary panicle branches shorter than this;
buds 2.5-3 mm. long, obovoid, rounded and slightly apiculate at tip; calyptra
membranous, domelike, 1.7-2 mm. across; hypanthium after opening campanulate,
2 mm. long; style 4-5 mm. long; stamens 50-60, up to 3.5 mm. long; petals tiny,
adhering to the calyptra and falling with it; fruit dark red or black, globose or
oblate, 5-8 mm. in diameter.

The taxonomic position of this species is somewhat doubtful.
It is similar in the characteristics of the inflorescence and especially
of the pubescence to C. pattens Griseb. From the Central American
and Mexican varieties of C. pollens, however, C. pendula may be
distinguished by its larger leaves, which are often rounded at base
and usually apparently eglandular above, by its longer petioles,
and by the absence of wing-like angles on the leafy branchlets.
From C. Chytraculia, flowering specimens of C. pendula may be
distinguished readily by the differences in abundance and quality
of the pubescence. Fruiting specimens, from which most of the
pubescence has disappeared, may be difficult to identify. As far
as known C. Chytraculia is confined to lowlands near the Atlantic
Coast, whereas C. pendula is a species of middle elevations. From
C. Millspaughii, a species of the lowlands of the Yucatan Peninsula,
C. pendula may be distinguished by the more slender inflorescence
and the usually somewhat crisped pubescence.

Calyptranthes perlaevigata Lundell, Contr. Univ. Mich. Herb.
7: 29. 1942.

Subdeciduous forest, river-banks, 500-1500 meters. To be ex-
pected in Guatemala. The type, Matuda 4387, came from near
Siltepec, Chiapas, less than 50 km. from the Guatemala border.

A tree 3-5 meters high, glabrous as far as known except for a few sessile
reddish dibrachiate hairs 0.2-0.3 mm. long, on the vegetative buds, the branches
of the panicle, and the base of the hypanthium; leaves elliptic, or sometimes
broadest above the middle, 3-6 cm. wide, 10-16 cm. long, 2-3 times as long as
wide, rounded distally to a tapering acumen up to 1-2 cm. long, rounded below
the middle to a broadly acute or cuneate base, the thick and somewhat revolute
margins cuneately decurrent on the inner angles of the ventrally concave petiole

308 FIELDIANA: BOTANY, VOLUME 24

5-9 mm. long, 1.5 mm. thick; midvein channeled above near base, for most of
its length with a shallow central channel separating low convex parallel ridges;
lateral veins close and parallel, 20 or more on each side including some inter-
mediate ones, ascending, inconspicuous but apparent on both surfaces; marginal
vein about equaling the lateral ones and arched between them, forming a firm
line nearly parallel to the margin and 1.5-3 mm. from it; blades lustrous and
abundantly but minutely impressed-punctate above, sparingly resinous-dotted
beneath; inflorescence relatively short and about 9-flowered, the paired panicles
(2-) 5-8 cm. long, arising from an abortive axillary axis or from the lowest (brac-
teate) node of a leafy branchlet; peduncles 3.5-6 cm. long, up to 1.7 mm. wide,
more than half as long as the whole panicle; lateral flower-bearing branchlets usually
one pair, 5 mm. long and 3-flowered, but the lateral flowers sometimes sessile
on the main axis; buds before anthesis broadly fusiform, obtuse, 7-8 mm. long;
calyptra conic, 4 mm. high, 4 mm. in diameter; hypanthium after anthesis cam-
panulate with flaring margins, 4 mm. long; style about 11 mm. long; stamens
about 100, up to 7 mm. long, the anthers 0.5-0.7 mm. long; petals not seen; calyptra
(persistent on the fruit) up to 4.5 mm. wide, obtusely broad-conic, thickened,
apparently fleshy, 2 mm. high; fruit globose or oblate, probably up to 1 cm. in
diameter, the persistent collar-like neck of the hypanthium up to 2 mm. high,
its rim 4 mm. wide.

Calyptranthes zanquinensis A. Molina, Ceiba 3: 168. 1953.

To be expected in Guatemala. Known only from Liquidambar
forest, Honduras, Dept. of Morazan, Montana Zanquin, Quebrada
de la Paz, 1600 m., A. Molina R. 2977, the type.

A tree 5 meters high, with compressed branchlets; inflorescence, branchlets
and petioles bristly-tomentose with very coarse hollow erect somewhat ascending
pale yellowish-brown forked hairs up to 0.3-0.7 mm. high; leaves elliptic to ovate-
lanceolate, (2-) 2.5-4 cm. wide, (7-) 9-12 cm. long, 2.6-3.4 times as long as
wide, gradually acuminate with tapering tip, rounded from the middle or below
toward an acute base, the margins finally somewhat cuneately decurrent on the
acute inner angles of the channeled petiole 5-7 mm. long; midvein sulcate above;
lateral veins 12-15 pairs including some intermediate ones, inconspicuous above,
evident beneath; marginal vein about equaling the lateral ones and slightly
arched between them, 1-2 mm. from the margin; leaves nearly concolorous,
lustrous and obscurely impressed-punctate above, the glands scarcely apparent
beneath; paired panicles from the lowermost (bracteate) nodes of a leafy axillary
branch, the bracts foliaceous, lanceolate, 18 mm. long, 4 mm. wide; panicle 10
cm. long, 2 or 3 times compound, the peduncle 4-6 cm. long, up to 2 mm. wide
at the apex; buds clustered near the tips, obovoid, about 5 mm. long; hypanthium
after anthesis campanulate, flaring at the mouth, 3-4 mm. long; style about
7 mm. long; stamens probably about 150, up to 4-6 mm. long, the anthers 0.4
mm. long; calyptra dome-shaped, 2.5 mm. wide, 2 mm. high including the slender
apiculum 1 mm. long; petals 2-3, minute, attached to the calyptra and falling
with it; fruits mostly solitary on short pedicel-like branchlets, apparently black,
nearly globose, about 8 mm. in diameter, surmounted by a prominent collar-
like rim 2.7-3.2 mm. wide.

McVAUGH: FLORA OF GUATEMALA 309

EUGENIA L.

References: Berg, Linnaea 27: 140-306. 1856; and in Mart. Fl.
Bras. 14, pt. 1: 214-326. 1857.

Calyx-lobes 4, distinct and imbricated in bud and in anthesis, and usually
persistent on the fruit (in some species loosely coherent below the tips until an-
thesis). Hypanthium little or not at all prolonged beyond the summit of the
ovary. Petals as many as the calyx-lobes, usually conspicuous. Ovary usually
bilocular, the ovules numerous (or rarely only 2) in each locule, usually in a sub-
capitate group attached near the middle of the central dissepiment. Seeds one
or two, the ovules as far as known undivided, pseudomonocotyledonous, the sep-
arate structures of cotyledon, radicle and plumule not discernible. Inflorescence
fundamentally a raceme, i.e. the individual flowers in decussate pairs; terminal
flower of the axis usually wanting; axis sometimes extremely shortened, the flowers
then appearing as if in axillary fascicles, umbels or glomerules; flowers, if occa-
sionally solitary, arising from the basal bracteate nodes of new branches that bear
leaves at the upper nodes, or from the bracteate nodes of abortive axillary axes;
bracteoles at base of flower often broad and persistent, sometimes connate. Fruit
a "berry," usually with thin pulp over the massive seed.

One of the very large genera of flowering plants; the American
species, according to an estimate made by Berg in 1859, number 537.
The actual number may be smaller than this; according to Bentham
the whole genus includes no more than about 500 species. According
to Merrill and Perry (Mem. Am. Acad. 18, pt. 3: 136. 1939) about
2500 species in the New and the Old Worlds have been described
in, or transferred to, Eugenia. The limits of the inclusive genus
Eugenia have been the subject of much controversy. Many modern
authors prefer to segregate the very numerous species of south-
eastern Asia as a distinct genus, Syzygium Gaertn. (see below).
American authors generally have been reluctant to recognize the
segregate genera proposed by Berg, e.g. Stenocalyx, Phyllocalyx,
Siphoneugena, Myrcianthes and Myrciaria. For the sake of con-
venience the few Old World Eugenias introduced into Guatemala
may be treated as forming a distinct section, distinguished as follows:

Hypanthium much prolonged beyond the summit of the ovary, forming a funnel-
form, campanulate or bowl-shaped "calyx-tube" bearing the stamens about
its margin; base of the hypanthium attenuate above the place of attachment
of the deciduous bracteoles into a narrow stipelike pseudostalk; inflorescence
terminal or if lateral then usually at leafless nodes on old wood; natives of
the Old World, cultivated or escaped Sect. Syzygium (p. 310).

Hypanthium not much prolonged beyond the summit of the ovary, the stamens
borne on a flat disk surrounding the base of the style and at about the same
level (the base of the style sometimes in a shallow concave depression) ; base
of the hypanthium usually not stipelike, the expanded portion of the hypan-
thium subtended (usually closely) by a pair of bracteoles, these mostly per-
sistent until well after anthesis; inflorescence always lateral, usually at leafy
nodes; native American species Sect. Eugenia (p. 315).

310 FIELDIANA: BOTANY, VOLUME 24

Sect. I. SYZYGIUM, sensu M. R. Henderson, Gard. Bull.
Singapore 12: 11. 1949.1

There has been no general agreement on the validity of Syzygium
as a genus distinct from Eugenia. Merrill and Perry, in studies
on the Myrtaceae of China and of Borneo, respectively (Journ.
Arnold Arb. 19: 191-247. 1938; and Mem. Am. Acad. 18: 135-202.
1939), argued for the separation of Syzygium on the basis of the
divided embryo (i.e. the distinct cotyledons) and the roughish seed
coat "loosely or closely adhering to the pericarp." In this stand
they have been followed by perhaps a majority of recent workers
on Old World Myrtaceae. In his comprehensive treatment of
Eugenia in Malaya, however, Henderson maintained after study of
various living species that neither the characteristics of the seed
coat nor those of the embryo can be used as a reliable basis for generic
distinction. Henderson thus proposed to regard Syzygium as a
section of Eugenia, distinguished from Eugenia proper chiefly by
having the "calyx-tube" (hypanthium) prolonged above the ovary,
with the stamens borne along the margin of the tube.

Recent studies on wood anatomy and on pollen morphology
suggest that there may indeed be some valid basis for the division
of the inclusive genus Eugenia and the recognition of Syzygium
(Ingle, H. G., & Dadswell, H. E., The Anatomy of the Timbers
of the South-West Pacific Area. III. Myrtaceae. Austral. Journ.
Bot. 1: 353-401. 1953; Pike, Kathleen M., Pollen Morphology of
Myrtaceae from the South-West Pacific Area. op. cit. 4: 13-53. 1956).

In Asia, Malaya and Australia there are numerous native species
of Syzygium. A few species have been introduced into America
for ornament. One species not treated below, S. samaragnense
(Blume) Merr. & Perry (Eugenia javanica Lam.), has been reported
from Panama.

Inflorescence compound (i.e. a panicle, its lower branches branched again, bearing
several or many flowers) ; panicle usually lateral at leafless nodes, sometimes
terminal; buds calyptrate, circumscissile, the petals falling as a unit and the
truncate margin of the hypanthium persisting as a tube or collar in flower
and in fruit E. Cumini.

Inflorescence a raceme, the branches ("pedicels") 1-flowered (occasionally in
E. myrtifolia 3-flowered); raceme terminal or lateral; calyx-lobes 4, persist-
ent and conspicuous in flower and in fruit.

1 This name was derived from the generic name Syzygium Gaertn., published
in 1788. It was published as a sectional name without Latin description and with-
out any specific reference to the original publication as a genus. It is used here
more or less in the sense of Henderson, merely as a convenient way of designating
the small group of Asiatic species that are cultivated or naturalized in Guatemala.
Nomenclaturally it is probably unacceptable.

McVAUGH: FLORA OF GUATEMALA 311

Flowers very large (buds 1.5-3 cm. long including the pseudostalk; style 2.5-

4 cm. long; calyx-lobes up to 1-1.5 cm. wide); fruit 2.5-6 cm. in diameter.

Flowers pink to red, nearly sessile in crowded lateral clusters. .E. malaccensis.

Flowers white, or somewhat greenish or yellowish; pedicels mostly 1-1.5 cm.

long; raceme usually terminal, not crowded E. Jambos.

Flowers smaller (buds 6-8.5 mm. long; style 8-10 mm. long; calyx-lobes 3 mm.
wide or less); fruit probably about 1 cm. in diameter or less; inflorescence
terminal, racemose, or the flowers on the most vigorous branches subum-
bellate in threes E. myrtifolia.

Eugenia Cumini (L.) Druce, Rep. Bot. Exch. Club Brit. Isles
3: 418. 1914. Myrtus Cumini L. Sp. PI. 471. 1753. E. jambolana
Lam. Encyc. 3: 198. 1789. Syzygium Cumini (L.) Skeels, U. S. Dept.
Agric. Bur. PI. Ind. Bull. 248: 25. 1912. Calyptranthes 0' Neillii
Lundell, Bull. Torrey Club 64: 554. 1937 (type from an old clearing
near Belize, British Honduras, H. 0' Neill 8764).

Cultivated grounds about towns and villages, spreading from
cultivation in many parts of the tropics. British Honduras; Suri-
nam; Brazil; widely distributed in the Indo-Malaysian region, in-
troduced in other tropical regions.

Tree 6-20 (-30) meters high, glabrous; leaves coriaceous, oblong-ovate to
elliptic-oblong, 3-8 cm. wide, 7-18 cm. long, 2.5-3 times as long as wide, the tip
shortly or abruptly acuminate, rarely obtuse; base acute to cuneate, the margins
decurrent on the channeled petiole 1-2.5 cm. long, 1-2 mm. thick; midvein broadly
sulcate above; lateral veins 25-40 pairs in addition to some intermediates, slightly
elevated on both surfaces when dry; marginal vein about equaling the laterals and
somewhat arched between them, 1-3 mm. from the margin; blades olive-green
and lustrous above, paler beneath, both surfaces often finely glandular-punctate;
inflorescence many-flowered, sometimes terminal, often lateral at leafless nodes
below the tips of the branches, cymose-paniculate, 3 or 4 times compound, 4-
12 cm. long with several pairs of distant horizontal branches, the peduncle 1-3 cm.
long, the flowers sessile in groups of 3 or more near the tips; bracteoles and upper
bracts similar, deciduous at or before anthesis, reddish brown, deltoid, acute,
0.5-1.5 mm. long; buds pyriform, 5-6 mm. long, rather abruptly contracted be-
low into a stout pseudostalk 1 (-2) mm. long; calyx truncate, 2.5-3 mm. across
the mouth, brownish-pink, the tube prolonged 1.5-2 mm. above the ovary, the
4 lobes very broad and low, obscure, quickly deciduous; style 5-7 mm. long; sta-
mens about 100, inserted on the margin of the calyx tube; filaments pinkish,
2-6 mm. long; petals falling in a flat orbicular calyptra ca. 2.5 mm. in diameter;
ovary bilocular, the ovules 12-15 in a compact group near the summit of each
locule; fruit black when ripe, edible, oblong to oblong-ellipsoid, asymmetric, 1.5-
2 cm. long, 1-1.5 cm. wide, umbilicate; seed 1-1.5 cm. long, the two cotyledons
unequal, not fused.

Eugenia Jambos L. Sp. PI. 470. 1753. Jambosa vulgaris DC.
Prodr. 3: 286. 1828. Syzygium Jambos (L.) Alston, in Trimen, Fl.

312 FIELDIANA: BOTANY, VOLUME 24

Ceylon (Suppl.) 6: 115. 1931. Manzana rosa; Pomarrosa; Manzana;
Manzanita; Ros (Coban, Quecchi).

Often abundantly naturalized in forest, pastures, and hedges,
especially in Alta Verapaz, Izabal, and along the Pacific bocacosta;
cultivated commonly at low and middle elevations almost every-
where in Guatemala; native of the Indo-Malaysian region (according
to Merrill & Perry, Journ. Arnold Arb. 19: 218. 1938), now pantropic
in cultivation and as an escape.

A shrub or small tree, glabrous; leaves lanceolate or elliptic-lanceolate, mostly
3-5 cm. wide, 12-20 cm. long, 4-5 times as long as wide, long-tapering to the
narrowly acuminate apex, subcuneate or gradually rounded at the base, the mar-
gins shortly decurrent on the stout petiole 1-2 mm. thick, about 1 cm. long;
mid vein impressed above, prominent beneath; lateral veins 10-15 pairs, slightly
elevated both sides but more prominent beneath; marginal vein about equaling
the laterals, very slightly arched between them, 2-5 mm. from the margin, with
a much fainter nerve very near the margin; upper surface drying pale or dark
green, the lower usually greenish brown; glands scarcely apparent on either sur-
face at maturity; inflorescence a terminal raceme up to 2.5 cm. long, the axis
4-angled, stout, 2-3 mm. thick, bearing 2-4 decussate pairs of flowers, the terminal
flower usually abortive; flowers large, 7-8 cm. across the stamens; pedicels up to
1.5 cm. long, 1-1.5 mm. thick (2 mm. in fruit); bracts and bracteoles fugacious;
hypanthium obconic, about 1.5 cm. long and 1 cm. across the mouth, gradually
narrowed to the base and slightly contracted into a stout, poorly defined pseudo-
stalk 2-3 mm. long;1 calyx-lobes 4, persistent, concave, reflexed after anthesis,
broadly rounded, the larger about 1 cm. wide at base, 6 mm. long; petals white,
orbicular, gland-dotted, about 1.5 cm. broad and long; stamens very numerous
(about 300) and conspicuous, up to 4 cm. long, creamy- white; anthers oblong,
1.2 mm. long; style 3.5-4 cm. long; fruit broadly depressed-globose, up to 6 cm.
in diameter, 3-4 cm. long, rose-scented, pale yellow with pink flush; seed 1, about
2.5 cm. in diameter.

The following notes were supplied by Mr. Standley: Every-
where in Central America the tree is called "Manzana rosa" or
"Pomarrosa." It is planted chiefly for ornament and shade, for
which it has few superiors in the tropics, the foliage being very
dense, dark green, and persistent through the dryest seasons. The
flowers are large and rather showy but scarcely pretty. The fruits
have a characteristic flavor of rose water, and remind one of toilet
soap. Their flavor is little superior to the taste of that article,
and the fruits are eaten principally by children, birds, and various
domestic animals. The seeds germinate readily, and probably are
dispersed by the animals that eat them, for the seedlings often

1 The majority of species of Syzygium have the hypanthium narrowed abruptly
or gradually to a slender pedicel-like base which is actually not a pedicel but part
of the flower. This is the pseudostalk (cf. Henderson, Card. Bull. Singapore 12:
12. 1949). When a pedicel is present, the articulation between it and the pseudo-
stalk is evident.

McVAUGH: FLORA OF GUATEMALA 313

are found in quantity remote from mature trees. The trees have
naturalized themselves thoroughly in the wetter parts of Guate-
mala, particularly about Coban, where they often look like native
trees, especially when scattered through open pine forest. The
trees frequently are planted as living fence posts. Sometimes the
plant is used for hedges, especially in the coffee regions of the
bocacosta. If the young plants are pruned severely, they become
very dense and form an effective if not particularly handsome
hedge. Very infrequently the fruit is stewed and served as a dessert,
as we have noted at Quezaltenango and elsewhere. It seldom is
offered for sale in the markets. The wood is described as reddish
gray, close-grained, and durable. The bark is reported to yield
a brown dye, and in some parts of the world is employed for tanning.

Eugenia malaccensis L. Sp. PI. 470. 1753. Syzygium malac-
cense (L.) Merr. & Perry, Jour. Arnold Arb. 19: 215. 1938.

Planted rarely, especially about Guatemala; doubtless also in
the Atlantic lowlands. British Honduras; El Salvador (where called
marandn japones) ; Panama; West Indies; tropical South America;
native of some part of the Indo-Malaysian region, now more or
less pantropic in cultivation.

A tree 8-20 meters high, glabrous; leaves drooping, coriaceous, usually large,
variable in shape and size, from oblong-oblanceolate to oblong-elliptic or some-
what obovate, up to 18 cm. wide, 35-36 cm. long, 1.7-2.7 times as long as wide,
the apex abruptly acuminate or obtuse, the base cuneate or somewhat rounded,
the margins decurrent on the stout, adaxially flattened or concave petiole 1-1.5 cm.
long, 2-3 (-5) mm. thick; midvein impressed above; lateral veins 10-15 pairs,
distinctly stronger than any intermediate ones, 1-2.5 cm. apart, diminishing dis-
tally, ascending and overarching 0.5-1.5 cm. from the margin and each ultimately
joining the next adjacent vein, a definite marginal vein usually not formed as such;
blades dark green and somewhat lustrous above; glands obscure on both surfaces;
inflorescence a stout axillary raceme formed at leafless nodes below the ends of
the branchlets (rarely at leafy nodes), the axis woody, 1-1.5 cm. long, 1.5-3 mm.
thick, bearing 2-5 opposite decussate pairs of conspicuously jointed nearly sessile
flowers (the pedicels 1-1.5 mm. thick, 1-2 (-8) mm. long); bracts and bracteoles
1-2 mm. long, deciduous before anthesis and leaving prominent narrow scars;
buds 1.5-3 cm. long; calyx obconic or funnelform, about 1 cm. across the mouth,
abruptly narrowed below into a stout pseudostalk 6-10 mm. long; lobes 4, unequal,
persistent, broadly rounded with thin pale margins, at maturity 6-10 (-15) mm.
wide, the larger ones 3-6 (-10) mm. long; styles 2.5-3.5 cm. long; stamens about
100; filaments 2-3 cm. long, red, flattened below; anthers 1 mm. long; petals 4,
orbicular-spatulate, pink to red, 1-1.5 cm. long; ovary bilocular, multiovulate;
fruit oblong or obovoid, 4-7.5 cm. long, 2.5-6 cm. in diameter, pinkish to dark red,
umbilicate; seed subglobose, 1.5-2 cm. in diameter, the cotyledons equal or un-
equal, not fused.

314 FIELDIANA: BOTANY, VOLUME 24

Mr. Standley notes that it is a handsome tree, in appearance
much like E. Jambos, but the flowers are rose-purple, and produced
on thick branches inside the foliage, so they are not visible except
when one is under the tree. The fruits are red, pyriform, about 5
cm. long, with an acid and highly agreeable flavor. When in flower
this is one of the most delightful of trees, the petals falling in vast
numbers and covering the ground with a lovely carpet of bright
and handsome color.

Eugenia myrtifolia Sims, Bot. Mag. t. 2230. 1821. ?E. rheedi-
oides Standl. & Steyerm. Field Mus. Bot. 23: 131. 1944.

Known in Guatemala from four collections only, all these without
special locality and without habitat-data. "Dept. Escuintla," in
1942, Aguilar 1754, 1757 (type of E. rheedioides); "Dept. Guate-
mala," in 1938-39, Aguilar 185, 351.

Tree or shrub, glabrous even to the vegetative buds; leaves elliptic and acute
at both ends, varying to oblanceolate or obovate and then often obtuse or rounded
at apex; apex often sharply acute or acuminate, or in obtuse leaves shortly cuspi-
date; margins long-de current on the channeled and usually thin-edged petioles
2-5 mm. long; blades 1-2.5 cm. wide, 2-7.5 cm. long, usually 2-3 times as long as
wide; midvein sulcate above; lateral veins indefinite, somewhat apparent beneath,
rather closely parallel, 10-12 on each side of the midvein; marginal vein about
equaling the laterals and sometimes arched between them, up to 1 mm. from the
margin; glandular dots scarcely apparent on the leaves; inflorescence terminal,
few-flowered, corymbosely cymose, the axis 1-5 cm. long beyond the last pair of
foliage leaves, the branches in 2-4 decussate pairs, naked at anthesis or rarely with
small leafy bracts, the flowers terminal and solitary or on the most vigorous
branches subumbellate in threes on pedicels 1-4 mm. long; buds 6-8.5 mm. long,
much wrinkled in drying, the hypanthium long-drawn-out beneath the calyx into
an attenuate pseudostalk; calyx-lobes 4, often unequal, rounded-triangular, con-
cave, wrinkled in drying, 1.5-2 mm. long in bud, up to 3 mm. in fruit; disk 3 mm.
in diameter, the deeply concave center 1.5 mm. in diameter; style 8-10 mm. long,
its base depressed as much as 2 mm. in the concave disk; stamens about 150, the
filaments 8-9 mm. long, the anthers 0.5-0.6 mm. long; petals glabrous, ragged-
edged, concave, up to 5 mm. long; ovary bilocular, the ovules 6-8 in each locule,
radiate in a subcapitate group affixed near the middle of the central dissepiment;
fruit not seen.

The type of E. rheedioides perhaps belongs to another species,
closely allied to the one represented by the other specimens cited
above. In the type, Aguilar 1757, the flower-buds are turbinate,
4-5 mm. long, acutely narrowed to the base but lacking the very
long attenuate base that marks the other flowering specimens. It
is quite clear that the specimens of E. rheedioides and the others
here referred to E. myrtifolia belong to Old World species recently

McVAUGH: FLORA OF GUATEMALA 315

introduced into Guatemala. No American Eugenia known to me
has the terminal corymose cyme tending to branch trichotomously.
The two flowering specimens of E. myrtifolia from Guatemala,
Aguilar 185 and 351, match precisely in every particular a specimen
of E. myrtifolia collected by Adams Jewett in the Jardin des Plantes,
Paris, Sep. 4, 1834, a little more than a decade after the introduction
of the species from "New Holland." They also appear to match
the original illustration in the Botanical Magazine; thus, although
I have not seen the type of E. myrtifolia, I am confident the name
is correctly used for the Guatemala plant. It seems likely that
this species has been introduced for ornament, as have so many
other Australian Myrtaceae throughout the tropics and subtropics.

Sect. II. EUGENIA.

All native American species as far as known belong to this section.
Numerous additional species, thus far unknown from within or near
the limits of Guatemala, occur in Mexico. Relatively few others
are known from Central America; the number of species known from
Guatemala far exceeds that known from any other part of Central
America.

Many Guatemalan species are still imperfectly known. Twelve
are thus far known from the types only; ten are known in fruit
only; and six are known only from flowering specimens. Because
of these gaps in our knowledge of these plants, the construction
of a satisfactory general key to species is well-nigh impossible. The
difficulties are increased by the vegetative uniformity that prevails
throughout this genus — and indeed throughout the Myrtaceae—
and by the fact that the majority of the species change considerably
in appearance, in pubescence and in other characteristics between
the time of flowering and that of fruiting.

The following key makes primary use of the various features
of the inflorescence, because the structures involved are relatively
long-persistent, and because they change little after attaining their
full growth. Pubescence-characters are similarly useful except that
in mature specimens only small remnants of the original extensive
covering may persist. In the interpretation of the key it should
be remembered that most of the data, including those relative to
such things as color, texture, wrinkling, and appearance of veins
and glands, have necessarily been taken from herbarium specimens.

The inflorescence in most American species is a simple leafless
raceme bearing a number of opposite decussate pairs of flowers

316 FIELDIANA: BOTANY, VOLUME 24

subtended by small bracts; the raceme in each species is distinctive
as to length and as to the number of nodes. In almost all species,
however, the raceme may upon occasion be extraordinarily elongated
and axillary branches (under the influence of as yet unknown stimuli)
may bear flowers in the axils of small bracts at the lower nodes,
and flowers in the axils of ordinary leaves at the upper nodes. Per-
sons using the key, therefore, will do well to remember that the same
species may appear in several places; an effort has been made to
take into account the variation in the most plastic species (e.g.
E. biflora, E. Oerstedeana) , but a key to all possible forms of all spe-
cies would be so cumbersome and equivocal as to defeat its own
purpose.

Flowers solitary, axillary, large (disk 5-6 mm. wide; calyx-lobes 4.5-5.5 mm. wide) ;
fruit 1.5 cm. long; plants tomentose; Honduras and Nicaragua.

E. hondurensis.

Flowers racemose, the axis of the raceme often very short and the flowers in
approximate pairs, sometimes appearing glomerate or fasciculate Key A.

A. Appressed-pubescent or tomentose species with relatively slender and elongate
racemes (axis of the racemes mostly 1-3 cm. long, the internodes much longer
than the diameter of the axis, and at least some of them 3-5 mm. long or
more).

Bracteoles thin and conspicuous, mostly glabrous or nearly so and contrasting
with the heavily pubescent hypanthium, persistent in flower and in fruit,
0.7-1.5 mm. long, usually connate along their proximal margins and form-
ing an involucre beneath the flower; plants with copious appressed pubes-
cence of pale sordid hairs.

Bracteoles acute or acuminate, broadly ovate-deltoid, 1-1.5 mm. long; bracts
thin, persistent, the upper ones usually longer, often 1-2 mm. long;
racemes usually elongate, the axis 1 cm. long or more, with well-separated
pairs of flowers; lateral veins close and parallel, 10-20 on each side of
the midvein.

Leaves lanceolate, 3-4 times as long as wide, bristle-tipped, impressed-
punctate above, the glandular depressions numerous and conspicuous;
lowlands, widely distributed on the Atlantic slope E. biflora.

Leaves elliptic to ovate or broadly lanceolate, not bristle-tipped (acuminate
but the tip obtuse or sometimes subacute), not at all or sparingly and
obscurely impressed-punctate above.

Leaves mostly 3-4 times as long as wide; eastern Mexico to Guatemala.

E. Karwinskyana.
Leaves mostly 1.7-2.5 times as long as wide; Yucatan Peninsula.

E. yucatanensis.

Bracteoles obtusely pointed or rounded, less than 1 mm. long; bracts ovate-
deltoid, somewhat indurated at base, less than 1 mm. long; axis of the
raceme usually 2-6 mm. long (rarely up to 1 cm.), the pairs of flowers
approximate; lateral veins 4-6 (-8) on each side E. argyrea.

Bracteoles not conspicuous, usually pubescent like the hypanthium and pedicel,
persistent (and then often somewhat thickened at base), or deciduous at
flowering time, not connate and involucre-like even if persistent; plants

McVAUGH: FLORA OF GUATEMALA 317

with closely appressed or loose pubescence or the hairs all forked or di-
brachiate.

Leaves narrowly elliptic-lanceolate, 1.5 cm. wide or less, 3.5-4.5 times as long
as wide; known in fruit only, the raceme-axis 1.5 cm. long, bearing 2-3
pairs of flowers; immature fruit oblong-ellipsoid, 12 mm. long; calyx-
lobes 2.5 mm. wide; disk 2-2.5 mm. wide E. sasoana.

Leaves broader, mostly 2-4 (-7) cm. wide, 1.5-3 times as long as wide.

Flowers small; disk in flower mostly 1.3-2 mm. wide; stamens 50-75;
larger calyx-lobes 1.5-2.5 mm. wide; axis of the raceme usually less
than 1 cm. long; leaves less than 8 cm. long.

Leaves glabrous beneath even at flowering time, elliptic, 2-2.5 times as
long as wide, 4-8 cm. long, on petioles 6-9 mm. long; axis of the
raceme often 6-10 mm. long; Pete"n E. Shookii.

Leaves appressed-pubescent with pale hairs, these often persisting on
one or both surfaces until the fruit matures; blades broadly elliptic
to ovate or obovate, 1.6-2 (-2.5) times as long as wide, on petioles
4-6 mm. long; axis of the raceme usually less than 5 mm. long;

Pacific lowlands, Quezaltenango to western Mexico E. Rekoi.

Flowers large; disk in flower 2-3 mm. wide or more; stamens 100-200;
larger calyx-lobes up to 2.5-4 mm. wide and long; axis of the raceme
1-2.5 (-8) cm. long, seldom shorter; leaves up to 16 cm. long, often
shorter, closely appressed-pubescent or tomentose beneath at least at
flowering time.
Axis of the raceme 2-5 (-8) cm. long; bracteoles deciduous at flowering

time or before.

Pubescence appressed, dull coppery red; leaves acute at the base, at
maturity glabrous or sparingly pubescent; disk in flower 2.7-
3.3 mm. wide; bracteoles 1.3-1.5 mm. long; lowlands near the
Atlantic coast E. aeruginea.

Pubescence of contorted and often matted hairs, yellowish white to
pale reddish brown; leaves rounded and often subauriculate at
base, usually permanently pubescent beneath; disk in flower (3-)
4-5 mm. wide; bracteoles 2.5-5 mm. long; middle elevations,

Guatemala to Mexico E. salamensis.

Axis of the raceme usually less than 2 cm. long; bracteoles persistent at
least through anthesis; pubescence closely appressed.

Leaves obovate or obovate-elliptic, rounded at apex, permanently sil-
very-scurfy beneath, the individual hairs so fine as to be scarcely
discernible; midvein flat or convex above; axis of the raceme

usually less than 1 cm. long E. hypargyrea.

Leaves mostly elliptic with acuminate tip, glabrescent, at maturity
glabrous beneath or with fine scattered pale hairs; midvein im-
pressed above; axis of the raceme usually 1-2 cm. long.
Leaves larger, those on vegetative branchlets 3-6.5 cm. wide, 7-
15 cm. long, the smaller leaves often 2.5-3 cm. wide, 6-7.5 cm.
long; blades usually broadest at the middle or below, somewhat
rounded to the base and gradually narrowed to the tip; Pete'n;

eastern Central America; Lesser Antilles E. octopleura.

Leaves smaller, those on vegetative branchlets 3-4 cm. wide, 6-
9 cm. long, the smaller leaves often 2-2.5 cm. wide, 4.5-6 cm.
long; blades mostly broadest near the middle, some of them
usually oblanceolate or obovate, with relatively long-drawn-out

318 FIELDIANA: BOTANY, VOLUME 24

base; mountains of Guatemala, 800-2000 meters, to south-
eastern Mexico E. guatemalensis.

A. Pubescence various or none; racemes short (usually much less than 1 cm.)
and the flowers approximate, or if the racemes elongate and slender then the
pubescence (if any) mostly of minute upstanding hairs.

Racemes slender and elongate, 1-2.5 (-4) cm. long; plants glabrous or nearly
so at maturity, the branchlets and petioles minutely hispidulous, the tender
growth bearing tiny appressed coppery dibrachiate hairs as well, a few of

these persisting near the base of the lower leaf -surf ace E. Oerstedeana.

Racemes short and crowded, if more than 1 cm. long then with numerous
approximate pairs of flowers; flowers often fasciculate or glomerate, some-
times solitary at the lowermost nodes of new leafy branchlets and on the
same plants in short racemes.

Cauliflorous species, the flowers long-pedicellate in clusters at usually leafless
nodes on old wood, the axis of the raceme almost none, or 1-3 mm. long;
coarse plants with relatively large leaves, flowers and fruits (leaves 7-
18 cm. long; disk 2.5-6 mm. wide; pedicels 8-30 mm. long; fruit 1.5-
2.5 cm. long).

Leaves cordate-auriculate at base; petioles (3-) 4.5-7 mm. long; inflores-
cence hirsutulous E. jutiapensis.

Leaves acute to cuneate at base, or somewhat rounded to a cuneate-decur-
rent base, never cordate-auriculate; inflorescence glabrous or sparingly
ciliate.

Young branchlets and herbage densely brown-tomentose; leaves dotted
beneath with abundant small convex resinous glands; mid vein im-
pressed above; bracteoles connate, forming a persistent involucre

2-2.2 mm. wide E. chiapensis.

Plants glabrous or essentially so as far as known; leaves not much dotted
beneath; midvein flat or concave above; bracteoles distinct but per-
sistent, 1-1.5 mm. long.

Lateral veins about 6-8 on each side, strongly ascending and anasto-
mosing, scarcely forming a marginal vein; leaf 7-8 cm. wide;
pedicels in fruit 13-30 mm. long; disk 6 mm. wide; larger calyx-
lobes 4 mm. long, 5-6 mm. wide in fruit (flowers unknown); fruit

1.5-2 cm. thick, 2-2.2 cm. long E. Matudae.

Lateral veins about 10-12, nearly straight, connected by a series of
arches forming a marginal vein 3-5 mm. from the margin; leaf
2.5-6 (-8) cm. wide; pedicels in fruit 8-12 mm. long; disk 3.5-
4 mm. wide; larger calyx-lobes 2.5 mm. long, 3-3.5 mm. wide;

fruit 1-1.5 cm. thick, 1.5-2.3 cm. long E. choapamensis.

Species not cauliflorous, the inflorescences usually in the axils of new leaves,
if occasionally at leafless nodes then the raceme-axis prolonged or the
pedicels very short, or both; plants and flowers various KeyB.

B. Inflorescence (at the time of anthesis) or developing leafy branchlets, or usu-
ally both, tomentose (the hairs matted or contorted) or appressed-pubescent;
minute erect hairs if present sparse, or often obscured by the appressed hairs;
some or all the hairs often forked or dibrachiate. [Plants often glabrous or
nearly so by the time the fruit matures, a few of the characteristic hairs usually
persisting about nodes and in the inflorescence.] (See second "B," p. 322.)

Flowers large and tomentose, mostly few in close axillary clusters [calyx-lobes
4-8 mm. long and wide; disk 3.5-5.5 mm. wide; axis of the raceme none or
scarcely apparent, 5 mm. long or less, 1-2 mm. thick; pedicels 0-3 (-5) mm.
long).

McVAUGH: FLORA OF GUATEMALA 319

Midvein of leaves convex above; known in fruit only, the fruit about 2 cm.
long; pedicels 2-2.5 mm. long and thick; disk 5.5 mm. wide; tomentum
pale brown; axis of raceme almost none E. toledinensis.

Midvein impressed above, often sharply and narrowly so; tomentum bright
red or coppery or silvery brown.

Leaves (3-) 4 times as long as wide, 9-20 cm. long, with caudate-acuminate
tip 1-2 cm. long; petioles 6-8 mm. long; bracteoles persistent, forming
a tomentulose involucre beneath the fruit; flowers unknown; fruit
nearly sessile, its disk 3.5-4.5 mm. wide; tomentum pale coppery to
"silvery brown" or "buff brown," persistent on the lower leaf-surface.

E. cervina.

Leaves 2-2.7 times as long as wide, 5-12 cm. long, broadly short-acuminate;
petioles 10-15 mm. long; bracteoles deciduous at anthesis, 2-5 mm.
long; tomentum bright red or coppery.

Flowers very large and almost sessile (disk 5 mm. wide; style 12 mm.
long; calyx-lobes 6-8 mm. long and wide); fruit 2-3 cm. long,
tomentose; surfaces of racemes and young growth densely tomen-
tose, the mat of contorted hairs up to 1 mm. thick . . E. pachychlamys.

Flowers moderately large, on pedicels 3-5 mm. long (disk 3.5-4 mm.
wide; style 8 mm. long; calyx-lobes 4-4.5 mm. long and wide); fruit
unknown; branchlets and inflorescence more loosely and thinly to-
mentose E.savannarum.

Flowers neither large nor tomentose, the hairs if abundant and only loosely
appressed then mostly longitudinally oriented on the branchlets and ra-
cemes; calyx-lobes at most 3.5-4 mm. long and wide, usually much less;
disk at most 3.5 mm. wide, usually much less; axis of the raceme often
well developed, the flowers then evidently racemose.

Hairs of the branchlets and pedicels mostly or partly appressed and dibrachi-
ate, one branch of the hairs often considerably longer than the other;
bracteoles and calyx-lobes usually pubescent to about the same extent
as the hypanthium.

Flowers glabrous or nearly so (the bracteoles and perianth-segments often
ciliate) ; young branchlets sparingly or rather densely pubescent with
small evanescent partly dibrachiate hairs, the plants as a whole soon
glabrate.

Pedicels (3-) 6-10 (-25) mm. long; flowers up to 5-6 pairs in slender
axillary racemes, or solitary in some axils; fruit 8-10 mm. in diam-
eter E. Oerstedeana.

Flowers sessile, as many as 50 or more in dense glomerules; fruit 2.7-
3.3 cm. in diameter, 3-3.5 cm. long E. citroides.

Flowers and pedicels appressed-pubescent, usually densely so, like the
young leafy branchlets and the rest of the inflorescence, with reddish,
sordid or yellow-white hairs; hairs more or less persistent, the plants
eventually glabrate.

Flowers small; disk in flower mostly 1.3-2 mm. wide; stamens 50-75
(-100); larger calyx-lobes 1.5-2.5 mm. wide; axis of the raceme
usually less than 1 cm. long; leaves 8 cm. long or less.
Leaves glabrous beneath even at flowering time; lowlands of the At-
lantic slope.

Leaves caudate-acuminate, the slender obtuse apex 7-12 mm. long,
or shorter in small leaves; pedicels and buds including the calyx

320 FIELDIANA: BOTANY, VOLUME 24

subtomentose, the pubescence copious; bracteoles 1-1.5 mm.
long; branchlets often hispidulous after the appressed hairs fall.

E. bumelioides.

Leaves abruptly or gradually acuminate, or obtuse or rounded, if
acuminate the tip shorter than above; calyx-lobes glabrous or
merely strigose except near the tips; bracteoles 1 mm. long or
less.

Flowers glomerate, the axis of the raceme usually 1-3 mm. long;
leaves often obtuse, sometimes acuminate; hairs pale, yellow-
white to pale reddish; hypanthium little elongated after an-
thesis, the fruit globose or nearly so E. laevis.

Flowers definitely racemose, the axis often 6-11 mm. long; leaves
obtusely but often slenderly acuminate; hairs coppery brown;
hypanthium soon elongate, the fruit oblong or ellipsoid.

E. Shookii.

Leaves appressed-pubescent beneath with pale hairs, these often per-
sisting on one or both surfaces until the fruit matures; axis of the
raceme usually less than 5 mm. long; Pacific lowlands, Quezalte-
nango to western Mexico E. Rekoi.

Flowers larger; disk in flower 2-3 mm. wide or more; stamens 100-200;
larger calyx-lobes up to 2.5-4 mm. wide; axis of the raceme often
1-2 cm. long; leaves up to 16 cm. long, often shorter, closely ap-
pressed-pubescent or tomentose beneath at least at flowering time
[E. hypargyrea, E. guatemalensis, E. octopleura]. (See key above,
p. 317).

Hairs of the branchlets and pedicels simple, appressed or sometimes in part
contorted or spreading-ascending; hypanthium often heavily pubescent
and often contrasting with the more nearly glabrous bracteoles and
calyx-lobes.

Bracteoles acute or acuminate, thin and conspicuous, 1-1.5 mm. long,
broadly ovate-deltoid, glabrous or nearly so and contrasting with the
heavily pubescent, canescent hypanthium, mostly connate along the
proximal margins and forming an involucre beneath the pedicellate
flower; bracts persistent, the upper ones thinner and elongate, 1-2.5
mm. long.

Mid vein concave or broadly sulcate above, glabrous or at first with a
few long hairs; leaves elliptic or ovate, about twice as long as wide,
obtusely acuminate; axis of the raceme very short, usually less than
5 mm. long; Pete"n, British Honduras E. Trikii.

Mid vein sharply and narrowly impressed above, the furrow usually per-
sistently hispidulous; axis of some or all the racemes usually 1 cm.
long or more [E. biflora, E. Karwinskyana, E. yucatanensis]. (See
key above, p. 316.)

Bracteoles obtuse or rounded, mostly inconspicuous and less than 1 mm.
long; hypanthium (except in two species) not canescent or strongly
colored by the pubescence; bracts mostly 1 mm. long or less; flowers
approximate, the internodes of the rachis usually less than 1.5 mm.
long; axis of the raceme usually 3-6 mm. long or less.

Hypanthium densely appressed-pubescent, whitened or colored by the
pubescence.

Flowers pedicellate; inflorescence and young growth densely clothed
with closely appressed lustrous coppery or golden silky hairs;

McVAUGH: FLORA OF GUATEMALA 321

bracteoles and calyx-lobes nearly glabrous, contrasting with the
pubescent hypanthium E. argyrea.

Flowers sessile; inflorescence and young growth rather sparingly ap-
pressed-pubescent with stiff pale hairs; calyx-lobes silky-strigose,
scarcely less pubescent than the hypanthium . . .E. amatenangensis.

Hypanthium sparingly pubescent or glabrous, the bracteoles and calyx
similarly pubescent; inflorescence and young growth with some ap-
pressed, ascending or flexuous hairs, often also with numerous small
erect hairs. [Here are to be found notably pubescent individuals
of species that will ordinarily be located in Key C on p. 324.]

Flowers sessile and glomerate, 2-3 pairs, the raceme-axis 1 mm. long;
bracteoles broad and membranous, suborbicular or broadly del-
toid-ovate, connate and persistent; pubescence of short appressed
reddish hairs; Peten E. vesca.

Flowers evidently pedicellate, the pedicels sometimes only 1-2 mm.
long; axis of the raceme usually 3-6 mm. long.

Young branchlets and inflorescence more or less persistently hispid-
ulous, sometimes also with some appressed or silky hairs in the
inflorescence; lowlands, chiefly of the Atlantic slope.

Silky hairs in the inflorescence and on the flowers abundant, up
to 0.5 mm. long, yellowish-brown or coppery, soon falling,
the branchlets strongly and permanently hispidulous; fruit
7-8 mm. in diameter, usually broadly short-stipitate; bracte-
oles distinct, persistent and hairy beneath the fruit, 0.5-0.8
mm. long E. vacana.

Flowers glabrous or somewhat hispidulous; inflorescence hispid-
ulous, glabrous, or with some short appressed hairs on the
raceme; fruit as far as known not stipitate; bracteoles distinct
or connate, glabrous or minutely hispidulous.

Bracteoles persistent but not connate, broadly deltoid-ovate,
0.5 mm. long; racemes and branchlets rather coarsely se-
tose-pubescent, the sordid hairs 0.2-0.5 mm. long; racemes
well developed, the axis often 5-12 mm. long with 4-8
well-separated pairs of flowers; leaves when dry opaque,
the veins scarcely apparent on the upper surface, the mid-
veins sulcate E. flavoviridis.

Bracteoles usually united by the proximal margins and forming
an involucre beneath the flower; plants mostly glabrous or
minutely hispidulous; axis of the raceme 3-6 mm. long, or
if longer then the flowers more numerous, and approximate.

Leaves ovate or rhombic-ovate to elliptic, 3-6.5 cm. long,
opaque, the veins scarcely evident above in dried leaves;
petioles ventrally flat or concave, dilated into the base
of the blade; pedicels in flower 3 mm. long or less; buds
1.5-2.7 mm. long; disk 1 mm. wide; style 3^4.5 mm. long;
fruit 7-8 mm. in diameter; plants usually glabrous.

E. axillaris.

Leaves elliptic or obovate, occasionally ovate, 6-12 cm. long,
veiny, the lateral and marginal veins pale above in dried
leaves and contrasting with the darker mesophyll; peti-
oles deeply channeled, usually with an abrupt transition
into the blade; midvein often flat and wrinkled near the

322 FIELDIANA: BOTANY, VOLUME 24

base, often somewhat depressed below the surface of the
blade, for most of its length often narrowly sulcate be-
tween two low parallel convex ridges; pedicels (at least
the lower) 5-13 mm. long; buds 2.5-4 mm. long; disk
1.3-2 mm. wide; style (5-) 7-8 mm. long; fruit 1 cm. or
more in diameter; plants often hispidulous.

E. acapulcensis.

Young branchlets and inflorescence not hispidulous, but more or
less appressed-pubescent with coarse coppery hairs up to 0.5
mm. long or more, the whole plant soon glabrate; leaves ellip-
tic, acuminate; pedicels 1.5-2 mm. long; mountains of Chiapas.

E. nigrita.

B. Plants glabrous, hispidulous, hirsute or setose, the hairs mostly erect, some-
times very short; appressed hairs, if present, comparatively few; dibrachiate
(appressed) hairs present in 4 species [included in this part of the key because
mature plants may be quite glabrous, or merely hispidulous].

Plants nearly glabrous, or hispidulous, the young growth bearing small coppery
appressed dibrachiate hairs, a few of these hairs persisting on the lower
leaf -surf ace and the midvein near the base; at least some of the flowers in
slender elongated racemes 1-2.5 (-4) cm. long; bracteoles broad, mem-
branous, persistent, up to 1.5 mm. long and wide E. Oerstedeana.

Plants various; dibrachiate hairs if present conspicuous and abundant on the
pedicels at least at flowering time.

Plants hairy or setose, the erect stiff hairs often 0.5-1 mm. long or more,
densely and permanently covering the branchlets, petioles, racemes and
at least the veins of the lower leaf-surfaces.

Leaves large (blades mostly 7-19 cm. long, petioles 7-15 mm. long, 1.5-
2.5 mm. thick including the hairs); fruit 1-2 cm. in diameter; larger
calyx-lobes 2.5-4 mm. wide; flowers sessile or nearly so, glomerate.

Midvein flat or convex above; blades 12-19 cm. long; lateral veins arcu-
ate-ascending, 8-10 on each side, scarcely forming a marginal vein,
each recurving 5-7 mm. from the margin to join the next succeeding
one; flowers unknown; immature fruit 1.3 cm. long, its disk 2.5 mm.
wide E. lancetillae.

Midvein concave and somewhat impressed above; lateral veins not
strongly arcuate, the marginal vein well developed, about as strong
as the lateral ones and arched between them, 2-4 mm. from the
margin.

Leaves 2-4 (-6) cm. wide, acute or cuneate at base; plants densely
tawny-hirsutulous; bracteoles oblong, 1.5-1.7 mm. long, 0.7 mm.
wide; fruit 1.3-1.5 cm. long E. Yunckeri.

Leaves 6-8 cm. wide, rounded at base, the margins abruptly decurrent
on the sparingly hispid petiole; plants soon glabrate, known only
in fruit; bracteoles broadly rounded, 1 mm. long, 2 mm. wide;
fruit 2 cm. long or more E. papalensis.

Leaves small (blades 10 cm. long or less; petioles 4 mm. long or less); fruit
5-8 mm. in diameter; larger calyx-lobes 1-2 mm. long and wide.

Plants uniformly and coarsely but not very densely setose, the reddish
or tawny hairs in no way obscuring the surface of the branchlets or
the lower leaf-surface; setae on the midvein near the base of the
lower leaf -surf ace 1-1.3 mm. long; cilia of the calyx-lobes 0.5-0.7
mm. long; flowers sessile, glomerate; leaves oblong-elliptic, abruptly

McVAUGH: FLORA OF GUATEMALA 323

rounded to a subcordate base, the margins seeming to meet across
the flat summit of the petiole E. chinajensis.

Plants finely and often densely soft-hirsutulous, the pale hairs often very
numerous and covering the branchlets and principal veins and peti-
oles; hairs of the midvein near the base of the lower leaf -surf ace
0.5 (-0.8) mm. long; cilia of the calyx-lobes mostly 0.2-0.4 mm.
long or less; leaves (rarely subcordate in E. origanoides) cuneate
to acute or somewhat rounded at base.

Flowers closely glomerate, the racemes several, clustered, setose or
hairy, the pedicels and the axis of the raceme usually concealed
by the flowers; calyx-lobes 1-2 mm. long and wide; style 5.5-7 mm.
long; leaves mostly elliptic, 4-10 cm. long, the midvein impressed
above E. origanoides.

Flowers evidently racemose, the slender raceme-axis 1-6 mm. long,
bearing 2-5 pairs of flowers on pedicels 1-2 mm. long; racemes
puberulent or minutely hispidulous, the hairs and flowers not
concealing the racemes; calyx-lobes 1 mm. long and wide, or less;
leaves 2.5-5.5 cm. long, the midvein flat or convex above.

Leaves elliptic, bluntly acuminate; marginal vein evident; flowers
and mature fruit unknown E. rufidula.

Leaves obovate or elliptic, mostly rounded at apex; lateral veins
arcuate-ascending, a well-defined marginal vein not formed;
style 3.5 mm. long; fruit 5 mm. in diameter E. buxifolia.

Plants glabrous, puberulent or minutely hispidulous, the hairs 0.1-0.3 mm.
long, often incurved. [Bracts and flower-parts sometimes long-ciliate,
or branchlets with slightly longer hairs, but then the leaves glabrous or
nearly so.]

Hypanthium prominently 8-ridged, the ridges usually apparent at anthesis,
becoming broad and fleshy in fruit, sometimes as high as the thick-
ness of the body of the fruit.

Flowers sessile or essentially so, glomerate in the axils of leaves; hypan-
thium from the very earliest stages nearly globose and strongly
ridged, abruptly expanded above a narrow neck into the large erect
subfoliaceous calyx; mature calyx-lobes up to 4 mm. long and wide;
disk 3.5 mm. wide; flowers and mature fruit unknown; native species.

E. cacuminum.

Flowers on slender pedicels (5-) 10-18 mm. long, these usually subtended
by thin scarious or colored bracts 3-5 mm. long; hypanthium at first
campanulate, the ridges soon apparent at least just beneath the
calyx; calyx-lobes elongate, strongly reflexed at anthesis; disk 2 mm.
wide; cultivated, native of South America E. uniflora.

Hypanthium not prominently ridged, usually campanulate; fruit not
ridged.

Flowers sessile and glomerate; pedicels up to 1-1.5 mm. long in fruit;
axis of the raceme or racemes hardly developed, or 1-3 mm. long,
stout, hidden among the flowers and scarcely apparent; flowers gla-
brous, or some parts ciliate.

Leaves small and prevailingly obovate, coriaceous, 3-6 cm. long, blunt-
tipped or rounded at apex; midvein flat to convex above; petioles
2-4 mm. long; fruit 6-10 mm. in diameter; plants glabrous; Peten
and British Honduras. . . .E. Ibarrae.

324 FIELDIANA: BOTANY, VOLUME 24

Leaves larger and prevailingly elliptic or ovate, sometimes obovate,
thinner, mostly 6-13 cm. long, acuminate or subacuminate or
sometimes obtuse; mid vein channeled or narrowly impressed
above; petioles 4-10 mm. long; young shoots tomentose or
strigose.

Pet£n; fruit 5-7 mm. in diameter; young shoots probably reddish-
strigose with simple hairs; midvein narrowly channeled, often
bordered by parallel convex ridges; bracteoles broad and mem-
branous, connate, 1-1.3 mm. wide E. vesca.

Wet mountains above 2000 meters, Chiapas and Guatemala; fruit
2.7-3.3 cm. in diameter; young shoots tomentose, some of the
hairs dibrachiate; midvein narrowly impressed above; bracte-
oles ovate, pointed, scarcely connate, 0.7 mm. wide.

E. citroides.

Flowers definitely pedicellate (the pedicels sometimes only 1-2 mm. long
at flowering time); axis of the raceme usually slender and evident,

but sometimes very short Key C.

C. Flowers large [larger calyx-lobes 2.5-3.5 mm. wide; disk (1.5-) 2-3.5 mm.
wide]; axis of the raceme not much elongating (1-4 mm. long) ; pedicels except
in one species 6-20 mm. long.

Pedicels short and stout (1-2 mm. long, at anthesis 0.5 mm. thick or more);
leaf-blades 8-13 cm. long, the petioles 8-10 mm. long; inflorescence, includ-
ing the flowers, minutely puberulent E. Percivalii.

Pedicels usually 6-10 mm. long or more; inflorescence glabrous except that the

bracts and perianth-parts may be ciliate.

Leaves rounded-obovate to oblong-elliptic, 4-10 cm. long, broadly rounded
or obtuse at apex; petioles 2-4 mm. long, up to 2.5 mm. thick; midvein
prominently convex above, often nearly 1 mm. wide near base.

E. Winzerlingii.

Leaves ovate to elliptic-lanceolate, 2-6 cm. long, narrowly but obtusely
pointed or acuminate; petioles 1.5-7 mm. long, 0.7-1.2 mm. thick; mid-
vein shallowly sulcate at least toward the base of the blade, or flat, never

broad and convex E. rhombea.

C. Flowers small [larger calyx-lobes 2 mm. wide or less; disk 1-2 mm. wide];
axis of the raceme often slender and evident, 3-5 (-10) mm. long; pedicels
mostly 5 mm. long or less, occasionally longer; young branchlets or inflores-
cences, or both, often puberulent or hispidulous.

Pedicels very slender, (-5) 10-18 mm. long, usually subtended by thin scarious
or colored bracts 3-5 mm. long; bracteoles thin, deciduous before anthesis;
flowers few (1-6), solitary or seeming to arise from an involucre of bracts;

plants glabrous; developing ovary becoming 8-ridged E. uniflora.

Pedicels (at least most of them) 5 mm. long or less, subtended by indurated or

merely thin-tipped bracts 0.5-1 (-1.5) mm. long; bracteoles persistent,

usually until the fruit falls, often partly connate by the proximal margins;

branchlets and inflorescence usually hispidulous.

Plants glabrous (at least not hispidulous or puberulent, sometimes with a few

long or appressed hairs, the bracts and calyx-lobes often ciliate).
Bracteoles more or less connate and involucre-like beneath the flower;
lowlands mostly of the Atlantic slope [E. acapulcensis, E. axillaris}.
(See key above, p. 321.)

Bracteoles persistent, but distinct on the two sides of the hypanthium,
their proximal margins not or barely contiguous; mountains of Chiapas
and central Guatemala, one species in Tabasco.

McVAUGH: FLORA OF GUATEMALA 325

Midvein sharply impressed; leaves rather broadly elliptic or elliptic-
ovate, 2-2.3 times as long as wide; young herbage and inflorescence
with scattered coppery hairs; bracteoles obtusely pointed, divari-
cate, 1 mm. long E. nigrita.

Midvein flat or convex above, or shallowly channeled between low par-
allel ridges; leaves narrowly elliptic to lance-ovate, 2.5-4 times as
long as wide; plants essentially glabrous; bracteoles rounded or
truncate, 0.5 mm. long.

Leaf-bases mostly cuneate, the lowest 1-2 pairs of lateral veins leaving
the midvein at a narrow angle and ascending more or less parallel
to the margin; racemes proliferous, the flowers often 2-3 in the
axil of a bract E. capulioides.

Leaf-bases rounded to acute, the lowermost lateral veins divergent,
not noticeably more strongly ascending than the others; bracts
each subtending a single flower E. balancanensis.

Plants hispidulous or puberulent, the branchlets, petioles, inflorescences and
often leaves bearing numerous erect, ascending or incurved hairs 0.1-
0.3 mm. long.

Bracteoles connate and involucre-like beneath the flower; leaves 5-10 cm.
long, 2-6 cm. wide, thick and coriaceous but veiny, the lateral veins
in dried leaves usually pale and convex above; midvein usually de-
pressed beneath the general surface of the blade, itself sometimes
nearly flat, or sulcate, or even convex; axis of the raceme often 6-10
mm. long or even more.

Leaves drying yellow-green or yellow; midvein convex near the middle
of the blade on the upper surface, varying especially toward the base
of the blade to flat or even sulcate; hypanthium densely and very
minutely pubescent with pale appressed hairs smaller than those of
the pedicels, closely invested and partly covered by the bracteoles;
calyx-lobes finely puberulent on both sides; fruit transversely oval,
7-8 mm. wide in the longer axis, 5-6 mm. long E. farameoides.

Leaves drying brownish red, reddish, green or brownish green; midvein
usually sulcate between narrow parallel ridges, often flat and some-
what wrinkled near the base in dried leaves; hypanthium usually
glabrous, sometimes hispidulous like the pedicels, mostly or wholly
exserted beyond the bracteoles; calyx-lobes usually glabrous; fruit
globose, 1 cm. or more in diameter E. acapulcensis.

Bracteoles persistent, but distinct on the two sides of the hypanthium,
their proximal margins not or scarcely contiguous Key D.

D. Leaves flat in drying, the midvein not depressed beneath the general level of
the blade, the vein itself flat or slightly elevated on the upper surface, some-
times shallowly channeled especially near the base; leaves often opaque and
relatively featureless above, the veins then scarcely or not at all apparent.

Leaves mostly rounded at apex, obovate to elliptic, widest at or above the

middle, cuneate at base, the petioles 1.5-3 mm. long E. buxifolia.

Leaves acute or acuminate, elliptic or ovate to lanceolate, widest at or below

the middle.

Upper leaf-surface uniformly puberulent even in age, with minute curved
hairs 0.1 mm. long; petioles 2-2.5 mm. long; axis of the raceme usually

well developed, often 8-12 mm. long E. riograndis.

Upper leaf-surface glabrous at maturity, or pubescent along the midvein, at
most sparingly puberulent elsewhere.

326 FIELDIANA: BOTANY, VOLUME 24

Midvein (known in immature leaves only) channeled in the nearly plane
surface of the blade, the narrow channel bordered by low parallel
ridges; buds 3.5 mm. long, the hypanthium substipitate; mountains of
Chiapas E. ovandensis.

Midvein flat, or somewhat convex or elevated above; buds 2-2.5 mm. long,
the hypanthium closely invested by the bracteoles; low elevations,
mostly on the Atlantic slope.

Fruit (7-) 9-12 mm. in diameter; fruiting pedicels 0.4-0.6 mm. thick;
flowers usually evidently racemose, the axis of the longer racemes on
each plant 6-10 (-25) mm. long; calyx-lobes appearing about as
long as wide, the larger ones about 1.5 mm. wide E. tikalana.

Fruit 4-6 mm. in diameter; fruiting pedicels 0.3 mm. thick; flowers
usually in nearly sessile sub-umbellate clusters, the axis of the longer
racemes on each plant 4-5 mm. long or less; calyx-lobes often appear-
ing shorter than wide, the larger ones 1 mm. wide or less . .E. Capuli.

D. Leaves in drying downwardly curved toward the midline, the midvein de-
pressed below the general level of the upper surface of the blade, often very
narrow and appearing as if in a V-shaped furrow; lateral veins various.

Midvein (known in immature leaves only) channeled in the nearly plane surface
of the blade, the narrow channel bordered by low parallel ridges of vascular
tissue; buds 3.5 mm. long, like the pedicels finely hispidulous, the hypan-
thium substipitate; mountains of Chiapas E. ovandensis.

Midvein impressed in a narrow V-shaped furrow, or the furrow more open and
the vein apparent as a concave and often pale line; parallel vein-like ridges
not developed along the midvein, the laminar tissue descending smoothly
toward the furrow; hypanthium as far as known not stipitate; buds, if as
much as 3.5 mm. long, appressed-pubescent like the pedicels.

Leaves narrowly elliptic, bluntly acuminate, acute or cuneate at base, 2 cm.
wide or less, 2.5-3.5 times as long as wide; axis of the racemes (known
in flower only) 4-9 mm. long; inflorescence minutely puberulent; pedicels
1-1.5 mm. long; mountains of Chiapas E. letreroana.

Leaves usually broader, (1-) 2-5 cm. wide, 1.5-3 times as long as wide;
racemes and branchlets hispidulous or appressed-pubescent; lowlands,
Yucatan to eastern Guatemala and Honduras.

Leaves mostly of an oblong-elliptic type, 6-9 cm. long, acuminate, rather
rigidly coriaceous, drying very smooth above and often yellowish green,
concolorous, or darker beneath; midvein rather broadly sulcate; ra-
cemes in young fruit (flowers and mature fruits unknown) well devel-
oped and rather loosely flowered, the sparingly and rather coarsely
setose-pubescent axis mostly 5-12 mm. long; pedicels 1.5-4 mm. long,
similarly pubescent E. flavoviridis.

Leaves ovate to elliptic or obovate, varying to lanceolate or oblanceolate,
not especially smooth above when dry, the surface microscopically
roughened by very numerous small pale convex dots much smaller
than the foliar glands; blades often much darkening above in drying,
paler beneath; midvein usually narrowly impressed in a V-shaped fur-
row; raceme-axis 3-5 mm. long or less; pedicels and racemes appressed-
pubescent or loosely pilose at flowering time.

Branchlets at maturity smooth or essentially so, when young appressed-
pubescent, the pubescence including numerous dibrachiate hairs;
pedicels at flowering time densely appressed-pubescent with dibrachi-
ate hairs.

McVAUGH: FLORA OF GUATEMALA 327

Leaves caudate-acuminate, the slender obtuse apex 7-12 mm. long, or
shorter in some small leaves, the blades rather broadly elliptic-
ovate or obovate; calyx-lobes in flower densely pubescent on both
sides; fruit 10-12 mm. in diameter, sessile E. bumelioides.

Leaves if acuminate shortly and broadly so, often obtusely pointed or
rounded at the tip; blades elliptic to lance-ovate, or in Guatemala
mostly oblanceolate to obovate; calyx-lobes in flower rather
densely pubescent near the tips, otherwise merely strigose without,
and glabrous within; fruit 5-8 mm. in diameter, in one variety
stipitate E. laevis.

Branchlets rather persistently hispidulous with small erect stiff hairs,
when young shaggy-pubescent also with some long weak ascending
hairs, the hairs all or nearly all simple, including the contorted silky
ones of the pedicels and hypanthium; leaves elliptic, obscurely or
shortly and broadly acuminate, the tip seldom narrow and elongate;
calyx-lobes glabrous within; fruit 7-8 mm. in diameter, often broadly
short-stipitate E. vacana.

Eugenia acapulcensis Steud. Nom. ed. 2. 1: 601. 1840. Myrtus
maritima HBK. Nov. Gen. & Sp. 6: 146 [folio ed. 116]. 1823. Eu-
genia niaritima (HBK.) DC. Prodr. 3: 282. 1828, not E. maritima
DC. I.e. 271. ?E. Bonplandiana Berg, Linnaea 27: 228. 1856 (type
from Colombia, Humboldt & Bonpland). E. mosquitensis Berg, Lin-
naea 31 : 257. 1862 (type from Pearl Key Lagoon, Nicaragua, Wull-
schlagel). E. deUoidea Standl. Contr. U. S. Nat. Herb. 23: 1045. 1924
(type from Guerrero, Nelson 2292). E. antiquae Riley, Kew Bull.
1927: 121. 1927 (type from Panama, Riley 142). E. escuintlensis
Lundell, Phytologia 2: 4. 1941 (type from Chiapas, Matuda 4144).
E. ovatifolia Lundell, Bull. Torrey Club 69: 396. 1 Mai 1942 (type
from British Honduras, P. H. Gentle 30). E. Bartlettiana Lundell,
Contr. Univ. Mich. Herb. 7: 30. Mai 1942 (type from British Hon-
duras, Gentle 47). E. bracteolosa Lundell, I.e. 31 (type from British
Honduras, C. L. Lundell 4316). E. campechiana Lundell, I.e. 32
(type from Campeche, E. Matuda 3832). E. sibunensis Lundell,
I.e. 34 (type from British Honduras, Gentle 1619). E. comitanensis
Lundell, Wrightia 3: 12. 1961 (type from Chiapas, E. Matuda 5753).

Moist or wet thickets or forest, mostly in the Atlantic lowlands,
300 meters or less; Pete"n; Escuintla; Retalhuleu. Mexico (Tabasco
and Chiapas to Sinaloa, the type from Acapulco, Bonpland; Yucatan
Peninsula (Campeche, British Honduras); Honduras; Nicaragua;
Panama; northern Colombia; West Indies.

A tree mostly 5-10 meters high, up to 30 cm. in diameter, either quite gla-
brous, or the branchlets, petioles and inflorescence (usually) minutely hispidulous,
the bracts and the raceme-axis also thickly pubescent with crisped small golden
hairs; leaves generally elliptic, varying to obovate or rarely to ovate, (2-) 3-4

328 FIELDIANA: BOTANY, VOLUME 24

(-6) cm. wide, (6-) 8-12 cm. long, 2-3 (-3.5) times as long as wide, abruptly
short-acuminate or sometimes slenderly acuminate, or some or all the leaves ob-
tuse; blades acute or cuneate at base, or sometimes rounded in the broader leaves
on the same plants, the margins decurrent on the inner angles of the rather deeply
channeled petioles 5-8 (-12) mm. long, up to 1-1.5 (-2) mm. thick; midvein often
rather flat and somewhat wrinkled near base in drying, often somewhat depressed
below the surface of the blade and near the middle of the blade or below usually
narrowly sulcate between two convex ridges; lateral veins 6-10 on each side,
straight, passing directly into an arched marginal vein of about the same thick-
ness, or individual veins arching over and joining the next succeeding ones but
scarcely diminished distally; marginal veins often well separated (1-5 mm.) from
the margin, with small submarginal veins intervening; lateral and marginal veins
convex on both surfaces, pale above and contrasting with the darker mesophyll;
blades darker and somewhat lustrous above, paler and often drying brown beneath;
glands often apparent on both surfaces, or seen merely as dark dots beneath; in-
florescences racemose, stout and short, proliferous, the racemes usually 2 or 3
superposed in each axil, occasionally compound, or the lower nodes of individual
racemes bearing 2-3 flowers each; racemes 3-6 (-17) mm. long, crowded, bearing
3-6 (-15) opposite or subopposite, decussate pairs of flowers; axis of the raceme
angled and dorsi-ventrally compressed, 1.5-2 mm. wide at base, rapidly diminish-
ing distally; lower pedicels 5-9 (-13) mm. long, the upper (or sometimes all) 3-
5 mm. long; bracts scarious, persistent, ovate, acute or obtuse, up to 1.4 mm. long;
bracteoles variable in size from plant to plant, ciliate but otherwise usually gla-
brous, 0.5-1.5 mm. long and wide, connate by the proximal margins into a per-
sistent cuplike involucre partially concealing the usually glabrous cup-shaped
hypanthium; surface of the hypanthium, if minutely puberulent like the pedicel,
not obscured by the tiny erect hairs; buds pyriform, 2.5-4 mm. long, the globe
of the petals 2-2.5 mm. in diameter; calyx-lobes broadly rounded, usually glabrous,
1-1.5 mm. long and wide; disk 1.3-2 mm. wide, nearly glabrous or the staminal
ring hairy; stamens about (60-) 100, up to 7 mm. long; style (5-) 7-8 mm. long;
petals ciliate, conspicuously dotted with large dark glands evident even in the bud,
3-4 mm. long; ovary bilocular, the ovules 6-15 in a centrally attached sub-capitate
group in each locule; fruit globose, black, somewhat fleshy, 1 (-2?) cm. in diameter.

For notes on the taxonomy and nomenclature of this variable
species, see Fieldiana, Bot. 29: 421-425. 1963.

FIG. 50. Inflorescences in Myrtaceae. A, Racemose type, Eugenia acapul-
censis (primarily from Matuda 3194); X 4. B, Racemose type, E. axillaris (Steere
2640, type of E. cozumelensis) ; X 4. C, D, Forked branching at the terminal node,
Calyptranthes Karlingii (MaludaSlll) ; X 2. In Calyptranthes the smaller branches
are often forked and divaricate, two axillary branches developing at the terminal
node but the terminal bud aborting. The "paired panicles" (fig. 49, E) are ordi-
narily produced at the lowermost node of such an axillary branch, which may
produce leaves at the upper nodes. In D, the terminal bud and one axillary branch
have failed to develop; the left-hand axillary branch now falsely appears to be
terminal, having pushed aside in its growth the undeveloped remnants of the buds.

329

330 FIELDIANA: BOTANY, VOLUME 24

Eugenia aeruginea DC. Prodr. 3: 283. 1828. E.Fadyenii Krug
& Urb. ex Urb. Bot. Jahrb. 19: 622. 1895.

Wet mixed forest, often in mangrove swamps or along banks of
streams near the coast, at or little above sea level; Pete"n; Izabal
(Puerto Barrios, W. R. Hatch & C. L. Wilson). Campeche; British
Honduras; El Salvador; Jamaica and Cuba.

A tree up to 10-15 meters in height and to 60 cm. in diameter, glabrate, the
inflorescence and young growth appressed-pubescent with coppery dibrachiate
hairs (persistent hairs on the petioles and leaf-surfaces mostly white or whitish) ;
leaves at maturity sparingly pubescent or glabrous; blades elliptic or obovate,
usually narrowed about equally to both ends, 2.5-4.5 (-6) cm. wide, 5-9 (-14) cm.
long, 1.8-2.3 times as long as wide, bluntly short-acuminate and decurved at the
apex, the basal margins straight, forming a right or acute angle, often cuneately
decurrent on the channeled petiole up to 1.5 mm. in diameter, 8-16 mm. long;
midvein impressed above; principal lateral veins 8-12 pairs in addition to various
intermediate ones, ascending, slender, evident on both surfaces when dry, some
of the small connecting veins often joining at right angles; marginal vein about
equaling the laterals and somewhat arched between them, 1-3 mm. from the
margin; upper surface smooth, the glands usually not apparent; lower surface
finely gland-dotted; inflorescence an axillary raceme 3-5 (-8) cm. long with 3-5
opposite (or subopposite) and decussate pairs of flowers; terminal flower some-
times present; lowermost node (in the leaf -axil) often bearing a second short
raceme in addition to the primary one; pedicels at anthesis about 0.4 mm. in
diameter, up to 10 (-20) mm. long, the upper ones shorter; fruiting pedicels up to
1.5 mm. in diameter; bracts oblong to linear, scarious, 1.5-2 mm. long, mostly
deciduous before anthesis; bracteoles distinct, narrowly deltoid, 1.3-1.5 mm. long,
deciduous soon after anthesis; buds about 4.5 mm. long, the conic hypanthium
1-1.5 mm. long, abruptly expanded above into the relatively large calyx; calyx-
lobes in 2 unequal pairs, appressed-pubescent on both sides, the outer pair ovate,
the inner quadrate-rounded and larger, up to 2.5 mm. wide, 2.5-3.5 mm. long,
both pairs persistently concave in anthesis but loosely spreading or recurved from
the base; hypanthium soon elongating and becoming ellipsoid-oblong with a dis-
tinct neck below the persistent and somewhat accrescent calyx; disk 2.7-3.3 mm.
wide, often somewhat quadrate, the staminal ring short-bristly, the concave center
with a few long hairs; style 7-8.5 mm. long, sparingly pilose in the lower half; sta-
mens about 125, 6-8 mm. long, the anthers 0.5-0.7 mm. long; petals white, ca.
5-6 mm. long and wide; ovary bilocular, the ovules 9-12 in each locule, in a sub-
capitate group arising from the central dissepiment; mature fruit "red, size of a
cherry" (Schipp), ellipsoid (1.5cm. long, 1 cm. thick) to oblong (0.8cm. thick, 1.6
cm. long), sometimes curved, or enlarged distally, crowned by the persistent calyx.

For a note on the nomenclature of this species see Sandwith,
N. Y., Kew Bull. 1934: 124-126. 1934.

Eugenia amatenangensis Lundell, Wrightia 3: 10. 1961.

To be sought in Guatemala. Known only from the type, col-
lected on a river side, Amatenango del Valle, Chiapas, 1835 m.,
E. Matuda 5832.

McVAUGH: FLORA OF GUATEMALA 331

A tree 3 meters high, glabrescent, the branchlets and young leaves and the
inflorescence more or less densely appressed-pubescent with coarse pale gray or
sordid hairs 0.2-0.4 mm. long; leaves rigidly coriaceous, elliptic, 2.5-3.5 cm. wide,
4.5-7 cm. long, about 2-2.3 times as long as wide, short-acuminate, cuneate at
base, the margins decurrent on the inner angles of the channeled petiole 5-8 mm.
long; mid vein acutely but rather broadly impressed above; lateral veins about 8
on each side, inconspicuous above, more evident beneath, connected by a series of
irregular arches 1-2 mm. from the margin, the marginal vein as such not well
developed; leaves green and glossy above, gland-dotted and apparently paler be-
neath; flowers sessile and glomerate in the axils, the raceme-axis (sometimes 2
superposed) stout, 1-3 mm. long, bearing 3-5 pairs of flowers; bracts 1 mm. long
or more, their smooth indurate persistent bases rounded, about 1 mm. long and
wide; bracteoles broadly ovate, obtuse, persistent but not connate, dorsally some-
what silky-strigose, about 1 mm. long and wide; buds 2.5 mm. long; hypanthium
campanulate, 1 mm. long or a little more, densely pale-silky-strigose; calyx-lobes
rounded, glabrous within, up to 1.3 mm. long and wide, the outer surface silky-
strigose about as densely as the hypanthium; disk glabrous, 1.3 mm. wide; style
5-6 mm. long; stamens probably about 50, about as long as the style, the anthers
0.5 mm. long; petals about 3 mm. long; ovary bilocular, the ovules 5-8 in a com-
pact group in each locule; fruit (slightly immature) red, oblong-ellipsoid, about
6 mm. in diameter, 10 mm. long.

Eugenia argyrea Lundell, Wrightia 3: 10. 1961.

Thickets, hillsides, forest, dry stream beds, old pastures, mostly
in lowlands of the Atlantic slope, but up to 1200 meters or more;
Izabal; Alta Verapaz. Mexico [Veracruz to Chiapas (near Siltepec,
E. Matuda 5074, the type)]; Honduras; Nicaragua; Costa Rica; Pan-
ama; Colombia (Santa Marta, H. H. Smith 907).

A shrub 2-3 meters high, or a tree up to 6-15 meters, the inflorescence and the
youngest branchlets and leaves densely clothed with lustrous appressed (antrorse)
simple coppery or golden hairs up to 0.5 mm. long; older leafy branchlets, and
leaves (at least the upper surface), glabrous at flowering time; leaves elliptic-ovate,
or elliptic, often broadly so, 1.5-3 (-5) cm. wide, 3-6 (-10) cm. long, 1.6-2.5 times
as long as wide, rather gradually deltoid-acuminate or narrowly acuminate,
somewhat rounded or usually cuneate at base, the margins decurrent on the
inner angles of the channeled petiole 4-7 (-10) mm. long; midvein sulcate above
in the otherwise plane surface of the blade; lateral veins obscure, 4-6 (-8) on each
side, scarcely apparent above in dried leaves, more prominent beneath, ascending;
marginal vein about as strong as the lateral ones or a little weaker, sometimes
ill-defined, 1-2.5 mm. from the margin; blades nearly concolorous, apparently dull
and nearly featureless above, sometimes sparingly appressed-pubescent beneath
at maturity but opaque, the glands apparent only in young leaves; inflorescence
an axillary raceme, 2-3 usually superposed, the axis 2-6 (-10) mm. long, bearing
up to 6 (rarely -9) pairs of flowers on pedicels 2-7 mm. long, up to 0.5 mm. thick
in fruit; bracts dark, somewhat indurated at base, ovate-deltoid, less than 1 mm.
long; bracteoles glabrous or nearly so, glandular-verrucose, broadly ovate, con-
nate and forming a boat-shaped involucre 1.3-1.5 mm. wide, 0.5 mm. high; buds

332 FIELDIANA: BOTANY, VOLUME 24

2.5-3.5 mm. long; hypanthium campanulate, 1 mm. long, densely tawny pubescent
with appressed glistening hairs; calyx-lobes in unequal pairs, rounded, nearly gla-
brous, the outer pair about 1.3 mm. wide, nearly 1 mm. long, the inner pair up
to 2 mm. wide and 1 mm. long; disk 1-1.4 mm. wide, hairy around the base of the
style; style 5-6.5 mm. long, hairy in the proximal third; stamens about 50, up to
5 mm. long, the anthers 0.6 mm. long; petals greenish white; ovary bilocular, the
ovules 8-10 in each locule; fruit obovoid or short-ellipsoid, black, often substipi-
tate, 5-8 mm. in diameter, 6-10 mm. long.

Eugenia axillaris (Sw.) Willd. Sp. PL 2: 970. 1800. Myrtus
axillaris Sw. Prodr. 78. 1788. E. guttata Lundell, Wrightia 2: 207.
Sep. 1961. E. cozumelensis Lundell, Wrightia 3: 13. Dec. 1961.

Moist or wet forests or thickets, at low elevations; Pete"n; Santa
Rosa. Veracruz; Yucatan Peninsula of Mexico; British Honduras;
Honduras; southern Florida; West Indies, the type from Jamaica.

A small tree to 7 meters high, 15 cm. in diameter, glabrous except for appressed
small coppery hairs on the vegetative buds and similar hairs on the margins of the
bracts, bracteoles and calyx-lobes; leaves coriaceous, ovate or rhombic-ovate,
sometimes elliptic, 1.5-3.5 cm. wide, 3-6.5 cm. long, 1.7-2.2 times as long as wide,
bluntly acuminate or merely obtusely pointed, rather abruptly narrowed to the
acute or cuneate base, the thick cartilaginous margins decurrent on the splayed
summit of the flat or concave petiole 5-8 mm. long, up to 1.5 mm. thick near base;
mid vein sulcate above; lateral veins 5-8 pairs in addition to some intermediates,
inconspicuous but in dried leaves apparent as fine convex lines on both surfaces;
marginal vein about equaling the laterals and strongly arched between them, well
separated (1-3 mm.) from the margin; leaves smooth (lustrous?) above, dull and
usually drying brown beneath; glands evident beneath, or in leaves dried at an-
thesis often apparent as convexities on both surfaces; inflorescence a short axillary
raceme (often with 1-2 additional superposed shorter racemes or with an extra
flower at some nodes of the axis), the crowded axis somewhat quadrangular, 3-4
(-7) mm. long, with (3-) 5-7 opposite decussate pairs of flowers on pedicels up
to 3 mm. long; bracts ovate or broader, incurved, scarious, subpersistent, about
0.6 mm. long and 0.8 mm. wide; bracteoles usually united by their proximal mar-
gins, forming beneath the hypanthium a persistent compressed cup 0.4-0.5 mm.
high, 0.7-1 mm. long; buds 1.5-2.7 mm. long, the hypanthium cup-shaped; calyx-
lobes in 2 unequal pairs, broadly rounded, 0.7-1.2 mm. broad, 0.5-0.7 (-1) mm.
long, concave; disk about 1 mm. wide, glabrous; style 3-4.5 mm. long; stamens
about 40, up to 3-4 mm. long, the anthers 0.4 mm. long; petals 2-3 mm. long,
white; ovary bilocular, the ovules 7-10 in each locule, radiating in a subcapitate
group from a centrally affixed placenta; fruit blue-black, globose or oblate, 7-8
mm. in diameter.

Perhaps as many as a score of species, and hundreds of specimens,
from Mexico and Central America, have in the past been misidenti-
fied as Eugenia axillaris, and so cited in various publications. The
true axillaris is primarily West Indian in distribution, but appears
to be well scattered also in the Yucatan Peninsula as far south as

McVAUGH: FLORA OF GUATEMALA 333

Pete"n. Records of axillaris from other localities in Mexico and Cen-
tral America should be regarded with caution.

Most West Indian specimens of axillaris are quite glabrous ex-
cept that some floral parts are ciliate. Some specimens that are
otherwise indistinguishable, however, especially in Florida and in
Central America, but also rarely in the West Indies, are bristly-
pubescent on the young growth and in the inflorescence. The taxo-
nomic significance of this is unknown. The type of Eugenia cozu-
melensis (from Cozumel Island, Steer e 2640) is pubescent but would
otherwise be referred to E. axillaris. The original specimens of
E. guttata (Pete"n, Tikal National Park, E. Contreras 25, the type,
and Contreras 338) are glabrous, and are exactly like West Indian
specimens of axillaris as I understand that species.

Eugenia balancanensis Lundell, Phytologia 1: 481. 1941.

To be sought in Guatemala. Known only from the type collec-
tion, from advanced forest near Balancan, Tabasco, E. Matuda 3361.

Tree 4 meters high, quite glabrous except for a very few stiff appressed hairs
on the youngest vegetative growth, and the finely ciliate margins of the bracts,
bracteoles and calyx-lobes; leaves lance-ovate, 1.5-3.5 cm. wide, 4-9.5 cm. long,
2.5-3 times as long as wide, gradually and bluntly but rather slenderly acuminate
at tip, rounded to acute at base, the margins at the very base cuneately decurrent
on the inner angles of the channeled petiole 3-6 mm. long; midvein in mature
leaves convex above when dry; lateral veins 7-10 on each side, inconspicuous
above, more evident beneath; marginal vein about as strong as the lateral ones
and arched between them, 1.5-3 mm. from the margin; blades green and appar-
ently dull above, paler beneath and minutely dark-dotted (glands evident in
young leaves but not at maturity) ; inflorescence an axillary raceme (or 2 racemes
superposed), the axis 4-10 mm. long, bearing 3-5 opposite or subopposite and de-
cussate pairs of flowers on pedicels 2-5 mm. long; bracts 0.5-0.8 mm. long, strigose
on the backs, deciduous at anthesis except the glabrous indurated base; bracteoles
0.5 mm. long, broadly ovate, obtuse, persistent, not connate; buds and flowers
not seen; hypanthium just after anthesis broadly campanulate, rounded at base,
0.7-1 mm. long; calyx-lobes thin, rounded, longer than wide or suborbicular, the
larger ones 1.5 mm. long; disk about 1.5 mm. wide; style not seen; stamens about
60; ovary bilocular, the ovules 5-6 in a compact group in each locule; fruit un-
known.

Eugenia biflora (L.) DC. Prodr. 3: 276. 1828. Myrtus biflora
L. Syst. Nat. ed. 10. 1056. 1759.

Wet mixed forest, in British Honduras sometimes growing in
mangrove swamps, at or near sea level; Izabal. Mexico (Chiapas);
British Honduras; West Indies; Costa Rica; Panama; lowlands of
northern South America, to eastern Peru and Bolivia.

334 FIELDIANA: BOTANY, VOLUME 24

Shrub or small tree up to 6-8 meters high, pubescent, often with conspicuous
pale appressed silvery-silky pubescence on the inflorescence and young growth,
the hairs up to 1 mm. long; leaves usually lanceolate, varying to ovate or to very
narrowly lance-linear, (0.5-) 1-1.5 (-3) cm. wide, 5-8 (-10) cm. long, 3-4 times
as long as wide (twice as long as wide in ovate-leaved forms, and up to 15 times
in very narrow-leaved forms) ; blades flat and silky or glabrous beneath, the upper
surface with numerous impressed glands, occasionally glabrous, usually sparingly
beset with white cobwebby hairs up to 1 mm. long; blades prominently long-
acuminate, almost caudate, the acumen often slender and curved, normally prom-
inently mucronate; base of blade rounded or cuneate; petiole channeled above;
stout, 2.5-4 mm. long; midvein impressed above, prominently raised beneath,
lateral veins very slender, 12-20 pairs, often not apparent in dried material or
evident on the lower surface only; marginal vein about as conspicuous as the lat-
erals and little arched between them, 1 mm. or less from margin; inflorescence
axillary, usually short and racemoid, the axis 0.5-1.5 (-3) cm. long, about 6-flow-
ered, the terminal flower usually aborting; pedicels 4.5-12 mm. long, 0.5 mm.
thick, often longer than the axis, appressed or ascending in flower, spreading in
fruit; bracts somewhat persistent, paired, narrowly triangular, acute, 2-3 mm.
long; bracteoles like the bracts but broader and connate-perfoliate, forming a
boat-shaped cup about 1 mm. high and 3.5-4.5 mm. long, which is persistent
until after the fruit matures; margins of the bracteoles often scarious, glabrous;
calyx-lobes 4, oblong with rounded tip, silky on both sides, at maturity 2 mm.
wide, 2-2.5 mm. long; disk flat, strongly 4-angled, about 2 mm. wide, somewhat
pubescent or the center glabrous; style 4-6 mm. long, glabrous; petals white,
elliptic, about 3 mm. wide, 5 mm. long, ciliate-f ringed at tip; stamens about 50,
4-7 mm. long; fruit oblong or subglobose, glandular-verruculose, about 6 mm.
long and nearly as thick.

The inflorescence in this species is extremely plastic. It may be
regarded as an axillary branch which may at one extreme be so much
reduced that the axis is not readily apparent and the "raceme" ap-
pears as a solitary axillary flower. At the other extreme the "raceme"
may bear one or two pairs of flowers at the base and continue above
this as a leafy vegetative branch. The inflorescence may be strictly
"racemose," that is, with 2-5 pairs of opposite, decussate flowers;
in slightly more expanded variants, the upper nodes may bear 2
flowers each, or the lowermost node may bear 2 short racemes in-
stead of 2 solitary flowers; in this latter case the lowermost node is
actually in the axil of the leaf, and the inflorescence appears as 3
racemes, one above another in the axil.

Because of this plasticity of the inflorescence, and because of the
unusually high degree of variability in the foliage of this species,
many segregate species have been described. For a synopsis of the
West Indian taxa belonging to this complex, see Bot. Jahrb. 19:
628-632. 1895; for additional synonymy pertaining to South Amer-
ican taxa, see Field Mus. Bot. 13, pt. 4: 684. 1958. Two taxa treated

McVAUGH: FLORA OF GUATEMALA 335

in this Flora as distinct species, Eugenia Karwinskyana and E. yuca-
tanensis, are perhaps too closely related to E. biflora (cf. Fieldiana,
Bot. 29:428-430. 1963).

Eugenia bumelioides Standl. Carneg. Inst. Wash. Publ. 461:
75. 1935. E. tapirorum Standl. Field Mus. Bot. 9: 321. 1940.

Dense wet mixed forest, 650 meters or less; Pete"n. Mexico (Ta-
basco) ; British Honduras, the type from Camp 32, British Honduras-
Guatemala boundary, W. A. Schipp 1270; Honduras (Dept. Atlan-
tida, Mt. Cangrejal, near La Ceiba, Yuncker et al. 8700, type of
E. tapirorum).

A tree 6-8 meters high, 10 cm. in diameter, ferruginous-tomentose when
young; leaves and branchlets glabrate, or the latter persistently puberulent;
young growth and inflorescence (including the hypanthium and both surfaces
of the calyx-lobes) densely matted with coppery-brown shining irregularly con-
torted and appressed dibrachiate hairs up to 0.5 mm. long, these sparingly per-
sistent even in fruit; leaves elliptic-ovate or -obovate, caudate-acuminate, 1.7-
4 cm. wide, 4-8 cm. long, 1.7-2.7 times as long as wide, the slender obtuse apex
7-12 mm. long, shorter in some small leaves; base of blade acute, the margins
cuneately decurrent on the channeled petiole 4-8 mm. long; midvein impressed
above; lateral veins about 10-15 pairs in addition to some intermediate ones,
obscure, ascending and parallel; marginal vein about equaling the laterals and
somewhat arched between them, 0.5-1.5 mm. from the margin; blades strongly
discolorous in drying, dark green and apparently lustrous above, paler and spar-
ingly dark-dotted beneath; inflorescence a short axillary raceme up to 3-5 mm.
long, bearing 1-3 opposite and decussate pairs of flowers on pedicels 5-6 mm. long
(or a few flowers solitary at leafless nodes near the bases of new shoots); buds 3.5-
4 mm. long, pyriform, the hypanthium narrowly cup-shaped, 1.5 mm. long; bracts
broadly ovate, dark, scarious, about 0.7 mm. long and wide; bracteoles distinct,
narrowly ovate, acute, persistent, 1-1.5 mm. long; calyx-lobes suborbicular, about
1.5 mm. long and wide (-2 mm. in fruit); disk 1.5-1.7 mm. wide, hairy; style
about 5.5 mm. long; stamens about 100, up to 5 mm. long, the anthers 0.4 mm.
long; petals obovate, 3.5 mm. long; ovary bilocular, the ovules 12-15 in each
locule, in a subcapitate group; fruit globose or essentially so, glandular- verru cose,
black, glabrous or nearly so, 10-12 mm. in diameter or more.

Eugenia buxifolia (Sw.) Willd. Sp. PI. 2: 960. 1800. Myrtus
buxifolia Sw. Prodr. 78. 1788. E. mayana Standl. Contr. U. S. Nat.
Herb. 23: 1042. 1924. Guayabillo.

In advanced deciduous forest, or in second growth, in calcareous
regions, near sea level; Pete"n. British Honduras (Caves, Stann Creek
Railway, W. A. Schipp 427) ; Yucatan Peninsula of Mexico (the type
of E. mayana from Yucatan, Gaumer 714); southern Florida; Ba-
hamas; Greater Antilles; St. Thomas; St. Croix.

336 FIELDIANA: BOTANY, VOLUME 24

A small tree, up to 12 meters high, the trunk up to 12 cm. in diameter; branch-
lets, inflorescence, petioles and at least the young leaves puberulent, bearing
numerous erect or incurved sharp hairs up to 0.3 mm. long; leaves obovate or
elliptic, widest at or above the middle, 1-2.5 cm. wide, 2.5-5.5 cm. long, 1.5-
2.3 times as long as wide, mostly rounded at apex, cuneate at base, the margins
often revolute, decurrent on the straight terete or plano-convex petioles 1.5-3 mm.
long; mid vein on the upper surface flat or sulcate, often persistently pubescent;
lateral veins few and irregular, 2-5 on each side, at least 1 or 2 basal ones strongly
ascending, the apical ones usually recurving and joining the next succeeding ones,
but a definite marginal vein not formed; glands not apparent on the upper sur-
face, usually small but evident on the lower; inflorescence an axillary raceme, or
2-3 racemes superposed, the axis 1-4 mm. long with 2-5 decussate pairs of flowers,
the terminal flower usually aborted; pedicels 1-2 mm. long, ascending; bracts sub-
persistent, 0.5 mm. long; bracteoles persistent, ovate, acute or obtuse, scarious,
convex on the backs, not connate, 0.4-0.6 mm. long; buds 1.5 mm. long, the hypan-
thium cup-shaped, lightly pubescent; calyx-lobes 4, about equal, scarious, rounded,
ciliate, glabrous within, 0.5-0.7 mm. long and wide; disk 1-1.5 mm. wide, some-
times quadrate; style glabrous, 3.5 mm. long; stamens about 40, about 2.5 mm.
long, the anthers 0.3 mm. long; petals glabrous, about 1.7 mm. long and wide;
ovary bilocular, the ovules 5-6 in a subcapitate group in each locule; fruit globose
or oblate, about 5 mm. in diameter, black.

The Maya names of Yucatan are "sacloob" and "khilnich."
There seems to be no obvious difference between Central Amer-
ican and West Indian material of this species, except that in some
Antillean specimens the branchlets are glabrous or nearly so, a fact
long ago noted by Urban (Bot. Jahrb. 19: 639. 1895).

Eugenia cacuminum Standl. & Steyerm. Field Mus. Bot. 22:
357. 1940.

Known only from the type, Dept. Chiquimula, Montana Nube
(Montana Volcancitos), between Socorro Mountain and Cerro Brujo,
southeast of Conception de las Minas, 1500-1700 meters, Steyermark
30903.

A shrub 3 meters high, nearly glabrous at maturity, the young branchlets and
petioles and the inflorescence finely hispidulous; leaves coriaceous, rigid in drying,
elliptic or varying to narrowly ovate, 1.5-3 cm. wide, 3-7 cm. long, 2-2.5 times as
long as wide, obtusely pointed or bluntly acuminate, acute at base, the thick
revolute margins decurrent on the nearly terete shallowly channeled petiole 4-
6 mm. long; midvein pale above, concave in the flat surface of the blade; lateral
veins 5-6 on each side, inconspicuous especially above, more evident beneath,
connected by irregularly angled arches 1.5-3 mm. from the margin, the marginal
vein ill-defined; blades dark green and glossy above, paler and somewhat obscurely
gland-dotted beneath; inflorescence an abbreviated axillary raceme, the axis 4-
angled, 2 mm. long or less, bearing 1-2 approximate pairs of sessile flowers (or the
pedicels up to 0.7 mm. long and thick); bracts indurate, ovate, 1 mm. long; brac-
teoles elliptic-ovate, blunt, distinct, probably deciduous, about 0.7 mm. long and

McVAUGH: FLORA OF GUATEMALA 337

wide; buds not seen; hypanthium probably 1.5-2 mm. long in anthesis, minutely
pubescent, its body from the earliest stages nearly globose and prominently 8-
ridged, constricted above into a narrow neck, then abruptly expanded into the
large subfoliaceous calyx; wing-like ridges of the half-grown fruit 2-2.5 mm. high,
extending through the narrow neck to the base of the calyx, their height about
equal to the thickness of the body; calyx-lobes rotund, membranous, ciliate, mi-
nutely puberulent without, glabrous within, concave, up to 4 mm. long and wide,
pale and prominent on the immature fruit; disk 3.5 mm. wide; immature fruit
apparently fleshy, shrunken in drying, the body including the wings about 8 mm.
in diameter, 1 cm. long including the neck; ovary bilocular, the ovules 5 in each
locule.

Eugenia Capuli (Schlecht. & Cham.) Berg, Linnaea 27: 238.
1856. Myrtus Capuli Schlecht. & Cham. Linnaea 5: 561. 1830.
Eugenia Schiedeana Schlecht. Linnaea 13: 418. 1839. E. Capuli a
micrantha Berg, Linnaea 27: 239. 1856. E. Capuli ft macroterantha
Berg, Linnaea 27: 239. 1856. E. Capuli 7 rigida Berg, Linnaea 29:
238. 1858. E. Contrerasii Lundell, Wrightia 2: 206. Sep. 1961.
E. tenuissima Lundell, Wrightia 3: 19. Dec. 1961.

Forested hills, ridges, river bluffs, mostly at low or very low ele-
vations; Pete"n (Tikal National Park, Lundell 16597, the type of
E. Contrerasii); Alta Verapaz; Izabal; Escuintla; perhaps also in
Chiquimula, Retalhuleu, and Huehuetenango. Mexico (west to
Jalisco and Sinaloa; north to Tamaulipas, the type from Papantla,
Veracruz, Schiede); British Honduras; Honduras.

Shrub or small slender tree to 8 meters high, 10 cm. in diameter, hispidulous,
the foliage glabrate, the inflorescence, branchlets and young growth densely beset
with small sharp erect or ascending hairs 0.1-0.2 mm. long; leaves ovate, lanceolate
or sometimes elliptic, usually broadest below the middle, (0.8-) 1.5-2.5 cm. wide,
(2-) 3-6.5 cm. long, 2-3 (-4) times as long as wide, about equally narrowed to
both ends, or (usually) prolonged distally, the apex narrow and blunt, acute or
acuminate, often caudate-acuminate with slender tip 1 cm. long; base cuneate
(usually) or acute, the margins decurrent on the inner angles of the flat or concave
petiole 3-5 mm. long; midvein flat or convex above; lateral veins ascending,
6-10 pairs, inconspicuous above, more evident beneath; marginal vein about as
thick as the laterals, arched between them, 1-2 mm. from the margin; leaves
dark green above, paler and often drying brown beneath; glands in young dried
leaves often seen as convexities on both surfaces, in older leaves usually evident
as dark points on the lower surface; inflorescence a short axillary raceme, sometimes
with 1-2 additional racemes superposed, the axis 3-6 (-16) mm. long, bearing 3-4
opposite decussate pairs of flowers on pedicels 3-5 mm. long; buds 2 mm. long,
pyriform; bracts scarious, 0.5-1 mm. long; bracteoles ovate, obtuse, persistent,
about 0.5 mm. long and wide, nearly contiguous at base; hypanthium campanu-
late, very finely puberulent, 1 mm. long; calyx lobes in 2 unequal pairs, thin,
pubescent on both sides, broadly rounded, 0.8-1 mm. wide and nearly as long;
disk about 0.9 mm. wide, glabrous or nearly so; style about 4-5.5 mm. long; sta-

338 FIELDIANA: BOTANY, VOLUME 24

mens ca. 50, up to 3 mm. long; petals white; ovary bilocular, the ovules in each
locule ca. 8-10, radiating from a centrally attached placenta; fruit probably black,
globose, ca. 5 mm. in diameter.

This species is not clearly distinct from a West Indian species,
Eugenia monticola (Sw.) DC., and by some it is regarded as a variety
of E. monticola (cf. Urban, Bot. Jahrb. 19: 636. 1895). For notes
on the taxonomy of E. Capuli and related species, see Fieldiana,
Bot. 29:432-435. 1963.

A very narrow-leaved plant that may be a form of E. Capuli,
namely E. Lindeniana Berg, Linnaea 29: 240. 1858, is known from
Tabasco (Linden 619, the type) and Oaxaca, but apparently has not
been found in Guatemala. The leaves are narrowly lanceolate or
linear-lanceolate, about 1 cm. wide or less, 6-8 times as long as wide.

Eugenia capulioides Lundell, Contr. Univ. Mich. Herb. 7: 32.
1942. Escoba (Huehuetenango, according to Standley).

Forested mountains, in advanced forest, or along rivers, 1000-
1800 meters; Huehuetenango (between La Libertad and Paso del
Boqueron, along Rio Trapichillo, Standley 51106). Mexico (Chia-
pas, near Siltepec, E. Matuda 4521, the type).

A shrub or small tree up to 5 meters high, quite glabrous except the ciliate
tips of the bracts; leaves narrowly elliptic or lance-elliptic, often about equally
narrowed to a slender but obtuse acumen and an acute or cuneate base, 1.3-
2.6 cm. wide, 3.5-8.5 cm. long, about 2.6-4 times as long as wide, the margins
at base cuneately decurrent on the inner angles of the channeled petiole 2.5-
5 mm. long; midvein flat or slightly elevated above on the plane surface of the
blade, usually appearing as a pair of closely parallel convex ridges with a shallow
furrow between them; lateral veins 5-7 on each side, inconspicuous, more evident
beneath, the lowest 1-2 pairs leaving the midvein at a narrow angle and ascend-
ing more or less parallel to the margin, the distal veins diverging and nearly
straight, passing into a nearly straight marginal vein 1-2 mm. from the margin;
blades nearly concolorous, usually gland-dotted beneath, sometimes with gland-
ular convexities above; inflorescence an axillary raceme (usually solitary, but
occasionally 2-3 superposed), the axis up to 5-7 mm. long, with 4-6 opposite and
decussate floriferous nodes; racemes proliferous, the flowers often 2-3 in the axil
of each bract; pedicels 1.5-3 mm. long; bracts scarious, subpersistent, ovate,
divaricate, 0.5-0.7 mm. long; bracteoles rounded or truncate, thickened and
glandular at base, persistent but not connate, 0.5 mm. long; buds about 2.5 mm.
long, pyriform, the campanulate hypanthium 1 mm. long or a little more, dotted
with convex glands; calyx-lobes broadly rounded, the larger ones 1-1.2 mm. long
and wide; disk 1-1.2 mm. wide in flower; style not seen; stamens about 50; petals
white, obovate, about 2.5 mm. long; ovary bilocular, the ovules 5-6 in each locule,
pendent in a loose group; fruit apparently globose, 5-6 mm. in diameter.

Eugenia cervina Standl. & Steyerm. Field Mus. Bot. 23: 128.
1944. Cacho de venado.

McVAUGH: FLORA OF GUATEMALA 339

Wet forest, 400 meters or less; Alta Verapaz, the type from near
Rio Icvolay, 5 miles northwest of Cubilgiiitz, Steyermark 44691.
Endemic.

Tree 8-12 meters tall, the young branchlets and petioles, inflorescence and
lower leaf-surface densely appressed-pubescent with lustrous coppery dibrachiate
hairs up to about 0.5 mm. long; leaves persistently tomentose beneath with a
close coat of similar but paler ("silvery brown" or "buff brown") hairs; leaves
elliptic or elliptic-oblanceolate, 2.5-5.5 cm. wide, 9-20 cm. long, (3-) 4 times as
long as wide, about equally narrowed to the caudate-acuminate tip 1-2 cm. long,
and to the acute base, the margins rather abruptly decurrent on the petioles 6-
8 mm. long, 1.3-1.5 mm. thick; midvein sharply and narrowly impressed above;
lateral veins 15-20 on each side, inconspicuous, ascending, rather close and par-
allel, convex above, mostly obscured by the tomentum beneath; marginal vein
about equaling the lateral ones and arched between them, 0.5-2 mm. from the
margin; leaves dark green and probably glossy above, obscurely glandular- verru-
cose, the lower surface covered by the tomentum; inflorescence racemose, the
axis almost none or up to 6 mm. long, 2 mm. thick, the flowers 4 pairs or fewer,
nearly sessile, the fruiting pedicels up to about 1.5 mm. long and thick; bracts
probably short and indurated; bracteoles persistent, becoming indurated, ovate-
orbicular, in fruit 2-2.5 mm. long and wide, glabrous within, barely connate and
forming a tomentulose involucre that persists when the fruit falls; flowers and
buds not seen; fruit "buff -brown or tinged with rose," subglobose, tomentulose,
1.8-2.3 cm. in diameter, crowned by the broadly rounded calyx-lobes, these up
to 4 mm. long and wide, appressed-pubescent on both sides with pale reddish
hairs; disk (3.5-) 4.5 mm. wide.

Eugenia chiapensis Lundell, Amer. Midi. Nat. 20: 238. 1938.

A mountain species, collected at an elevation of 1600 meters;
to be expected in Guatemala. Known only from near Siltepec,
Chiapas, less than 50 km. from the Guatemala border, Matuda 1904,
the type.

A tree 4 meters high, 10 cm. in diameter, the youngest branchlets and leaves
densely tomentose with brown contorted hairs up to 1 mm. long; hairs soon de-
ciduous, the mature foliage, branchlets and inflorescence glabrous except for a
few cilia on the perianth-segments; leaves elliptic, 3-5.5 cm. wide, 8-15 cm. long,
about 2.5-3.5 times as long as wide, about equally narrowed to the bluntly short-
acuminate tip and the somewhat rounded base, the margins at the very base
cuneately decurrent on the sulcate petiole 8-12 mm. long, 1.5 mm. thick; mid-
vein impressed above; lateral veins about 10 on each side, evident on both sur-
faces, more conspicuous beneath, connected by symmetrical rounded arches about
2-4 mm. from the margin; blades probably dark green and lustrous above, paler
beneath and abundantly dotted with small convex resinous glands; flowers 1-3
in sessile umbel-like groups from short spurs at leafless nodes, surely racemose
but the crowded axes suppressed, the slender pedicels 8-16 mm. long and 0.7 mm.
thick seeming to arise directly from the node; bracts broad, indurated; bracteoles
membranous, rounded, divaricate, broadly ovate, connate and forming a per-
sistent involucre 2-2.2 mm. wide; buds pyriform, 7-8 mm. long; hypanthium

340 FIELDIANA: BOTANY, VOLUME 24

2.5 mm. long, substipitate, abruptly expanded into the globe of the perianth;
calyx-lobes rounded, membranous, gland-dotted, in unequal pairs, the larger ones
up to 6 mm. long and wide; disk quadrate, 3.5-4 mm. wide; style 10 mm. long;
stamens about 200, the anthers 1.5-1.7 mm. long; petals gland-dotted, 10-12 mm.
long and wide; ovary bilocular, the ovules about 8 in each locule, stalked, radiating
in one plane and forming a peltate group; fruit fleshy, probably globose, about
2.5 cm. in diameter.

Eugenia chinajensis Standl. & Steyerm. Field Mus. Bot. 23:
129. 1944.

Savannahs and savannah borders, 100-300 meters; endemic;
Alta Verapaz, the type from the south side of Cerro Chinaja, Steyer-
mark 45142.

A shrub 1.5-4.5 meters high, setose, the branchlets, inflorescence and lower
leaf-surfaces covered with erect stiff sharp hollow reddish-brown hairs up to 1-
1.2 mm. long; leaves glabrate above except the midvein, oblong-elliptic, 1.2-4 cm.
wide, 3.5-9.5 cm. long, 2.5-3.5 times as long as wide, gradually and broadly acumi-
nate or the tip acute; blades somewhat narrowed below the middle, then abruptly
rounded to a subcordate base, often appearing subsessile, the strongly revolute
margins passing abruptly onto the petiole and seeming to meet across the flat
base of the midvein in a zone marked by elevated persistent bases of hairs; petiole
2.5-3 mm. long; midvein (above the flat base) depressed below the surface of the
blade, rather persistently setose; lateral veins (7-) 10-12 on each side, distant,
clearly marked as fine lines above, evident but less conspicuous beneath; marginal
vein about equaling the lateral ones, forming long high asymmetrical arches 2-
4 mm. from the margin; blades "dull green" above, paler and drying reddish brown
beneath; glands inconspicuous; flowers in small sessile axillary glomerules, the
stout racemose axis 2-2.5 mm. long, 1.5 mm. thick, bearing about 2-3 approxi-
mate decussate pairs of flowers, the axis itself concealed even after the fruit has
fallen by the prominently setose-fringed narrow bracts, and the similar but
broader bracteoles which are round-obovate, glabrous within, about 0.7 mm.
long and wide, not connate but forming a persistent involucre; buds probably
about 2 mm. long, obovoid, the glabrous hypanthium campanulate or hemi-
spheric; calyx-lobes 4, concave, in unequal pairs, glabrous within, setose without
and on the margins, the outer ones rounded or broadly pointed, 1 mm. long, the
inner ones broadly rounded, 1.5-1.7 mm. long and wide; disk glabrous, 1.5-1.8 mm.
wide; style not seen; stamens 35-40, the anthers 0.4 mm. long; petals suborbicu-
lar, nearly eciliate, about 2.5 mm. long; ovary bilocular, the ovules 3 in each
locule, collateral; fruit nearly glabrous, coarsely glandular-verruculose, subglobose
or in 2-seeded forms laterally elongate, about 6 mm. long just before maturity,
crowned by the erect persistent calyx-lobes; seed round or hemispheric, 5 mm.
long, with very lustrous cartilaginous testa free from the homogeneous embryo.

This is one of the most distinctive species of Eugenia discovered
in Central America in recent years. The coarsely setose herbage,
oblong subcordate leaves and sessile clustered flowers and fruits com-
bine to give it a unique aspect.

McVAUGH: FLORA OF GUATEMALA 341

Eugenia choapamensis Standl. Field Mus. Bot. 22: 93. 12 June
1940. E. simiarum Standl. & Steyerm. I.e. 359. 31 Oct. 1940.

Wet lowland forest, 375 meters or less; Izabal (Montana del
Mico, Steyermark 38274, type of E. simiarum}. Mexico (Oaxaca,
Mexia 9254, type; Tabasco; Chiapas).

Tree 7-12 meters high, quite glabrous except the finely ciliate bracts, bracte-
oles and perianth-parts, and the bristly-hispid staminal ring; leaves at maturity
coriaceous, elliptic or elliptic-lanceolate, (2.5-) 4-8 cm. wide, (7.5-) 12-16 cm.
long (-22 cm. long on shoots), 2.8-3.3 times as long as wide, about equally nar-
rowed to the gradually and bluntly acuminate tip and to the acute or cuneate
base, the thickened callous margins decurrent on the dilated flat or concave sum-
mit of the petiole 5-8 mm. long, 1.5 mm. thick; midvein broad and flat or convex
above, sometimes with very narrow median channel near base; lateral veins 10-14
on each side in addition to some intermediate ones, straight, inconspicuous above,
evident and somewhat elevated beneath, connected by a series of marginal arches
3-5 mm. from the margin; upper surface "rich green," when dry nearly feature-
less or the veins evident, the lower "silvery pale green" (according to Steyermark),
often drying reddish brown, the few large glands scarcely apparent; flowers in
nearly sessile umbel-like fascicles on very short spurs at leafless nodes, the inflores-
cence a much-crowded raceme (or 2-3 superposed), its axis up to 2-3 mm. long
(usually shorter), 2 mm. thick, bearing 1-5 approximate pairs of flowers on slender
pedicels (4-) 10-18 mm. long (in fruit up to 1 mm. thick); bracts broadly ovate,
imbricate, indurate, 1 mm. long; bracteoles persistent, distinct, ovate, obtuse,
1-1.5 mm. long; buds 4.5-6 mm. long, pyriform, the globe of the petals much
broader than the cup-shaped hypanthium 1.5-2 mm. long; calyx-lobes deltoid-
ovate or broadly rounded, deeply concave, membranous, the larger ones 3-3.5 mm.
wide, 2-2.5 mm. long; disk 2.5 mm. wide; style about 7 mm. long; stamens more
than 100, longer than the style; petals white; ovary bilocular, the ovules 10-16
in each locule, in a compact group arising centrally from the dissepiment; fruit
smooth, black, ellipsoid, 1-1.5 cm. in diameter, 1.5-2.5 cm. long.

Eugenia citroides Lundell, Contr. Herb. Univ. Mich. 7: 33.
1942.

In high wet mountain forests, often in barrancas with Abies,
2100-2700 meters; San Marcos; Huehuetenango; Quiche". Mexico
(Chiapas, the type from near Siltepec, E. Matuda 4554).

A large shrub, or a tree 6-10 meters high, soon glabrate, the young branchlest
and foliage softly and sometimes densely tomentose with pale brown somewhat
appressed hairs 0.2-0.6 mm. long, some of the hairs dibrachiate but attached very
near one end; leaves elliptic or on slowly growing branches obovate, abruptly or
gradually acuminate, or occasionally obtuse in obovate leaves, acute at base, the
rather thick revolute margins somewhat decurrent on the stout petioles 5-10 mm.
long (or longer on vigorous shoots), 1-2 mm. thick; blades 3-5.5 cm. wide, (6-)
8-13 cm. long, 1.8-2.4 times as long as wide; midvein narrowly impressed above;
lateral veins 6-10 on each side in addition to some intermediate ones, evident as
pale convex lines above, elevated and more prominent beneath, there seeming

342 FIELDIANA: BOTANY, VOLUME 24

stronger than the intermediate veins, connected by a series of irregularly angled
arches 3-6 mm. from the margin, a series of small arching submarginal veins be-
yond these; blades dark green and lustrous above, paler beneath; small convex
glands usually evident beneath, sometimes also above; flowers in dense axillary
or partly supra-axillary glomerules, as many as 50 or more at a node, often fewer,
racemose in arrangement, the racemes usually several crowded together, the indi-
vidual axes up to 3 mm. long, bearing 3-4 pairs of sessile flowers; bracts ciliate,
small and indurate, persistent; bracteoles ciliate, ovate, 1-1.3 mm. long, 0.7 mm.
wide, becoming indurated and persistent; buds 3-4 mm. long, pyriform, nearly
glabrous, the campanulate hypanthium 1.5 mm. long, strigose; calyx-lobes mem-
branous, glabrous, up to 2 mm. long and wide, the larger ones in fruit 2 mm.
wide, 2.5 mm. long; disk 1.7-2 mm. wide, glabrous; style about 6 mm. long; sta-
mens about 50, the anthers 0.5-0.6 mm. long; petals white; ovary bilocular, the
ovules 4 in a subcapitate group in each locule; fruits on pedicels up to 1.5 mm.
long, 2 mm. thick; fruit subglobose, dark red (according to Standley), 2.7-3.3 cm.
in diameter, 3-3.5 cm. long, with thin flesh and massive seed 2.5 cm. long.

Eugenia farameoides A. Rich. Ess. Fl. Cub. 588. 1845. E. flan-
folia Standl. Field Mus. Bot. 8: 320. 1931. E. ceibensis Standl. Field
Mus. Bot. 9: 319. 1940.

Pine ridges or open swampy places, at or little above sea level.
British Honduras (the type of E. flavifolia from Stann Creek Rail-
way, W. A. Schipp 450); Mexico (Oaxaca); Honduras (the type of
E. ceibensis from near La Ceiba, Dept. of Atlantida, T. G. Yuncker
et al. 8309) ; Cuba.

A shrub or small tree, sometimes 8 meters high with trunk 12-15 cm. in diam-
eter, nearly glabrous, the young branchlets and petioles and the inflorescence
minutely pubescent with sharp erect hairs up to 0.1 mm. long; leaves when dried
markedly yellow-green, coriaceous, elliptic to ovate, 2-5.5 cm. wide, 5-10 cm. long,
2-3 times as long as wide, acute or acuminate or infrequently broadly rounded,
the very tip blunt or rounded even in narrowly acute forms; base gradually or
somewhat abruptly rounded, the margins usually (and sometimes cuneately) de-
current on the rough, channeled petiole 1-1.5 mm. thick, 4-8 mm. long; midvein
convex on the upper surface, or especially toward the base of the blade flat or
even sulcate; lateral veins evident on both surfaces and somewhat elevated (con-
vex) when dry, 5-8 pairs in addition to some intermediate ones, scarcely forming
a marginal vein but each arching over (often in a series of angular bends) to join
the next succeeding one; blades lustrous and smooth above, paler and dull beneath,
the glands usually not apparent on the upper surface, the lower surface usually
minutely dark-dotted or even slightly roughened by the glands; inflorescence a
short axillary raceme (or two racemes superposed), the axis somewhat 4-angled,
4-12 mm. long, with 4-6 opposite, decussate pairs of flowers; terminal flower
usually aborting; pedicels filiform, straight, 3-5 mm. long; bracts scarious, some-
what persistent, 0.5-1 mm. long; bracteoles united by their proximal margins, in
anthesis forming a boat-shaped envelope surrounding and concealing the cup-
shaped hypanthium, the envelope 0.7 mm. high, 1.5 mm. long; bracteoles in fruit
explanate, closely investing the base of the hypanthium, forming a scarious in-

McVAUGH: FLORA OF GUATEMALA 343

volucre 2 mm. long, 1.5 mm. wide; hypanthium densely and very minutely pubes-
cent with pale appressed hairs unlike those of the pedicels; calyx-lobes 4, broadly
rounded, appressed-pubescent on both surfaces, about 1.5 mm. long in anthesis,
accrescent, persistent and conspicuous in fruit, when up to 2 mm. wide and long;
disk 1.5-1.7 mm. wide, bristly in the staminal ring, and with a dense ring of hairs
surrounding the sparsely hairy base of the style; style 7 mm. long; stamens 60-75,
about 5 mm. long, the anthers about 0.5 mm. long and wide; petals white, puberu-
lent without, 3-4 mm. long; ovary 2-locular, the ovules 6-7 in each locule, radiating
from a small attachment on the central dissepiment; fruit when ripe black (Schipp),
transversely oval, 7-8 mm. in the longer axis, about 5 mm. in the shorter, 5-6 mm.
high (i.e. from bracteoles to calyx).

Central American specimens of E. farameoides, including the
plant described as E. flavifolia, rather consistently have the leaves
2-3 times as long as wide and the axis of the racemes 6 mm. long or
more. Some Cuban specimens are precisely similar, but in others
the leaves are often less than twice as long as broad, and the axis
of the raceme is much abbreviated.

Eugenia flavoviridis Lundell, Amer. Midi. Nat. 29: 479. 1943.
E. dissitiflora Lundell, Wrightia 2: 207. Sep. 1961. E. flavida Lun-
dell, Wrightia 3: 14. Dec. 1961. Guayabillo.

In low forest, often associated with Metopium or Haematoxylum
(chechemal or tinted'); Pete"n (Santo Toribio, E. Contreras 2684, the
type of E. dissitiflora) . British Honduras (Toledo District, Monkey
River, Jenkins Creek, P. H. Gentle 4085, the type).

Treelike, 2-3 meters high, nearly glabrous, the young branchlets and inflores-
cence sparingly setose-pubescent with coarse sordid antrorsely curved or almost
erect hairs 0.2-0.5 mm. long; leaves ovate to oblong or elliptic, 2-4.8 cm. wide,
(4-) 5.5-9 (-11) cm. long, 2-3 times as long as wide, obtusely acuminate, rounded
or acute at base, the margins finally cuneately decurrent on the channeled petiole
4-7 (-11) mm. long; midvein impressed above; lateral veins 8-10 including some
intermediate ones, very inconspicuous above, more evident beneath; marginal
vein about equaling the lateral ones and arched between them, often irregularly
angled, 1.5-4 mm. from the margin; leaves smooth and probably darker green
above, often drying yellowish green especially beneath, finely dark-dotted beneath;
inflorescence an axillary raceme (or 2-3 racemes superposed), the axis up to 5-
12 mm. long, bearing 4-8 pairs of flowers on pedicels 1.5-4 mm. long and up to
0.5 mm. thick in fruit; bracts 0.5-0.8 mm. long, divaricate, subpersistent, or their
bases ovate, indurate-persistent; bracteoles broadly deltoid-ovate, ciliate, thin but
subpersistent, scarcely connate, broader than long, 0.5 mm. long; buds probably
somewhat more than 2 mm. long; flowers not seen; hypanthium hemispheric or
short-campanulate, glabrous or nearly so; style about 5 mm. long; fruit globose,
5-6 mm. in diameter or more, probably black, its disk a little more than 1 mm.
wide, the calyx-lobes appressed over the disk, flat, obtusely deltoid, the larger ones
0.5-0.6 mm. long, 0.7-1 mm. wide.

344 FIELDIANA: BOTANY, VOLUME 24

Eugenia guatemalensis Bonn. Sm. Bot. Gaz. 23: 245. 1897. E.
patalensis Standl. & Steyerm. Field Mus. Bot. 23: 130. 1944. Guaya-
billo; silich (Huehuetenango) ; rutzache-ska (according to Aguilar).

Wooded moist forests, oak or pine forests, or according to Stand-
ley on dry open hillsides, 800-2000 meters; Baja Verapaz, the type
from Santa Rosa, Tuerckheim 1318; ?Huehuetenango; Santa Rosa;
Guatemala; Sacatepe"quez; Chimaltenango. Southern Mexico (Edo.
de Mexico; Oaxaca; Veracruz; Chiapas); Costa Rica.

A shrub or small tree 2-6 meters high, the branchlets and the lower leaf-
surfaces and the inflorescence cinereous- or pale reddish-pubescent (or the inflores-
cence golden) with very closely ap pressed small dibrachiate hairs; leaves elliptic
to oblanceolate or obovate, 1.5-4 cm. wide, (4-) 5-9 cm. long, 2-2.7 times as long
as wide, often about equally narrowed to both ends, gradually or abruptly acumi-
nate (the tip blunt), acute at the base, the margins decurrent on the channeled
petiole 7-10 mm. long, 1 mm. thick; midvein impressed above; lateral veins incon-
spicuous, 8-10 on each side including some intermediate ones; marginal vein about
equaling the lateral ones and arched between them, 0.5-2 mm. from the margin;
blades lustrous and dark green above, dark- or convex-dotted; inflorescence an
axillary raceme (often 1-2 additional ones superposed), the axis 1-2.5 cm. long,
bearing up to 5 opposite and decussate pairs of flowers on pedicels 3-7 mm. long;
bracts lanceolate, up to 1.5 mm. long; bracteoles scarious, distinct, similar to the
bracts, 1-1.5 mm. long, subpersistent; buds 3.5 mm. long; hypanthium densely
appressed-pubescent, obovoid, substipitate, 1.5 mm. long; calyx-lobes in unequal
pairs, appressed-pubescent on both surfaces, separating in anthesis by splitting in
the sinuses between them; smaller lobes deltoid-ovate, 1.5-2 mm. long and wide, the
larger ones cucullate-concave, 2.5-3 mm. long and wide; disk 2-2.7 mm. wide,
hairy about the base of the style; style 5-6 mm. long; stamens about 125, up to
5 mm. long; petals glabrous, about 4 mm. long; ovary bilocular, the ovules 15-18
in a compact group in each locule; developing fruit pyriform, the mature fruit 1 cm.
long or more, probably black and oblong or obovoid.

According to Standley (mss.) the mature fruit may be subglobose;
no Guatemalan specimens with mature fruits are at hand, but rather
numerous specimens from cloud-forest areas in the mountains of
Honduras have been referred to E. guatemalensis by Standley and
others. These specimens have nearly globose fruits, and rather more
symmetrically elliptical leaves than Guatemalan plants. No ade-
quate flowering specimens are available; it is probable that these
represent another species.

The type of E. patalensis Standl. & Steyerm., from Fatal, Baja
Verapaz, Standley 69568, is a specimen with half-grown fruit. This
and other specimens from the type-region of E. guatemalensis have
more stiffly coriaceous leaves than flowering specimens of that spe-
cies, and have lost most of their characteristic pubescence except a
little on the fruits and the pedicels. For the present this species
may be regarded as synonymous with E. guatemalensis.

McVAUGH: FLORA OF GUATEMALA 345

Eugenia hondurensis A. Molina, Ceiba 1: 261. 1951. E. crassi-
folia A. Molina, Ceiba 3: 169. 1953. E. nicaraguensis Amsh. Act.
Bot. Ne"erl. 5: 278. 1956.

Wooded moist ravines, moist brushy rocky hills, dense wet mixed
forest, 1800 meters or less; to be sought in southeastern Guatemala.
Honduras (Choluteca; Morazan, Molina 2559, the type, and Molina
3031, type of E. crassifolia) ; Nicaragua (Chinandega; Chontales,
Standley 9427, type of E. nicaraguensis; Jinotega).

A shrub or small tree up to 6 meters high, the branchlets, inflorescence, peti-
oles and lower leaf-surfaces tomentose with yellowish-brown or nearly colorless
stalked forked hairs, the tomentum reddish in the young growth; leaves soon gla-
brate above except the midvein; leaves elliptic, oblong, or obovate, (2-) 4-6 (-9)
cm. wide, (4-) 6-9 (-15) cm. long, 1.5-2.5 times as long as wide (or even broader
on shoots), rounded or obtuse or rarely very shortly acuminate at apex, narrowed
from the middle or above to a truncate-auriculate base, the margins abruptly
decurrent on the inner angles of a nearly terete channeled petiole 3-5 mm. long,
1.3-1.7 mm. thick; midvein depressed a little on the upper surface, nearly flat or
sulcate, rather persistently tomentose; lateral veins 5-6 on each side, evident as
pale lines above, more prominent beneath, strongly arcuate-ascending, diminish-
ing distally, curving in at a point 2-4 mm. from the margin to join the next suc-
ceeding ones; leaves dull green above, pale beneath because of the persistent
tomentum; glands inconspicuous; flowers solitary, axillary, or at the lowest 1-2
(bracteate) nodes of leafy branchlets, the pedicels 9-15 (-30) mm. long, 0.7-
1.2 mm. thick, in fruit compressed and 1.5 mm. wide; bracts indurate-scarious,
oblong-linear, deciduous at anthesis, 3-4.5 mm. long; bracteoles similar, deciduous
at anthesis, 2.5-4.5 mm. long; buds not seen; calyx-lobes rounded, tomentose on
both sides, reflexed at anthesis, 4.5-5.5 mm. wide, 4-6 mm. long; disk 5-6 mm.
wide, tomentose about the base of the style; style pilose in the proximal one-third,
10-11 mm. long; stamens very numerous (250-300), up to 12 mm. long, the anthers
0.7-1 mm. long; petals white, obovate, ciliate, 8-10 mm. long; ovary bilocular, the
ovules (according to Amshoff) 5-10 in each locule; immature fruit ellipsoid-obo-
void, obtuse at apex, acute at base, crowned by the calyx, 8-14 mm. in diameter,
about 1.5 cm. long.

This species is variable in leaf -form and in pubescence. Flower-
ing specimens look quite unlike those in fruit. With maturity the
pubescence falls to a considerable extent, and becomes paler; the
leaves become firm, and nearly glabrous beneath. The original
specimens of E. hondurensis were from mature plants with half-
grown fruit; flowering specimens were later described under the
name of crassifolia.

Eugenia hypargyrea Standl. Contr. U. S. Nat. Herb. 23: 1044.
1924.

In wet forest or partly deciduous forest, probably 1000 meters
or less; to be expected in Guatemala. Mexico (Veracruz, near Za-

346 FIELDIANA: BOTANY, VOLUME 24

cuapan, C. A. Purpus 6171, the type; Chiapas); Honduras (Olan-
chito, Record & Kuylen H55, H55A.

A tree 10-12 meters high, the branchlets and inflorescence and the youngest
leaves closely appressed pubescent with silvery or pale reddish dibrachiate hairs,
the leaves permanently silvery-scurfy beneath; leaves obovate or obovate-elliptic,
rounded at apex, cuneate at base, 2-4.5 (-7) cm. wide, 5-11 cm. long, (1.5-) 1.8-
2.2 times as long as wide, the margins long-decurrent on the nearly terete petiole
5-9 mm. long, 1.3-2 mm. thick; midvein flat or convex above; lateral veins 4-7
on each side, strongly ascending, more conspicuous beneath and stronger than the
intermediate ones if any, connected by a series of oblique angular arches 2-8 mm.
from the margin; blades dark green, lustrous and glabrate above, the lower sur-
face obscured by hairs so small and densely aggregated as to make the surface
appear silvery-cellular; inflorescence a short axillary raceme, the axis 4-7 (-10)
mm. long, bearing 1-3 pairs of flowers on pedicels 4-7 mm. long, up to 0.7-1 mm.
thick in fruit; bracts ovate, indurate-persistent, 1 mm. long or less; bracteoles per-
sistent, broad, not connate, 0.5-1 mm. long; buds 4-4.5 mm. long; calyx-lobes
broadly rounded, in unequal pairs, the larger ones 3-4 mm. long and wide; hypan-
thium broadly campanulate, 2 mm. long, pale-reddish-tomentulose like both sur-
faces of the calyx-lobes; disk 2.5-3.5 mm. wide, hairy in the staminal ring and
about the base of the style; style about 7 mm. long; stamens about 100; petals
glabrous or nearly so; fruit apparently black, oblong-ellipsoid, 8-10 mm. in diam-
eter, 13-15 mm. long, topped by the concave tomentulose calyx-lobes.

Eugenia Ibarrae Lundell, Wrightia 2: 56. 1961. Chilonche.

Wooded swamps (tintales) dominated by species of Haematoxy-
lum, Pete"n (the type from Bajo de Santa Fe", E. Contreras 19). Brit-
ish Honduras.

Slender shrub, or a small tree 2-5 meters high, with smooth gray bark, quite
glabrous except that the bracts and the perianth-lobes are finely reddish-ciliate;
leaves rigidly coriaceous, obovate or elliptic, 1.5-3.5 cm. wide, 3-6 cm. long, 1.6-2
times as long as wide, blunt-tipped or rounded, sometimes broadly and obscurely
acuminate, cuneate at base, the cartilaginous revolute margins decurrent on the
stout short sulcate petiole 2-4 mm. long, up to 1.5 mm. wide; midvein flat to con-
vex above or narrowly sulcate especially near base; lateral veins 4-6, evident on
both surfaces in dried leaves, the lowermost pair strongly ascending and passing
directly into a marginal vein about equaling the lateral ones and arched between
them, the arches well separated (2-4 mm.) from the margin; leaves dark green
and lustrous above, paler beneath; glands seen as small convexities above, and as
abundant dark dots beneath; flowers 1-2 pairs in sessile (actually racemose) clus-
ters in the axils of foliage leaves or the axils of indurated bracts at the lowest
nodes of new leafy branchlets; axis of the raceme up to 1 mm. long and about as
thick, the flowers on gross pedicel-like bases 0.5 mm. long and thick; bracts ovate
and up to 1.5 mm. long, or the uppermost ones paired, persistent and coriaceous,
1 mm. long and wide, resembling the bracteoles; bracteoles broadly rounded, cori-
aceous, persistent, connate and forming an involucre 1-1.5 mm. long, 0.5 mm.
high in bud, more than 1 mm. wide when expanded; buds 2-2.5 mm. long, pyri-
form, the campanulate hypanthium 0.7-1 mm. long, abruptly expanded into the

McVAUGH: FLORA OF GUATEMALA 347

large globe of the perianth; calyx-lobes strongly imbricate, concave, broadly
rounded, in unequal pairs, the larger ones 1.5-2.3 mm. wide, 1.5-1.7 mm. long;
disk rather ill-defined, about 1.5 mm. wide; style 5-6 mm. long; stamens about 50,
the longer ones 5-5.5 mm. long; petals white, ovate or suborbicular, 3-3.5 mm.
long; ovary bilocular, the ovules 4 in each locule; fruit globose, reddish purple,
6-10 mm. in diameter, with thin flesh.

Eugenia jutiapensis Standl. & Steyerm. Field Mus. Bot. 23:
129. 1944.

In forests and wooded ravines, 1000 meters or less; Jutiapa, the
type from near Jutiapa, J. R. Johnston 1480. El Salvador (Santa
Tecla, Calderon 1423; Dept. Ahuachapan, Standley 20129, 20268).

A tree 4-6 meters high, the branchlets, inflorescence and lower surfaces of the
leaves (at least on the veins) hirsutulous with fine sharp pale erect hairs 0.5-
0.8 mm. long; leaves glabrate above except the midvein; leaves broadly ovate-
elliptic, 5-8.5 cm. wide, 9-18 cm. long, 1.6-2.3 times as long as wide, broadly and
bluntly short-acuminate, broadly rounded at the cordate base, the two rounded
auricles (1-) 2.5-3.5 mm. long; petioles (3-) 4.5-7 mm. long; midvein on the upper
surface broadly channeled and hirsutulous nearly its whole length; lateral veins
5-6 on each side, sometimes in addition to a few weaker intermediate ones, widely
spaced, diminishing distally, arching over and joining the next succeeding ones,
forming a series of asymmetrical arches 5-10 mm. from the margin; veins and
small veinlets elevated as pale lines on both surfaces, forming a conspicuous anasto-
mosing network of angular areolae; blades probably dark green above and paler
beneath, at least the upper surface drying somewhat purplish brown and sometimes
[the upper surface] sparingly beset with resinous glands 0.1-0.2 mm. wide; flowers
in sessile umbels at leafless nodes, the racemose axes 1 or more, almost suppressed,
up to 1.5 mm. long and thick, bearing up to 3 pairs of flowers on slender pedicels
(1-) 2-2.5 cm. long, 0.5-0.8 mm. thick; buds pyriform, 6-7.5 mm. long, the cam-
panulate hypanthium 2.5-3 mm. long, densely velutinous-hirsute; bracts somewhat
indurated, hirsute, up to 1 mm. long; bracteoles distinct, ovate, obtuse, 1 mm.
long and wide; calyx-lobes "brown-red" (according to Standley), membranous,
oblong, short-hirsute outside, minutely puberulent or almost glabrous inside, in
unequal pairs, the larger ones 4 mm. wide, 5-6 mm. long, the smaller ones 3 mm.
wide, 3.5-5 mm. long; disk glabrous, 2.5-3.5 mm. wide; style 8-10 mm. long;
stamens about 125; petals broad, nearly glabrous, ciliate, up to about 8 mm. long;
ovary bilocular, the ovules about 12 in each locule, in a subcapitate group from
near the middle of the central partition; fruit probably oval-oblong and about
2-2.5 cm. long.

Eugenia Karwinskyana Berg, Linnaea 29: 244. 1858. Eugenia
tabascensis Limdell, Phytologia 1: 482. 1941.

Wet or dry forests, at about 1500 meters; Alta Verapaz (region
of Chelae, northeast of Carcha, Standley 70388). Mexico (San Luis
Potosi; Hidalgo, Huejutla, Karwinsky 242, the type; Tabasco, Teno-
sique, E. Matuda 3561, type of E. tabascensis).

348 FIELD IANA: BOTANY, VOLUME 24

Shrub or small tree, rather generally antrorsely appressed-pubescent or stri-
gose, with sordid or silvery simple hairs 0.3-0.5 mm. long; leaves finally glabrate,
ovate or elliptic, 1.5-3.5 cm. wide, 5-11 cm. long, (2.6-) 3-4 times as long as wide,
gradually long-acuminate, the tip obtuse; base acute to cuneate, the margins de-
current on the inner angles of the petiole 3-5 mm. long; mid vein sharply and rather
narrowly impressed above, the furrow persistently short-hispid; lateral veins rather
close and parallel,. 12-20 on each side including some intermediate ones, arching-
ascending, scarcely apparent above in mature leaves, convex but inconspicuous
beneath; marginal vein about equaling the lateral ones and arched between them,
0.5-1 mm. from the margin; blades dark green and lustrous above, paler beneath;
leaves gland-dotted, the glands usually appearing as scattered resinous convexities
on the lower surface, more frequent above, where dark or translucent, or the older
leaves impressed-punctate; inflorescences racemose, 2-4 cm. long [or some flowers
solitary in the lowermost (bracteate) axils of leafy branchlets], the axis dorsi-
ventrally compressed, 1-1.5 mm. wide at base, bearing 4-5 (-9) pairs of flowers
on pedicels 4-7 mm. long, 0.2-0.4 mm. thick, the terminal flower often present
also; bracts persistent, convex on the back, ovate to lanceolate, 0.5-2 mm. long,
the distal ones longer and narrower; bracteoles broadly ovate-deltoid, convex on
the backs, somewhat coherent by their proximal margins and closely investing
the base of the hypanthium, acuminate and sharp-pointed, 1.3 mm. long, thinly
strigose and contrasting with the densely strigose, canescent hypanthium; buds
2.5 mm. long, pyriform, the campanulate hypanthium more than 1 mm. long;
calyx-lobes rounded-ovate, strigose without, closely silvery-silky within, the larger
ones 1.5 mm. wide, 1.7 mm. long; disk 1.5 mm. wide, hairy; style 4.5-5 mm. long;
stamens 50-60, about as long as the style, the anthers 0.4 mm. long; petals about
3 mm. long, white; ovary bilocular, the ovules 5 in each locule; fruit black or
reddish black, depressed-globose, about 1 cm. in diameter.

This species is certainly closely related to E. biflora and is per-
haps not specifically distinct. From other known Central American
representatives of E. biflora, E. Karwinskyana is distinguished by
having the leaves mostly elliptic (not lanceolate as is usual in E. bi-
flora), slender-pointed but not bristle-tipped, and obscurely if at all
impressed-punctate. The flowers are usually 8-10 in each raceme,
as against the usual number of about 6 (-8) in E. biflora; other
distinctions are noted in the key.

Eugenia laevis Berg, Linnaea 27: 177. 1856.

A shrub or small tree up to 10 meters high and 6-8 cm. in diameter, soon gla-
brate, the youngest branchlets and the inflorescence silky-pubescent with loosely
attached short-stalked somewhat appressed equally 2-branched lustrous pale yel-
low-white or reddish hairs 0.2-0.5 mm. long; petioles, and branchlets of the current
season sometimes minutely hispidulous; plants usually nearly glabrous at flowering
time except the inflorescence; leaves coriaceous, rather small, ovate (sometimes
narrowly so) to elliptic or obovate, about 7.5 cm. long or usually less, acute or
usually cuneate at base, the thick and often revolute margins decurrent on the
dilated summit of the channeled petiole; midvein rather sharply impressed above;
lateral veins 5-10 on each side, stronger than the few intermediate ones, obscure

McVAUGH: FLORA OF GUATEMALA 349

above, more evident beneath; marginal vein about equaling the lateral ones and
arched between them, 0.5-2 mm. from the margin; blades strongly discolorous,
the upper surface lustrous and minutely verruculose (or dotted) with innumerable
pale convex elevations much smaller and more numerous than the few (if any)
translucent glands; lower surface paler, often drying brown or yellow-green, usu-
ally with some translucent convex glands; inflorescence a very short axillary
raceme (or 2-3 racemes superposed), or some of the flowers solitary and opposite
at the lowermost (bracteate) nodes of a new leafy shoot; raceme-axis 1-3 (-5) mm.
long, bearing 1-4 opposite decussate pairs of flowers on slender pedicels (2-) 3-8
(-13) mm. long, up to 0.5 mm. thick in fruit; bracts 1 mm. long or less, ovate,
somewhat persistent-indurate; bracteoles persistent, not connate, ascending, some-
what glandular-thickened at base, 1 mm. long or less; buds 2.5-4 mm. long,
pyriform, the campanulate hypanthium usually densely and conspicuously pale-
pubescent, 1-1.5 mm. long; calyx-lobes in unequal pairs, concave in flower and in
fruit, ovate, densely silky-pubescent near the apices and sometimes on as much
as the distal one-third of both surfaces, but otherwise merely strigose without and
glabrous within, the larger ones 1.5-2.7 mm. long, 1.5-2 mm. wide; disk 1.5-2 mm.
wide; style (4-) 5-6 mm. long; stamens 75-100, about as long as the style, the
anthers up to 0.7 mm. long; petals white, suborbicular, 2.5-5 mm. long; ovary
bilocular, the ovules about 8 (5-10) in a compact group in each locule; fruit glo-
bose to ovoid or pyriform, probably black, tipped by the persistently concave
calyx lobes.

Represented in Guatemala and British Honduras by two vari-
eties distinguished as follows:

Leaves mostly 4.5-6.5 cm. long, acuminate; petioles 5-8 mm. long; fruit 7-8
mm. in diameter, as far as known not stipitate var. laevis.

Leaves mostly 5 cm. long or less, if longer often rounded or broadly and ob-
scurely acuminate at tip; petioles 2.5-5 mm. long; fruit 5-6 (-8) mm.
in diameter, usually contracted at base to a slender stipe 1-2 mm. long.

var. Gaumeri.

Eugenia laevis Berg, var. laevis. E. subverticillaris Berg, Lin-
naea 29: 235. 1858. E. caldphila Lundell, Wrightia 3: 11. 1961.

Well-drained upland forest of Brosimum (ramonal) ; Pete"n (Dos
Lagunas, E. Contreras 1541, type of E. caldphila). Yucatan; His-
paniola, the type region.

Leaves ovate or ovate-elliptic, 2.5-4 cm. wide, about 1.6-2 times as long as
wide, the tip bluntly acuminate or sometimes obtuse.

The only known Guatemalan collection is the type of Eugenia
caldphila. This is a specimen in bud, with a few open flowers. It
is only sparingly pubescent in the inflorescence, but otherwise agrees
precisely with specimens from Yucatan and from Hispaniola.

Eugenia laevis Berg, var. Gaumeri (Standl.) McVaugh, Fieldi-
ana, Bot. 29: 469. 1963. E. Gaumeri Standl. Field Mus. Bot. 8: 28.

350 FIELDIANA: BOTANY, VOLUME 24

1930. E.Lundellii Standl. Carneg. Inst. Wash. Publ. 461: 76. 1935.
Guayabillo.

Dry rocky thickets or scrub forest with Haematoxylum (tintal),
sometimes in pine-oak forest, mostly at low elevations; Pet&n; Baja
Verapaz (north of Santa Rosa, 1500 meters, Standley 69775) . Yuca-
tan, the type from Kancabtsonot, Gaumer 23843; Campeche (Tux-
pena, Lundell 1130, the type of E. Lundettii); British Honduras;
Honduras.

Leaves elliptic or lance-ovate, to elliptic-lanceolate, oblanceolate or obovate,
1-3 (-4) cm. wide, 1.5-2.3 (-2.7) times as long as wide, obtusely pointed, or
rounded, or broadly and bluntly acuminate.

For a note on the taxonomy of this variety, see Fieldiana, Bot.
29: 469. 1963. The cited collection from Baja Verapaz is sterile,
but agrees well in pubescence and in leaf -form with Yucatan material.

A very similar species, Eugenia coyolensis Standl. (Field Mus.
Bot. 9: 320. 1940) is known only from the Department of Yoro,
Honduras, the type from Coyoles, Yuncker et al. 8069. This may
not be specifically distinct, but seems superficially different; the
leaves are obovate, rounded or broadly obtuse at apex; the flowers
are somewhat larger than those of E. laevis (the larger calyx-lobes
2.5-3.5 mm. wide, the stamens about 125).

Eugenia lancetillae Standl. Field Mus. Bot. 4: 317. 1929.

Known only from the type, Honduras, Dept. Atlantida, wet for-
est, Lancetilla Valley near Tela, 20-600 meters, Standley 53301.

Tree 6-7 meters high, the branchlets, petioles, inflorescence and veins of the
lower leaf-surface hispidulous with pale short brown hairs; leaves thin, elliptic,
4-8 cm. wide, 12-19 cm. long, 2.5-3 times as long as wide, short-acuminate, nar-
rowed and rounded to the base, the margins finally cuneately decurrent on the
channeled petiole 8 mm. long, 1.5-2 mm. thick; midvein flat above; main lateral
veins about 8 on each side, seen above as pale somewhat depressed convex lines,
prominent beneath, arcuate-ascending, scarcely forming a marginal vein, often
diminishing distally, recurving 5-7 mm. from the margin and joining the next
succeeding veins; intermediate lateral veins scarcely developed, the small veinlets
forming a prominent network; blades probably dull above, paler beneath, the
glands hardly apparent; inflorescence a crowded axillary raceme, the axis in fruit
up to 3 mm. long, 1.5 mm. thick, bearing 1-3 pairs of nearly sessile flowers; bracts
persistent, indurated, broadly rounded, 1 mm. long; bracteoles persistent, distinct,
ovate-orbicular, 1-1.3 mm. broad and long; calyx-lobes in fruit in unequal pairs,
the larger ones up to 2.5-3 mm. wide, 3 mm. long, orbicular-ovate, the shorter
ones deltoid-ovate, 2.5 mm. wide, 2 mm. long; disk hirsute, 2.5 mm. wide; imma-
ture fruit ellipsoid-oval, 1 cm. in diameter, 1.3 cm. long.

McVAUGH: FLORA OF GUATEMALA 351

Eugenia letreroana Lundell, Wrightia 3: 15. 1961.

To be sought in Guatemala; known only from the type collec-
tion, Matuda 4336, from Letrero, near Siltepec, Chiapas, in virgin
forest at 2000 meters elevation.

A tree 3-4 meters high, the inflorescence and young branchlets and petioles
minutely puberulent; leaves narrowly elliptic, 1-1.8 cm. wide, 3-6 cm. long, 2.5-
3.5 times as long as wide, about equally narrowed to the slenderly but bluntly
acuminate tip and to the acute or cuneate base, the margins decurrent on the inner
angles of the deeply channeled petiole 4-6 mm. long; midvein pale, deeply incised
in the plane opaque surface of the blade; lateral veins 5-8 on each side, obscure
above, inconspicuous beneath, connected by irregularly angled arches, the mar-
ginal vein scarcely developed; blades dull dark green and almost featureless above,
when dry strongly discolorous, brown and finely dark-dotted beneath; inflores-
cence a well-developed raceme, the axis 4-9 mm. long, 4-angled and 1 mm. thick
at base, bearing (4-) 6-8 opposite and decussate pairs of flowers on pedicels
1-1.5 mm. long; bracts divaricate, subpersistent, indurate, 0.5 mm. long; bracte-
oles ovate, rounded on the backs, persistent but scarcely connate, 0.5 mm. long;
buds obovoid-pyriform, 2.5 mm. long, the globe of the petals much exceeding the
calyx; hypanthium puberulent, campanulate, 1 mm. long, prominently dotted
with convex yellowish glands; calyx-lobes broadly rounded, the larger ones 1 mm.
wide, 0.7 mm. long; disk 1.3-1.8 mm. wide; style about 4.5 mm. long; stamens
about 75; petals white, suborbicular, 3 mm. long; ovary bilocular, the ovules
about 9 in a compact group in each locule; fruit unknown.

Eugenia Matudae Lundell, Phytologia 1: 218. 1937. Stink-ya
(Huehuetenango, according to Steyermark).

A species of forested mountains, along rivers, to be expected in
Guatemala. Chiapas, the type from Mt. Madre Vieja, E. Matuda
648.

A tree 5-6 meters high, known only from over-mature fruiting specimens, the
whole plant at this stage glabrous; leaves elliptic, about equally narrowed to a
short blunt acumen and an acute base, 6.5-8 cm. wide, 14-18 cm. long, 2.3-2.7
times as long as wide, the margins cuneately decurrent on the channeled petiole
1.3-2 cm. long; midvein rather broadly concave above, the central channel bor-
dered by parallel convex ridges; lateral veins 6-8 on each side, arcuate-ascending,
much stronger than the intermediate ones if any, convex on both sides of the
blade, diminishing distally and arching over 5-6 mm. from the margin to join the
next succeeding ones, the marginal vein not well defined; leaves nearly concolorous,
probably paler beneath, with small convex glands on both surfaces; flowers appar-
ently several together in sessile clusters at leafless nodes on old wood, arising
from a short woody spur probably consisting of the fused axes of 2-3 very short
superposed racemes; pedicels in fruit 1.3-3 cm. long, 1.3-1.7 mm. thick; bracte-
oles persistent under the fruit, distinct, indurated, deltoid, 1.5 mm. long and
wide; fruit glandular-verrucose, glabrous, broadly ellipsoid or obovoid to nearly
globose, rounded at both ends, 1.5-2.3 cm. thick, 2-2.5 cm. long, surmounted

UNIVERSITY OR
ILLINOIS LIBRARY

352 FIELDIANA: BOTANY, VOLUME 24

by the concave, broadly rounded calyx lobes; lobes in unequal pairs, the larger ones
5 mm. wide at base, 6 mm. long; disk about 6 mm. wide.

This species much resembles E. choapamensis Stand!., from which
it may be superficially distinguished by the much wider staminal
disk, the larger fruit, and the smaller number of foliar veins.

The specific epithet was originally spelled Matudai. According
to the International Code of Botanical Nomenclature this is ortho-
graphically incorrect; the preferred spelling is Matudae, regardless
of the sex of the person for whom the species is named.

Eugenia nigrita Lundell, Wrightia 3: 16. 1961.

To be sought in Guatemala; known only from the type locality,
near Siltepec, Chiapas, where collected in advanced forest at 1600
meters elevation (Matuda 5183, the type).

A tree up to 10 meters high, darkening in drying, nearly glabrous, the young
branchlets and vegetative buds and the inflorescence more or less covered with
rather coarse lustrous simple copper-colored hairs up to 0.5 mm. long or more, the
whole plant soon glabrate; leaves broadly elliptic or elliptic-ovate, 1.5-2.8 cm.
wide, 3-6 cm. long, 2-2.3 times as long as wide, prominently acuminate (the very
tip obtuse), acute to cuneate at base, the margins decurrent on the inner angles of
the deeply channeled petiole 4-7 mm. long; midvein narrowly impressed above;
lateral veins about 5, inconspicuous above and a little more evident beneath,
connected by irregularly arching or branching subordinate veins, the marginal
vein scarcely developed; blades rather thin, darker and often with gland-filled
convexities above, the lower surface paler (brown in drying), eglandular or finely
dark-dotted; inflorescence an abbreviated raceme (or the flowers opposite at the
lowermost [bracteate] nodes of new leafy branchlets), the axis of the raceme often
3-5 mm. long, bearing up to 4 pairs of flowers on pedicels 1.5-2 mm. long; bracts
ovate, indurate, up to 0.8-1 mm. long; bracteoles broadly ovate, divaricate, thick-
ened at base and persistent but scarcely connate, 1 mm. long, coarsely hairy with-
out or at least apically fringed; buds pyriform, 3-4 mm. long including a con-
tracted broadly stipelike base; hypanthium campanulate, 1-1.3 mm. long, gla-
brous or sparingly strigose; calyx-lobes membranous, flat, fringed, hemi-orbicular,
the larger ones 1.5 mm. wide, 1.2 mm. long; disk about 1.5 mm. wide; style 5-6
mm. long; stamens about 75, about as long as the style, the anthers about 0.5 mm.
long; petals white, suborbicular, 3 mm. long; ovary bilocular, the ovules about
15 in a compact group in each locule; fruit globose or oblate, probably black,
1.3 cm. in diameter.

Eugenia octopleura Krug & Urb. ex Urb. Bot. Jahrb. 19: 653.
1895. E. Doubledayi Standl. Journ. Arnold Arb. 11: 36. 1930 (Hon-
duras, the type from Siguatepeque, Standley 56063). E. Koepperi
Standl. Field Mus. Bot. 9: 320. 1940 (Honduras, the type from Mt.
Cangrejal near La Ceiba, Yuncker et ol. 8400).

McVAUGH: FLORA OF GUATEMALA 353

Thickets and river banks, or wet forest, mostly at low elevations,
in Honduras up to 1500 meters; Pete"n (Dolores, km. 85, Machaquila
Road, E. Contreras 2953). Honduras; Costa Rica; Lesser Antilles.

Tree or shrub 3-6 meters tall, up to 20 cm. in diameter, appressed-pubescent
but glabrate, the inflorescence densely and closely beset with yellowish-white to
pale brown or rufous dibrachiate hairs, the branchlets and petioles and lower
leaf-surfaces with very fine numerous pale hairs; leaves elliptic-ovate, 3-5 (-6.5)
cm. wide, 7-10 (-15) cm. long, 2-3 times as long as wide; blades prominently acu-
minate at tip, acute or somewhat rounded at base, the margins somewhat cuneately
decurrent on the inner angles of the channeled petiole 8-15 mm. long, 1-1.5 mm.
thick; midvein impressed above; lateral veins 8-12 pairs, ascending, rather incon-
spicuous above, more prominent beneath, stronger than the intermediate veins
if any, distally forming strong irregular interconnecting arches 1.5-5 (-10) mm.
from the margins; glands inconspicuous; inflorescences rather stout clustered axil-
lary racemes (usually 2-3 superposed) 1.5-2.5 cm. long, the axis 1-1.8 cm. long,
bearing 3-4 opposite and decussate pairs of flowers; terminal flower abortive;
pedicels 3-9 mm. long, 0.5 mm. thick in flower; bracts ovate, acute, 1 mm. long,
rounded on the backs, becoming dark and indurated and probably falling after
anthesis; bracteoles similar, distinct, persistent through anthesis; buds 3-5 mm.
long, pyriform; hypanthium ridged at anthesis when dry, conic, 1.5-2 mm. long;
calyx-lobes in unequal pairs, broadly rounded, glabrate without, thickly appressed
pubescent with rufous hairs within, the smaller (outer) pair 1.5-2 mm. long, 2.5
mm. wide, the larger pair 2.5-3 mm. long, 3.5-4 mm. wide; disk 2.5-3 mm. wide,
with a few long hairs about the style, the staminal ring somewhat pitted; style
5-7 mm. long; stamens 125-150, up to 5 mm. long, the anthers 0.6 mm. long;
petals white, 4-5 mm. long; ovary bilocular, the ovules ca. 10-16 in each locule,
radiating in a subcapitate group from the central dissepiment; immature fruit
oblong or obo void-oblong, crowned by the persistent calyx-lobes, obtuse at apex,
narrowed a little toward the base, 8-ribbed, 6-7 mm. thick, 16-18 mm. long,
minutely pale pubescent, subtended by the persistent divaricate bracteoles.

Evidently very closely related to E. pleurocarpa Standl., of west-
ern Mexico, but differing in the relatively narrower and more prom-
inently acuminate leaves, the more closely appressed pubescence, the
more strongly proliferous inflorescence.

Eugenia Oerstedeana Berg, Linnaea 27: 285. 1856. E. vin-
centina Krug & Urb. ex Urb. Bot. Jahrb. 19: 621. 1895. E. Conzattii
Standl. Contr. U. S. Nat. Herb. 23: 1041. 1924. E. cocquericotensis
Lundell, Bull. Torrey Club 64: 554. 1937. E. petenensis Lundell,
Bull. Torrey Club 69: 397. 1942. E. eutenuipes Lundell, Wrightia 2:
106. 1960. "Jolteillo" (Pete"n, according to Lundell).

Moist or wet thickets or mixed forest, sometimes in pine forest,
mostly 1200 meters or less, most frequent at rather low elevations;
Pete"n (La Libertad, Lundell 3746, type of E. petenensis); Izabal;
Santa Rosa; Escuintla; Guatemala; Sacatepe"quez; Chimaltenango;

354 FIELDIANA: BOTANY, VOLUME 24

Solola; Retalhuleu; Quezaltenango; Huehuetenango. Mexico (north
to Nayarit and northern Veracruz; Oaxaca, Conzatti et al. 3113, type
of E. Conzattii; Tabasco; Chiapas, Siltepec, Matuda 411, type of
E. eutenuipes) ; British Honduras (Little Cocquericot, El Cayo Dis-
trict, C. L. Lundell 4090, type of E. cocquericotensis) ; Central Amer-
ica as far east as Costa Rica, the type from Nicaragua, between
Sapoa and Tortuga, Oersted; West Indies (St. Vincent).

A shrub or small tree 3-6 meters high, up to 15 cm. in diameter; leaves nearly
glabrous from the first, but the vegetative buds and youngest growth appressed-
pubescent with evanescent partly dibrachiate coppery hairs up to 0.4 mm. long,
and the petioles and a narrow zone on the upper leaf-surface along the midvein
persistently hispidulous with sharp erect colorless hairs 0.1-0.2 mm. long; branch-
lets similarly hispidulous, often glabrate; pedicels and flowers nearly glabrous;
leaves chartaceous or membranaceous, ovate to elliptic or broadly lanceolate,
(1.8-) 2.5-4 cm. wide, 4-9 cm. long, (1.75-) 2-3 (-3.5) times as long as wide, the
apex acuminate, the base rounded or acute, sometimes slightly inaequilateral,
the margins more or less cuneately decurrent on the deeply channeled petiole
4-7 mm. long; midvein deeply and narrowly impressed above, often elevated its
whole diameter beneath, and bearing a few coppery dibrachiate hairs; lateral
veins slender, 6-10 pairs in addition to some intermediate ones, curving-ascending,
more conspicuous and convex below, seen above as a fine and often slightly in-
cised line; marginal vein about equaling the laterals and arched between them,
well separated (2-5 mm.) from the margin, and paralleled by a thinner continu-
ous, submarginal vein near the margin; surfaces of the blade dull, pale or brownish
green and nearly concolorous when dry, the glands scarcely apparent, or a few
scattered on the lower surface; inflorescence variable, the flowers often solitary in
the axils of ordinary leaves or of small lanceolate scarious deciduous bracts, or
about equally often in slender axillary racemes 1-2.5 (-4) cm. long, less than
1 mm. thick at base, bearing up to 5-6 opposite or subopposite and decussate
pairs of flowers on very slender, sometimes hispidulous pedicels 6-10 (-25) mm.
long, about 0.3 mm. thick; terminal flower often present; bracteoles scarious,
thin-margined, persistent, with prominent convex glands near base, broadly
ovate-cordate but not contiguous, up to 1.5 mm. long and wide; buds 4-4.5 mm.
long, glabrous or essentially so except the ciliate perianth-segments; hypanthium
cup-shaped; calyx-lobes membranaceous, scarious, prominently gland-dotted, in
slightly unequal pairs, broadly rounded, up to 1.5-2 mm. long, 2-2.5 mm. wide;
disk 1.5-2.5 mm. wide; style 5 mm. long; stamens 60-75, about as long as the
style; anthers 0.5 mm. long; petals white, prominently glandular, 4 mm. long;
ovary bilocular, the ovules 2-6 in each locule, collateral or two-ranked; fruit ses-
sile, globose, glandular-roughened, 8-10 mm. in diameter, tipped by the pale mem-
branaceous calyx-lobes; seed usually 1, the embryo undivided.

One of the commonest species of Eugenia in Guatemala, widely
distributed and uniform in most of its characters. It has been re-
ported from Guatemala, and from British Honduras, as E. vincentina
Krug & Urban. Taxonomically E. Oerstedeana appears to be closely
allied to Eugenia florida DC., a widely distributed lowland species

FIG. 51. Eugenia Oerstedeana. A, flowering branchlet; X ca. 0.8. In this spe-
cies the inflorescence may be an elongate raceme, a 3-flowered raceme with the
terminal flower present (simulating a dichasium), or a leafy branch having the
individual flowers subtended by small bracts or even by ordinary leaves. Each
node in the inflorescence is at right angles to the one below it, and a similar arrange-
ment is carried through into the flower: B, Lateral view of flower, cut away to
show the orientation of the ovules. The bracteoles beneath the flower alternate
with the outer (shorter) calyx-lobes, and thus are opposite the inner pair of lobes.
C, Flower seen from below, the hypanthium and ovary cut away to show the orien-
tation of the locules of the ovary. The locules are opposite the outer calyx-lobes,
hence alternate with the inner pair of calyx-lobes and with the bracteoles.

355

356 FIELDIANA: BOTANY, VOLUME 24

of northern South America and Panama. In E. florida most of the
flowers are in racemes bearing 4-5 (or up to 10) pairs of flowers on
short pedicels 3-4 mm. long or less. The type of Eugenia cocquerico-
tensis answers this description; it is among the few Central American
specimens of E. Oerstedeana that closely resemble E. florida. For
notes on the taxonomy of E. Oerstedeana, see Fieldiana, Bot. 29:
446-448. 1963.

Eugenia origanoides Berg, Linnaea 29: 229. 1858. E. conglo-
bata Sesse" & Moc. Naturaleza II. 1: app. 83. 1888. E. Banghamii
Standl. Journ. Arnold Arb. 11: 125. 1930.

Moist or wet, mixed forest or thickets, frequently in pine forest,
often on limestone, sometimes in rather dry places, 1500 meters or
less; Alta Verapaz; Izabal; Zacapa; Retalhuleu; Huehuetenango.
Eastern Mexico (north to Tamaulipas, Palmer 295 in 1910; Papantla,
Veracruz, Karwinsky 238, the type) . British Honduras; Costa Rica;
Panama; Colombia (Triand).

Tree or shrub up to 6 meters high, usually densely tawny-hirsutulous on
branchlets and inflorescence, and somewhat less densely on the lower leaf-surfaces,
with erect or curved lustrous hairs about 0.5-1 mm. long; leaves elliptic, (1-)
2-3 (-6) cm. wide, (3-) 4-10 cm. long, (1.6-) 2-3 times as long as wide, abruptly
or gradually narrowed to the pointed or somewhat bluntly acuminate tip, nar-
rowed below the middle and rounded or often acute at base, the margins passing
abruptly into the sulcate hirsute petiole 3-4 mm. long, 1-2 mm. thick including
the hairs; midvein impressed and persistently hirsutulous above, prominent be-
neath; lateral veins about 6-10 pairs with some intermediate veins almost as
strong, slightly elevated in drying but inconspicuous or scarcely apparent above,
evident beneath, nearly straight and passing directly into the marginal vein,
which is about as strong as the laterals and arched between them, 2.5-5 (-8) mm.
from the margin; blades dark in drying, strigose and often otherwise almost fea-
tureless above, non-glandular, paler beneath and sometimes finely dark-dotted;
flowers closely glomerate, the inflorescence a group of abbreviated axillary ra-
cemes; axis of each raceme up to 4 (-7) mm. long, bearing up to 5 approximate
and decussate pairs of flowers on pedicels 1-1.5 mm. long or less; bracts rounded-
deltoid, about 0.7 mm. long and wide, glabrous on the inner surface, the outer
thickly velutinous with hairs 0.5 mm. long; bracteoles persistent, distinct, ovate,
acute, setose like the bracts, 0.5-1 mm. long; buds ca. 2.5 mm. long; hypanthium
subglobose, about 1-1.3 mm. long and wide, sparingly to very densely pilose;
calyx-lobes 4, membranous, glabrous or sparingly setose, long-ciliate, gland-dotted,
rounded-ovate, about as long as the hypanthium, 1-2 mm. wide and long; disk
flat, glabrous or hairy at the center, about 1-1.5 mm. wide; style 5.5-7 mm. long;
stamens about 35-60, up to 5-6 mm. long, the anthers 0.2-0.4 mm. long; petals
obovate, glabrous, 2.5 mm. wide, 4 mm. long; ovary bilocular, the ovules 4-5 or
up to 9-10 in each locule, radially attached to a short placenta near the center of
the central partition; fruit subglobose, 6-8 mm. in diameter, first red then finally
black, sparingly pubescent, finally glabrate.

McVAUGH: FLORA OF GUATEMALA 357

An imperfectly known species, Eugenia rubella Lundell, Wrightia
3: 18. 1961 (the type from Pete"n, Dolores, E. Contreras 2559) closely
resembles E. origanoides except that the pedicels are up to 4 mm.
long. Similar plants are known from the llanos of the Isthmus of
Tehuantepec, in Oaxaca. These specimens may represent some ab-
normality in E. origanoides or may perhaps be of hybrid origin.

Eugenia ovandensis Lundell, Wrightia 3: 17. 1961.

To be sought in Guatemala; known only from the type collection,
Matuda 1834, from Mt. Ovando, Chiapas.

A small tree, the branchlets, petioles and inflorescence hispidulous-puberulent
with small erect hairs; leaves elliptic or elliptic-ovate, 2-3 cm. wide, 4-6 cm. long,
2-2.3 times as long as wide, bluntly acuminate, acute or rounded at base, the
margins decurrent on the channeled petiole 4-6 mm. long; midvein channeled
above in the plane surface of the blade; lateral veins 5-8 on each side, incon-
spicuous on both surfaces, connected by irregularly angled arches 2-4 mm. from
the margin but a marginal vein scarcely developed; somewhat immature foliage
darkening in drying, the leaves glossy above, nearly concolorous, dull beneath,
the internal glands apparent as convexities on both surfaces; inflorescence an
abbreviated raceme (or the flowers opposite at the lowermost [bracteate] nodes
of new leafy branchlets), the axis of the raceme up to 4-5 mm. long, bearing up
to 4 pairs of flowers on slender pedicels 3-5 mm. long; bracts ovate, indurate,
0.5 mm. long; bracteoles ovate, divaricate, rounded on the backs, persistent but
scarcely connate, 0.5 mm. long, forming a 2-lobed involucre 1 mm. wide; buds
pyriform, 3.5 mm. long; hypanthium hispidulous like the pedicels, turbinate-
campanulate, stipitate and standing above the bracteoles, the expanded part
1 mm. long; calyx-lobes in unequal pairs, broad and low, often partly reflexed
at anthesis, glabrous within except near the ciliate margin, the smaller ones 1 mm.
wide, the larger ones 1.5 mm. wide, 1 mm. long; disk 1.3 mm. wide; styles 5-6 mm.
long, puberulent below the middle; stamens 60-75, up to 5 mm. long, the anthers
0.5 mm. long; petals white, obovate, 2.5 mm. long; ovary bilocular, the ovules
about 4 in each locule; fruit unknown.

Eugenia pachychlamys Bonn. Sm. Bot. Gaz. 27: 333. 1899.
E. chiquimulana Standl. & Steyerm. Field Mus. Bot. 22: 358. 1940
(type from summit of Volcan de Quezaltepeque, Steyermark 31453).
GuaydbiUo.

Moist or wet, broad-leaf forests, brushy hillsides, dry cleared
slopes, 1100-2150 meters; Chiquimula; Jalapa; Santa Rosa; Guate-
mala (type from Frai janes, Heyde & Lux 3061). El Salvador (Dept.
Chalatenango, Tucker 1204).

A compact shrub 1-2 meters high, or a tree up to 6 meters, densely rufous-
tomentose, the branchlets, petioles, inflorescences and youngest leaves quite ob-
scured by a mat of contorted hairs up to 1 mm. thick; leaves elliptic to ovate or
obovate, about equally narrowed to both ends, broadly short-acuminate, rounded

358 FIELDIANA: BOTANY, VOLUME 24

or acute at base, 2.5-5 cm. wide, 5-12 cm. long, 2-2.7 times as long as wide, the
margins passing abruptly into the petiole 10-15 mm. long and 2-2.5 mm. thick
including the hairs; midvein impressed above, tomentose; lateral veins 7-9 on
each side, slightly elevated on both sides when dry; marginal vein weaker than
the laterals, forming a series of angular arches 2-6 (-10) mm. from the margin;
blades opaque, dark green and lustrous above, reticulate-veiny, beneath paler
("silver green" according to Steyermark) and drying brown, tardily glabrescent
and finely gland-dotted; flowers large and nearly sessile in crowded axillary glom-
erules, usually one pair only on an abortive axis 1-5 mm. long; bracteoles ovate,
3-5 mm. long, deciduous at an thesis; buds 8-10 mm. long, pyriform; calyx-lobes
ovate-orbicular, 6-8 mm. long and wide, tomentose on both sides; disk 5 mm.
wide, tomentose at the center; style 12 mm. long; stamens about 200, up to about
12 mm. long; petals nearly glabrous, white; ovary bilocular, the ovules in a com-
pact group of about 10 in each locule; fruit hard, tomentose, globose to oval or
obovoid, 1.7-2.5 cm. in diameter, 2-3 cm. long (in 2-seeded fruits sometimes
didymous, 4 cm. wide).

In the protologue the number of the type collection was cited as
3961, but on the labels it is 3061.

Fine specimens of this species exist among the collections of
Sess£ & Mocino; these were presumably collected by Mocino dur-
ing his long sojourn in Guatemala from about 1796 to 1798 (cf.
Chronica Botanica 11: 37-38. 1947), and so must be among the old-
est of all known specimens of Guatemalan plants.

Eugenia papalensis Standl. & Steyerm. Field Mus. Bot. 23:
130. 1944. Guayabillo.

Known only from the type, Dept. Huehuetenango, Cumbre Papal,
between the summit and La Libertad, 1800-3000 meters, Steyermark
50964.

A tree 10-11 meters high, glabrate at maturity but the mature branchlets
hispid with pale coarse erect hairs up to 0.5 mm. long, the young herbage and
inflorescence probably hispidulous or appressed-pubescent with pale or reddish
hairs; leaves when dry rigid and leathery, broadly elliptic-ovate, 6-8 cm. wide,
12-15 cm. long, 2-2.4 times as long as wide, apparently blunt-tipped, rounded at
base, the thick revolute margins abruptly decurrent on the sparingly hispid chan-
neled petiole 12-15 mm. long, 2-2.5 mm. thick; midvein concave and somewhat
depressed above, sometimes persistently hispidulous; lateral veins about 10 on
each side in addition to some intermediate ones, seen as pale lines above, more
prominent beneath; marginal vein about equaling the lateral ones and arched
between them, 3-4 mm. from the margin; blades "deep green above," paler be-
neath, the glands inconspicuous; flowers probably in sessile axillary clusters, not
seen; pedicels in fruit 1 mm. long and 2 mm. thick; bracteoles persistent, broadly
rounded, hispid, 2 mm. wide, 1 mm. high; fruit "pale yellow," glandular- verru cose,
"bitter," globose or a little elongate, 1.7-2.2 cm. in diameter, about 2 cm. long,
crowned by the subpersistent calyx-lobes, these finely pubescent inside, the larger
ones 3-4 mm. long and wide; disk 3-3.5 mm. wide.

McVAUGH: FLORA OF GUATEMALA 359

Eugenia Percivalii Lundell, Wrightia 2: 124. 1961.

British Honduras: Toledo District, Edwards Road beyond Co-
lumbia, in high ridge, hilltop near Central Camp, P. H. Gentle 7332,
type. Known only from the type collection.

A tree 20 cm. in diameter, nearly glabrous, the inflorescence minutely puber-
ulent with erect or ascending pale hairs 0.1 mm. long or less; leaves coriaceous,
elliptic, 2.5-5.5 cm. wide, 8-13 cm. long, 2-3 times as long as wide, about equally
narrowed to a blunt or sometimes obscurely acuminate tip and to the rounded or
acute base, the margins cuneately decurrent on the channeled petiole 8-10 mm.
long, 1.5-2 mm. thick; midvein nearly flat or sulcate above; lateral veins about
10 pairs, apparent on both sides of dried leaves but inconspicuous, a little stronger
than the intermediate ones; marginal veins about equaling the lateral ones and
arched between them, 1.5-2.5 mm. from the margin; blades smooth and glossy
green above, paler and finely dark-dotted beneath; flowers clustered in the axils,
the inflorescence a very short axillary raceme (sometimes a second raceme super-
posed), the angular axis 1 mm. thick at base, 1-3 mm. long, bearing 1-4 approxi-
mate, decussate and opposite pairs of flowers on pedicels 1-2 mm. long, 0.5 mm.
thick, the terminal flower abortive; bracts scarious, persistent, ovate, broad-based,
0.7 mm. long; bracteoles persistent, ovate, somewhat flaring, 0.5-0.6 mm. long,
not coherent, thickened at base and convex dorsally, puberulent; buds [about
3 mm. long], pyriform, the campanulate hypanthium puberulent; calyx-lobes 4,
in unequal pairs, concave, broadly rounded, sparingly puberulent without, mi-
nutely appressed-puberulent within, the larger ones up to 3 mm. wide, 2.5 mm.
long; disk 2-2.5 mm. wide, bearing a few hairs around the base of the style, the
staminal ring somewhat bristly; style about 6 mm. long; stamens about 100;
petals white; ovary bilocular, the ovules 5-6 in each locule, in a compact group
on the central dissepiment; fruit unknown.

Eugenia Rekoi Standl. Contr. U. S. Nat. Herb. 23: 1044. 1924.

Deciduous forest near the Pacific Coast, rocky headlands, low
wooded hills, thickets near the beach, 300 meters or less; Quezalte-
nango (Caballo Blanco, J. D. Smith 1485). Mexico (Sinaloa to
Oaxaca, the type from Cafetal Apango [Cerro Huatulco], B. P. Reko
3356).

A tree up to 10 meters high, 15-20 cm. in diameter, the inflorescence and
young growth densely appressed-pubescent with coppery dibrachiate hairs mostly
0.2-0.3 mm. long, the youngest branchlets and foliage often reddened by the hairs,
the leaves soon glabrate or rather persistently appressed-pubescent with nearly
colorless hairs; blades broadly elliptic, varying to ovate or obovate, 2-4 cm. wide,
3-5 (-7) cm. long, 1.6-2.5 times as long as wide, the tip prominently or obscurely
acuminate or on short branchlets obtuse, the base cuneate or somewhat rounded,
the margins always cuneately long-decurrent on the dilated summit of the petiole
4-6 mm. long; midvein sulcate above; lateral veins 5-7 on each side, ascending,
obscure above, evident beneath, connected by irregularly angled arches 2-3 mm.
from the margin, the marginal vein often not well developed; blades dark green
and glossy and usually minutely glandular- verruculose above, paler and finely

360 FIELDIANA: BOTANY, VOLUME 24

gland-dotted beneath; inflorescence a very short axillary raceme (usually with
1-2 additional shorter racemes superposed), the axis up to 4 mm. long with 1-3
pairs of flowers on pedicels 1.5-3 (-5) mm. long; bracts ovate, 0.5 mm. long;
bracteoles persistent, convex, distinct, ovate-triangular, 0.5-0.7 mm. long; buds
pyriform, about 2 (-3) mm. long; hypanthium campanulate or obovoid, 1 (-1.5)
mm. long; calyx-lobes broadly rounded, membranous, appressed-pubescent on
both surfaces, the larger ones 1.5-2 mm. long and wide; disk 1.3-1.7 (-2) mm.
wide; style 4-4.5 mm. long; stamens about 50, about as long as the style; petals
white, up to 3 mm. long; ovary bilocular, the ovules 6-8 in each locule; fruit gla-
brous, glandular-verrucose, black, oblong or narrowly obovoid, obtuse at both
ends, 7-8 mm. in diameter, 9-13 mm. long, with very thin flesh.

This species is rather abundant and uniform along the Mexican
coast from Oaxaca westward. The single known Guatemalan col-
lection seems to have thinner and relatively narrower leaves and
somewhat longer pedicels than most Mexican specimens, but prob-
ably belongs to the same species.

Eugenia rhombea (Berg) Krug & Urb. ex Urb. Bot. Jahrb. 19:
644. 1895. E. foetida y rhombea Berg, Linnaea 27: 212. 1856. ?E. fis-
calensis Bonn. Sm. Bot. Gaz. 54: 235. 1912. ?E. leptopa Lundell,
Bull. Torrey Club 69: 396. 1942.

In deciduous forest and scrub, at or near sea level, or on rocky
hillsides, 700-1500 meters; Pete'n; Baja Verapaz; Guatemala (Fiscal,
C. C. Deam 6226, the type of E. fiscalensis) ; Huehuetenango; Quiche".
Veracruz; Chiapas; Yucatan (Lundell 7869, type of E. leptopa);
Honduras; southern Florida; Bahamas; Greater Antilles.

Shrub or treelike, up to 3 meters high, glabrous except for a few reddish hairs
on the vegetative buds, the bristly hairs of the staminal ring, and the cilia of the
bracts and flower-parts; leaves rigidly coriaceous, ovate, 1.5-3.5 cm. wide, 3-6 cm.
long, 1.5-2 times as long as wide, narrowed from the middle or below to a narrow
but rounded acute or acuminate tip, rounded or sometimes acute at base, the
heavy cartilaginous margins decurrent on the inner angles of the sulcate petiole
(1.5-) 3-7 mm. long, up to 1.2 mm. in diameter; midvein impressed above, or
sometimes nearly flat; lateral veins (4-) 6-10 pairs in addition to some interme-
diate ones, obscure above, evident beneath; marginal vein equaling or somewhat
more conspicuous than the laterals and irregularly arched between them, 1-3 mm.
from the margin; leaves dull green above, paler and often drying pale brown
beneath; glands scarcely visible above, evident beneath; inflorescence axillary,
actually racemose but the axis ordinarily not elongating (up to 1-2 mm. long)
and the 2-4 (-6) flowers seemingly arising directly from the axil [or 1-2 pairs of
flowers at the lowermost (bracteate) nodes of a new leafy shoot]; pedicels slender,
6-15 mm. long, up to 0.5 mm. thick; bracts scarious, deciduous, up to 1.5 mm.
long, the indurated bases persistent; bracteoles broadly or narrowly ovate, scari-
ous, 0.5-0.8 mm. long, ciliate, persistent or subpersistent; hypanthium hemi-
spheric, or in anthesis substipitate, in post-anthesis soon oblate and separated by
a narrow neck from the conspicuous green erect concave calyx-lobes; lobes in

McVAUGH: FLORA OF GUATEMALA 361

unequal pairs, rounded, broader than long, up to 3.5 mm. wide, ciliate; disk 2.5-
3.5 mm. wide, the staminal ring hairy, the margins nearly flat but the calyx-lobes
at first somewhat reflexed; style 4.5-5.5 mm. long; stamens about 80, up to
4-5 mm. long, the anthers 0.7 mm. long; petals ciliate, obovate, 4 mm. long;
ovary bilocular, the ovules 15-17 in each locule, radiating in a subcapitate group
from the central dissepiment; fruit black, globose or oblate, somewhat glandular-
roughened, 5-6 mm. long, 7-8 mm. broad.

The plant described as E. fiscalensis Bonn. Sm. is an abundant
shrub, according to notes by Standley, on the very dry rocky hill-
sides about Fiscal, the type locality; it is known from the Sierra de
las Minas opposite El Rancho, Baja Verapaz (Kellerman 7894), and
it has also been collected in sterile condition in the Departments of
Baja Verapaz and Quiche", and in Honduras. It is poorly known,
but on the basis of the available material almost impossible to sep-
arate from E. rhombea. In E. fiscalensis the leaves are usually smaller
(2-4 cm. long) and often obtuse rather than acuminate, somewhat
more generally rounded at base than the leaves of typical rhombea,
and a little thinner and more veiny; almost exactly similar leaves
may be found, however, on some West Indian specimens of rhombea.
In the only known flowering specimen of fiscalensis the flowers appear
to be smaller than usual in rhombea, but perhaps not significantly so,
and the calyx-lobes and margins of the disk are reflexed in anthesis.
More field-study is needed to determine the correct status of E. fis-
calensis.

Eugenia riograndis Lundell, Field & Lab. 13: 10. 1945.

In forest on ridges, British Honduras, the type from Toledo
District, Rio Grande, P. H. Gentle 4784. Mexico (Tabasco).

Tree up to 25 cm. in diameter, the branchlets and petioles and the inflores-
cence persistently hispidulous-puberulent; upper leaf -surf ace uniformly and per-
sistently puberulent with minute curved hairs 0.1 mm. long; leaves elliptic,
sometimes broader a little above or below the middle, usually about equally nar-
rowed to the obtusely but narrowly acuminate tip and to the cuneate base, 1-2.5
cm. wide, 4-8 cm. long, 2.5-3.5 times as long as wide, the margins gradually decur-
rent on the very short, ventrally concave or shallowly channeled petiole 2-2.5 mm.
long; midvein flat above, or toward the base shallowly channeled between parallel
elevated ridges; lateral veins 5-7 on each side, obscure above, more evident be-
neath, the marginal vein about as strong as the lateral ones and arched between
them, 0.5-1 mm. from the margin; blades nearly concolorous, a little paler and
sparingly dark-dotted beneath; inflorescence an axillary raceme (or 2-3 racemes
superposed), the axis usually well developed, slender, terete or compressed, 0.5-
0.7 mm. in diameter, up to 8-12 mm. long, bearing 3-4 pairs of flowers on slender
pedicels 4-5.5 mm. long; bracts divaricate, subpersistent, 0.5-1 mm. long; bracte-
oles divaricate, thin, persistent but not connate, broadly ovate, 0.5-0.7 mm. long;
flowers very small, the buds 1 mm. long; hypanthium broadly campanulate, 0.5-

362 FIELDIANA: BOTANY, VOLUME 24

0.7 mm. long in anthesis, minutely puberulent with erect hairs smaller than those
of the pedicels; calyx-lobes rounded-ovate or oblong-ovate, obtuse, membranous,
finely ciliate but otherwise essentially glabrous, strongly recurved at base but the
concave tips becoming erect, the larger lobes 1.3-1.5 mm. long and wide; disk
0.8-1 mm. wide (up to about 1.5 mm. wide in fruit); style 4-4.5 mm. long; sta-
mens 40-60; petals white, 2 mm. long; ovary bilocular, the ovules about 10 in a
compact group in each locule; fruit globose, up to 5-6 mm. in diameter.

This species superficially resembles Eugenia Capuli and E. tika-
lana, both of which grow in the same general area, but E. riograndis
is easily recognized by the unique small hairs of the upper leaf-
surfaces, and by the delicate, small-flowered racemes.

Eugenia rufidula Lundell, Wrightia 2: 209. Sep. 1961. Cacho
de venado; small-leaf turtle bone.

Known only from the type, from high ridge, on bank of creek,
Edwards Road beyond Columbia, Toledo District, British Honduras,
P. H. Gentle 6385.

A tree 7.5 cm. in diameter, the branchlets, petioles and main veins of the
lower leaf -surf ace hirsutulous with pale reddish hairs; inflorescence (when the
fruit is immature) with shorter and coarser pubescence, the pedicels merely mi-
nutely hispidulous or puberulent; leaves elliptic, about equally narrowed to the
bluntly acuminate tip and the acute base, 0.7-1.4 cm. wide, 2.5-5 cm. long,
about 3 times as long as wide, the margins somewhat decurrent on the terete peti-
ole 2-3 mm. long; mid vein flat or convex above, somewhat persistently hispid-
ulous; lateral veins 5-6 on each side, obscure above, more evident beneath; mar-
ginal vein about equaling the lateral ones and arched between them, 0.5-1 mm.
from the margin; blades discolorous in drying, the upper surface opaque, blacken-
ing, the lower green, dotted with convex glands; inflorescence an axillary raceme
(or 2 racemes superposed), the axis 2-6 mm. long, bearing up to 5 pairs of flowers
on pedicels 1.5-2 mm. long; bracts divaricate, 0.5-1 mm. long, subpersistent;
bracteoles membranous, broadly rounded, persistent but not connate, about 0.5
mm. long and wide, finally explanate somewhat below the immature fruit; buds
and flowers not seen; calyx-lobes membranous, prominently gland-dotted, erect
and concave on the immature fruit, finely ciliate, sparingly puberulent without,
glabrous within, the outer ones 1 mm. wide, 0.7 mm. long, the inner about 1 mm.
long and wide; disk 1-1.2 mm. wide; immature fruit globose, 2-3.5 mm. in diameter.

Eugenia salamensis Donn. Sm. Bot. Gaz. 27: 333. 1899, var.
sal a men sis. E. tomentulosa Standl. Contr. U. S. Nat. Herb. 23:
1043. 1924. E. Mexiae Standl. Field Mus. Bot. 4: 243. 1929.

Deciduous woodlands, often in ravines, mountainsides, often at
low elevations, 1500 meters or less; Salama (Llano Grande, Seler
2445, the type); Guatemala; Quiche"; Jalapa. Mexico (states bor-
dering the Pacific Ocean, Michoacan to Sinaloa).

McVAUGH: FLORA OF GUATEMALA 363

Tree 3-5 meters high, loosely appressed-tomentose, the lower leaf-surfaces
thinly pubescent to (usually) tomentose with yellowish- or grayish-white some-
what contorted dibrachiate hairs up to 0.5 mm. long; tomentum of the inflores-
cence and young growth yellowish- white to pale reddish-brown; leaves glabrate
above, usually permanently pubescent beneath; blades elliptic-oblong, varying to
rhombic-lanceolate or obovate, 3-7 cm. wide, 6-12 (-16) cm. long, 2-3 times as
long as wide, rounded to obtuse or bluntly short-acuminate at tip, narrowed or
rounded toward the base, and there often subauriculate, the thick revolute mar-
gins abruptly and often unequally rounded or contracted to the inner angles of the
sulcate petiole 4-10 mm. long, 1.5-2 mm. thick; midvein usually depressed above
near base, usually tomentulose, convex or flat, often with narrow elevated median
ridge; lateral veins 8-12 pairs, ascending, stronger than the intermediate ones,
somewhat elevated above and evident in dried leaves, more conspicuous beneath,
diminishing distally, and forming a series of strong arches 1-5 (-10) mm. from the
margin and separated from it by a strong net of minor veins; glands inconspicuous
on both surfaces; inflorescence an axillary raceme 2-5 cm. long (sometimes with
additional subordinate racemes or flowers below the primary ones in the axils),
with 1-3 (rarely -5) opposite or subopposite and decussate pairs of flowers; termi-
nal flower sometimes present; pedicels at anthesis up to 6-8 mm. long, 0.5-0.8 mm.
thick; fruiting pedicels 1-1.5 mm. thick; bracts and bracteoles linear, 2.5-5 mm.
long, densely pubescent, falling at or before anthesis; buds pyriform, 5-6 mm. long,
the conic hypanthium expanded into the large calyx that conceals the opening
corolla; calyx-lobes in 2 unequal pairs, broadly rounded, tomentulose on both
sides, 2.5-4 mm. long and wide, in anthesis persistently concave but spreading
from the base, persistent in fruit; hypanthium soon elongating and becoming ellip-
soid-oblong; disk (3-) 4-5 mm. wide, the staminal ring bristly, the concave center
with some long yellowish hairs; style 7-9 mm. long, sparingly pilose on the lower
half; stamens about 150-200, up to 10 mm. long, the anthers 0.6-0.8 mm. long;
petals white, about 5 mm. long and wide, ciliate; ovary bilocular, the ovules about
15-20 in each locule, in a sub-capitate group arising from the central dissepiment;
fruit first red, finally blue-black with thin flesh, elongate, 10-12 mm. thick, some-
times short-elliptic, little longer than wide, usually 15-20 mm. long, oblong;
seed 1, the testa thick-cartilaginous; embryo undivided.

This species is represented in Central America by two varieties
in addition to the var. salamensis (cf. Fieldiana, Bot. 29: 455-457.
1963). Only the var. Rensoniana (Standl.) McVaugh (Psidium
Rensonianum Standl.) is likely to be found in Guatemala. It is
known from El Salvador and from the Isthmus of Tehuantepec,
Oaxaca, Mexico. It may be recognized by the nearly complete lack
of pubescence on the leaves even as early as flowering time, and by
the thin, appressed (not loosely tomentose) covering of hairs in the
raceme at flowering time. The variety is of somewhat doubtful
status.

Eugenia sasoana Standl. & Steyerm. Field Mus. Bot. 23: 131.
1944.

364 FIELDIANA: BOTANY, VOLUME 24

Known only from the type, Dept. Chiquimula, rocky slopes along
Rio Lucia Saso, 3 miles southeast of Quezaltepeque, 1200-1500 me-
ters, Steyermark 31327.

A shrub 5 meters high, nearly glabrous at maturity, the young branchlets and
inflorescence probably abundantly appressed-pubescent with sessile dibrachiate
hairs, a few pale or coppery hairs 0.2-0.3 mm. long persisting on the leaves, the
nodes of the inflorescence and the base of the calyx; leaves "thin," elliptic-lanceo-
late, 0.7-1.6 cm. wide, 3.5-6.5 cm. long, 3.5-4.5 times as long as wide, about equally
contracted to a narrow blunt or bluntly acuminate tip and an acute base; margins
abruptly decurrent on the strong inner angles of the deeply channeled and wrinkled
pale petiole 3-5 mm. long; midvein a pale flat or concave line in the plane upper
surface; lateral veins 10-12 on each side, inconspicuous above, more evident but
not prominent beneath, straight; marginal vein about equaling the lateral ones
and slightly arched between them, less than 1 mm. from the margin; blades "bright
green above," "pale silvery green" and somewhat gland-dotted beneath; inflores-
cence an axillary raceme about 1.5 cm. long in fruit, the axis 4-angled, 1 mm.
thick at base, bearing 2-3 pairs of flowers, the fruiting pedicels up to 6 mm. long,
1 mm. thick; immature fruit oblong-ellipsoid, obtuse at both ends, up to 7 mm.
thick, 12 mm. long, "yellow-green with rose," topped by the firm erect incurved
calyx-lobes, these 2.5 mm. wide, 3 mm. long, ovate, blunt-tipped, appressed-
pubescent within; disk 2-2.5 mm. wide.

Eugenia savannarum Standl. & Steyerm. Field Mus. Bot. 23:
132. 1944.

Savanna between the base of Cerro Chinaja at Sachaj, and Saca-
cao, Alta Verapaz, 150-180 meters elevation, J. A. Steyermark 45710.
Known only from the type collection.

Tree 8 meters high, the branchlets and inflorescence tomentose with loose
coppery-red simple or irregularly dibrachiate hairs up to 0.5-1 mm. long; leaves
rigidly coriaceous, 4-7 cm. wide, 7-12 cm. long, 2-2.5 times as long as wide, the
tip short-acuminate, the base rounded or the margins nearly straight and meeting
at about right angles, finally cuneately decurrent on the inner angles of the deeply
channeled petiole 10-14 mm. long; midvein deeply and narrowly impressed above;
lateral veins rather close and parallel, inconspicuous above, more evident beneath,
10-15 on each side in addition to some intermediate ones, nearly straight; marginal
vein about as strong as the lateral ones, nearly straight and little arched between
them, 1-1.5 mm. from the margin; leaves hard and lustrous and deep green above,
pale beneath with scattered resinous dots; inflorescence a very short stout raceme
almost obscured by the dense tomentum, the axis up to 4 mm. long and 1.5 mm.
thick including the hair, bearing 1-3 opposite decussate pairs of flowers on tomen-
tose pedicels 3-5 mm. long; bracts broadly ovate, indurate, about 2 mm. long
and wide; bracteoles deciduous at anthesis, thin and scarious, oblong or ovate,
tomentose without, 1-1.3 mm. wide, 2-2.5 mm. long; buds not seen, probably
6-7 mm. long, the broadly campanulate densely tomentose hypanthium 3 mm.
long; calyx-lobes concave, broadly rounded, tomentose on both surfaces, reflexed
in anthesis but the tips erect, the larger pair 4-4.5 mm. wide and about as long;
disk 3.5-4 mm. wide, the staminal ring bristly, and a zone around the base of the

McVAUGH: FLORA OF GUATEMALA 365

style tomentose; style ca. 8 mm. long, glabrous; stamens very numerous (about
200), about as long as the style, the anthers 0.7 mm. long; ovary bilocular, the
ovules 15-20 in a compact group in each locule; fruit not seen.

Eugenia Shookii Lundell, Wrightia 2: 209. 29 Sep. 1961. Guaya-
billo.

High forest and bordering pine forest, Pete"n (the type from
Dolores, E. Contreras 2629). Endemic.

A tree 8-12 meters high, 15-20 cm. in diameter, the branchlets, inflorescence
and young foliage appressed-pubescent with short-stalked dibrachiate coppery-
brown hairs up to 0.5 mm. long; leaves elliptic, 2-3.5 cm. wide, 4-8 cm. long,
2-2.5 times as long as wide, abruptly or gradually acuminate (the tip often slender
but usually obtuse), acute at base, the margins cuneately decurrent on the inner
angles of a rather deeply channeled petiole 6-9 mm. long, 1 mm. thick; midvein
deeply and narrowly impressed above; lateral veins 10-12 on each side including
some intermediate ones, inconspicuous, more evident beneath, ascending; marginal
vein about equaling the lateral ones and arched between them, 1.5-2.5 mm. from
the margin; leaves lustrous and dark green above, eglandular, the lower surface
much paler (at least when dry), finely gland-dotted; inflorescence a short axillary
raceme (a second raceme often superposed), the axis up to 6-11 mm. long, bearing
up to 5 pairs of flowers on pedicels 1.5-3.5 mm. long, these up to 1 mm. thick in
fruit; bracts 0.5-1 mm. long, mostly deciduous; bracteoles lanceolate, scarious,
distinct, pubescent, 0.7 mm. long, subpersistent; buds not seen; hypanthium in
post-anthesis sparingly appressed-pubescent, obovoid, 1.8-2 mm. long, narrowed
to the mouth; calyx-lobes in unequal pairs, appressed-pubescent within at least
near the tips, gland-dotted and nearly glabrous without, reflexed at base but the
tips remaining concave, the larger ones 2-2.5 mm. long, up to 2.5 mm. wide; disk
glabrous, 1.5-2 mm. wide, often quadrate; style not seen; stamens 60-75; petals
(according to Lundell) glabrous, 2.2-2.5 mm. long; ovary bilocular, the ovules
12-15 in each locule; fruit "reddish-black" (according to Contreras), oblong or
ellipsoid, obtuse at both ends, 1-1.3 cm. long, 5-8 mm. in diameter, usually with
one massive elongate seed.

Eugenia tikalana Lundell, Wrightia 2: 210. 1961. Guayabillo;
chilonche.

In primary forest of Achras, Brosimum or Haematoxylum, or in
secondary associations; lowlands. Pete"n, the type from Tikal, C. L.
Lundell 15971. British Honduras; Campeche(?).

A tree up to 12 meters high, the branchlets, petioles and inflorescence minutely
hispidulous-puberulent; leaves membranous, glabrous except for a few hairs above
on the midvein, 1.5-3.5 cm. wide, (3-) 4-7 cm. long, (2-) 2.3-3 times as long as
wide, gradually but prominently acuminate (the very tip obtuse), acute or cuneate
at base, the margins decurrent on the inner angles of the ventrally concave petiole
2-4 mm. long; midvein flat or convex above; lateral veins 6-8 pairs, inconspicuous
but stronger than the intermediate ones between them; marginal vein about equal-
ing the lateral ones and arched between them, 1.5-3 mm. from the margin; glands

366 FIELDIANA: BOTANY, VOLUME 24

evident beneath in young leaves, often obscure on both surfaces when the leaves
mature; inflorescence an abbreviated axillary raceme (2 or 3 racemes usually
superposed), the axis up to about 10 (rarely to 25) mm. long, bearing 3-5 (-6)
opposite decussate pairs of flowers on pedicels 2.5-5 mm. long, 0.2 mm. thick (in
fruit up to 0.5 mm. thick) ; bracts scarious, ovate, 0.5 mm. long; bracteoles quadrate
or suborbicular, 0.3-0.5 mm. long, persistent but not connate, scarious, pale-ciliate,
dotted with large glands; buds 2-2.5 mm. long, pyriform, the campanulate puber-
ulent hypanthium 0.7-1 mm. long, the prominently gland-dotted perianth ex-
panded into a globe above it; calyx-lobes 4, in unequal pairs, nearly glabrous
without, glabrous within, rounded, thicker and gland-dotted near base, reflexed
at anthesis, the larger ones 1.3-1.7 mm. long and wide; disk 1-1.3 mm. wide, the
staminal ring sparingly hairy; style 4-5 mm. long; stamens 45-50, about as long
as the style; petals white; ovary bilocular, the ovules 6-10 in each locule; fruit
globose, soon glabrate, glandular-verrucose, at first turning reddish-yellow, then
purple-black, 9-12 mm. in diameter (occasional forms oval, 1 cm. long, 8 mm.
thick, or transversely oval, didymous, 2-seeded, 1 cm. the longer diameter).

This species is superficially almost indistinguishable from E.
Capuli.

Eugenia toledinensis Lundell, Phytologia 1: 307. 1939.

Known only from the type collection, British Honduras, on creek
bank in forest, ca. 600 meters, Camp 23 of the British Honduras-
Guatemala boundary survey, W. A. Schipp S-644.

A tree 10-11 meters high, 30 cm. in diameter, nearly glabrous at maturity,
the inflorescence and probably also the young branchlets and herbage pale-brown-
tomentose with coarse contorted and partly forked hairs up to 0.5 mm. long;
leaves glabrate, elliptic-oblong, 3-4.5 cm. wide, 10-12 cm. long, 3-3.5 times as
long as wide, about equally narrowed to a short-acuminate tip and an acute base,
the margins inconspicuously if at all decurrent on the stout rimose, nearly terete
petiole 5-6 mm. long, 1.5 mm. thick; midvein convex above, often drying into a
narrow elevated ridge except near the base of the blade; lateral veins 10-12 on
each side, straight, convex above when dry, similar but more evident beneath,
connected by a series of angular asymmetrical arches 3-7 mm. from the margin;
upper surface of blade green and lustrous, the lower probably paler, drying brown;
glands obscure on both sides; flowers apparently glomerate in much abbreviated
axillary racemes, the axis almost none; pedicels in fruit 2-2.5 mm. long and about
as thick; fruit brown-tomentose, subglobose, a little longer than wide, about 2 cm.
long and 1.8 cm. in diameter, the persistent calyx-lobes tomentose on both sides,
concave, broadly rounded, the outer ones about 4 mm. long and wide, the inner
pair about 6 mm. wide, 4 mm. long; disk in fruit about 5.5 mm. wide, the de-
pressed tomentose center 4 mm. wide.

Eugenia Trikii Lundell, Wrightia 2: 211. 29 Sep. 1961.

In climax forest of Brosimum (ramonal), and in lowland scrub
forest with Metopium (chechemal) ; Pete"n (the type from Santa Elena,
E. Contreras 2057). British Honduras (El Cayo, Chanek 191).

McVAUGH: FLORA OF GUATEMALA 367

A treelike shrub, or a tree up to 8 meters high, 8 cm. in diameter, the branch-
lets, petioles and inflorescences appressed-pubescent and more or less whitened
with pale simple hairs up to 0.3 mm. long; leaves elliptic or ovate, soon glabrate
but usually with some fine hairs persistent on the lower surface, 2-3.5 (-5.5) cm.
wide, 5-8 (-11) cm. long, 1.9-2.2 times as long as wide, gradually or abruptly
acuminate (the tip obtuse), rounded to the base, the margins somewhat decurrent
on the inner angles of the rather deeply channeled petiole 6-8 mm. long, 1-1.5 mm.
thick; midvein sulcate above; lateral veins 8-10 on each side in addition to some
intermediate ones, inconspicuous, somewhat elevated on both sides in dried leaves;
marginal vein about equaling the laterals and forming a series of angular arches
between them, 1.5-5 mm. from the margin; blades nearly concolorous when dry,
apparently lustrous above, with minute rather inconspicuous glands beneath;
inflorescence a very short axillary raceme, the axis up to 7 mm. long, bearing up
to 4 pairs of flowers on pedicels up to 3 mm. long; bracts ovate or the upper ones
oblong, sparingly pubescent, 1-2.5 mm. long; bracteoles deltoid-ovate, nearly gla-
brous, distinct, persistent, 1.3 mm. long; buds ca. 3-3.5 mm. long, pyriform;
hypanthium campanulate, canescent, contrasting strongly with the nearly gla-
brous calyx; calyx-lobes in unequal pairs, broadly rounded or the outer ones del-
toid-ovate, the larger ones 2 mm. wide, 1.3 mm. long; disk about 1.5 mm. wide;
style not seen fully developed; stamens about 50, the anthers 0.5 mm. long and
wide; petals white, rotund, ciliate, 2.5 mm. long and wide; ovary bilocular, the
ovules 4 in each locule; fruit fleshy, purple, globose and 8-10 mm. in diameter (or
didymous in two-seeded forms).

Eugenia uniflora L. Sp. PI. 470. 1753. E. Michelii Lam. Encyc.
3 : 203. 1789. Stenocalyx Michelii (Lam.) Berg, Linnaea 27: 310. 1856.
Cereza de Surin&m; pitanga. Surinam cherry.

Cultivated occasionally in Guatemala, especially about Guate-
mala and Coban and in the North Coast, for ornament and for its
edible fruit. Native probably of South America, now planted in
many tropical regions.

A shrub or small tree sometimes 9 meters high but flowering and fruiting when
a shrub of 1-2 meters only, essentially glabrous, with a few coarse reddish hairs
up to 0.5 mm. long on the youngest branchlets and leaves, the bracts and bracte-
oles, calyx-lobes and petals ciliate near the tips; leaves ovate, seldom ovate-elliptic,
1.5-3.8 cm. wide, 2.5-7 cm. long, 1.5-2 times as long as wide, bluntly acuminate
at tip, rounded at base, the margins passing abruptly into the subalate inner angles
of the adaxially concave petiole 3-4 mm. long, 1-1.3 mm. thick; midvein concave
or sulcate above; lateral veins 5-6 pairs, rather prominent beneath and stronger
than the intermediate veins, scarcely diminishing distally, but each arching over
irregularly to join the next, the marginal arches 1-3 (or often 5) mm. from the
margins, with a second series of arches forming a distinct submarginal vein be-
yond them; leaves dark green and glossy above, paler beneath, usually with numer-
ous pale translucent glands apparent on both surfaces; inflorescence a very short
axillary raceme bearing 1-3 approximate decussate pairs of flowers, these sub-
tended by scarious or colored thin bracts 1-2 mm. wide, 3-5 mm. long, and the
whole seeming to consist of a subumbellate cluster of flowers surrounded by an

368 FIELDIANA: BOTANY, VOLUME 24

involucre of bracts; floral axis sometimes abortive, but often produced into a new
leafy shoot developing with or after the flowers, the lowest sterile leaves (as well
as the floral leaves below them) often bractlike and the base of the branchlet
covered by the imbricated bracts; pedicels (5-) 10-18 mm. long, filiform, 0.3 mm.
thick (up to 0.7 mm. in fruit); bracteoles linear or filiform, distinct, 1-2 mm. long,
deciduous before anthesis; buds about 5 mm. long, the short conic hypanthium
1 mm. long, expanded abruptly into the globe of the petals; calyx-lobes somewhat
foliaceous, oblong-elliptic, nearly flat, little imbricated, 1.8-2 mm. wide, 2.5-4 mm.
long, strongly reflexed with the petals and the margins of the disk at anthesis;
disk glabrous, 2 mm. wide; style 4.5-5 mm. long; stamens about 60, up to 7 mm.
long; petals white, obovate, 7-8 mm. long; ovary bilocular, the ovules about 20 in
each locule, in a subcapitate group from the central dissepiment; fruit tomato-red,
very juicy, depressed-globose, 8-costate, 2-3 cm. in diameter.

This species was first described in 1723, from a cultivated plant
grown in a garden at Pisa, Italy, and supposed to have come origi-
nally from Goa, where also it must have been cultivated. It is now
widespread in cultivation in warm regions, but it is interesting to
note that its native region is still in doubt, or has been lost sight of.
In recent floras of tropical America, its range is usually given merely
as "South America," or "tropical America." Berg, in the Flora
Brasiliensis (1857) stated clearly that it was a native of tropical
Brazil, frequent in the province of Rio de Janeiro, and elsewhere
grown in gardens for its edible fruit. In view of its early occurrence
at the Portuguese colony of Goa, it may well have been carried at
an early date from the Portuguese colonies in America to those in
India.

The following notes are by Mr. Standley: Called "Guinda" in
El Salvador. The Surinam cherry is a usually densely branched
shrub of handsome appearance. It makes one of the best ornamental
hedge plants for Central America if kept low and well trimmed, but
if allowed to increase in height it becomes a rather straggling tree.
The fruit is very juicy and has a most pleasant spicy flavor, being
one of the best of all tropical fruits in the opinion of many people.
It makes excellent ice cream, sherbet, and beverages. In Guatemala
it grows well from sea level to an elevation of 1800 meters or more.
It is one of the few fruits of tropical America that have a pleasantly
acidulous flavor suggestive of cherries and other common fruits of
the North.

Eugenia vacana Lundell, Phytologia 1: 308. 1939.

On ridges and hillsides, wet forests, often in secondary forest, at
low elevations; Izabal. British Honduras, the type from Vaca, El
Cayo District, P. H. Gentle 2535.

McVAUGH: FLORA OF GUATEMALA 369

A tree 5-8 meters high, up to 15 cm. in diameter, the young branchlets, foliage
and pedicels minutely hispidulous with persistent, crowded erect sharp hairs up
to 0.2 mm. long, and also loosely pilose with deciduous yellowish-brown or coppery
hollow contorted hairs up to 0.5 mm. long or more; leaves glabrate, the branchlets
and inflorescence persistently hispidulous, a few long hairs sometimes persistent
on the inflorescence; leaves elliptic, (1-) 1.5-3 cm. wide, (2-) 3-6.5 cm. long,
2-2.5 (-3) times as long as wide, abruptly or gradually short-acuminate, the base
acute or cuneate, the margins decurrent on the inner angles of the sulcate petiole
3-5 (-8) mm. long; midvein impressed above; lateral veins 7-10 pairs, ascending
and parallel, passing directly into an arched marginal vein of about the same
thickness as the laterals, 0.5-1.5 mm. from the margin; leaves dark green above,
and nearly featureless, often drying much darker than the dark-dotted lower sur-
face; inflorescence an axillary raceme, the axis up to 3 mm. long, bearing usually
about 3 opposite decussate pairs of flowers on pedicels 4-7 mm. long; bracts
broadly rounded, 0.5-0.8 mm. long and wide; bracteoles ovate, acute, persistent,
0.5-0.8 mm. long, at anthesis loosely pilose; buds ca. 2.5 mm. long, the hypan-
thium 1 mm. long, thinly pilose with contorted hairs, otherwise glabrous; calyx-
lobes deltoid-ovate, thin, loosely spreading-reflexed in anthesis, up to 1.5 mm.
wide at base, 0.8-1.3 mm. long, glabrous within, thinly hairy without, dotted with
large glands; disk somewhat hairy, 1.3-1.5 mm. wide; style 4-4.5 mm. long; sta-
mens about 50, up to 3.5 mm. long; petals obovate, white, ciliate, about 2.5 mm.
wide, 3.5 mm. long; ovary bilocular, the ovules 4-6 in each locule, radiating from
a centrally attached placenta; fruit globose or oblate, black, 0.7-0.8 cm. in diam-
eter, broadly short-stipitate.

Eugenia vesca Lundell, Wrightia 2: 212. 1961. Guayabillo.
In high forest bordering an arroyo, known only from the type
locality; Pete"n (south of Dolores, E. Contreras 2080, the type).

A shrub, or treelike, 1-1.5 meters high, the bark of 1-year-old branchlets
scaling off in thin pale reddish-brown platelets; plants essentially glabrous, the
vegetative buds and the inflorescence and probably the young shoots (not seen)
sparingly strigose with short reddish hairs; leaves elliptic or ovate, 2-5.5 cm. wide,
4-13 cm. long, 2-2.8 times as long as wide, narrowed to a blunt apex or obscurely
acuminate, rounded to a cuneate base, the margins rather long-decurrent on the
dilated summit of the ventrally flat or concave petiole 4-6 mm. long; midvein
channeled above, often bordered by parallel convex ridges; lateral veins 7-10 on
each side, more conspicuous beneath; marginal vein about equaling the lateral
ones and somewhat arched between them, 1-4 mm. from the margin; blades
darker green and verrucose above, with numerous convex glands; lower surface
yellow green when dry, sparingly dark-dotted; inflorescence a tight axillary or
supra-axillary cluster of about 2-3 opposite and decussate pairs of sessile flowers
on a stout axis 1 mm. long or less; bracteoles partly hiding the immature buds,
broad and membranous, ciliate, suborbicular or broadly deltoid-ovate, 0.8-1 mm.
long, 1-1.3 mm. wide, connate, forming a persistent 2-lobed involucre; immature
buds 1.5 mm. long; calyx-lobes in unequal pairs, thin, ciliate, suborbicular, broadly
imbricate, the outer ones 1.3 mm. wide; fruit red, globose, 5-7 mm. in diameter,
surmounted by the thin erect calyx-lobes.

370 FIELDIANA: BOTANY, VOLUME 24

Eugenia Winzerlingii Standl. Trop. Woods 11: 20. 1927.

Savannah thickets, often or usually on limestone, 200 meters or
less; Pete"n; Alta Verapaz. Yucatan; Campeche; British Honduras,
the type from Orange Walk District, H. W. Winzerling VIII-7.

Shrub or small tree to 6 meters high, glabrous; leaves rigidly coriaceous,
rounded-obovate, oblanceolate or oblong-elliptic, 1.5-5.5 cm. wide, 4-10 cm. long,
1.6-2.3 (-3) times as long as wide, obtuse or rounded at apex, acute or some-
times rounded at base, the thickened cartilaginous margins often rather abruptly
narrowed at the very base, there decurrent on the inner angles of the very stout
petiole; petiole swollen (as seen on the abaxial surface) at the base of the midvein,
its free part 2-4 mm. long, up to 2.5 mm. thick, adaxially nearly flat or concave;
midvein prominently convex above, often 0.7-1.1 mm. wide near base; lateral
veins 6-9 principal pairs, ascending; marginal vein about equaling the laterals,
arched between them (strongly so in some broadly obovate leaves), 2-4 mm. from
the margin; veins in dried leaves visible on both surfaces, on the lower surface
often forming a conspicuous closed pattern well within the leaf-margin; blades
lustrous above, paler and dull and often drying brown beneath; glands obscure,
often evident but not abundant beneath; flowers in umbelliform clusters, the in-
florescence a very short axillary raceme 2-4 mm. long, 1-2 mm. thick, bearing
1-6 closely approximate opposite decussate pairs of flowers on filiform pedicels
10-20 mm. long (up to 0.6 mm. thick in fruit); bracts scarious, concave, ciliate,
ca. 0.7 mm. long and wide; bracteoles persistent, ovate, acute, ciliate, 1-1.5 mm.
long; buds 4-4.5 mm. long, the conic hypanthium surpassed by the dome of the
petals; calyx-lobes gibbous at base in bud, thin, rounded, in unequal pairs, ca.
2.5 mm. wide, 2 mm. long, somewhat accrescent and persistent; disk 1.5-2 mm.
wide; style 5-6 mm. long; stamens 40-60, up to 4 mm. long, the anthers 0.7 mm.
long; petals white, ciliate, broadly obovate, 5 mm. long; ovary bilocular, the
ovules 12-13 in each locule, radiating from a centrally affixed placenta; fruit glo-
bose, black, 6-10 mm. in diameter.

Eugenia yucatanensis Standl. Field Mus. Bot. 8: 28. 1930.
Guayabillo.

In forest, at low elevations; Pete"n (Dolores, E. Contreras 2592).
Yucatan, the type from Izamal, G. F. Gaumer in 1888; Quintana
Roo; British Honduras.

A tree as much as 12 meters high with a trunk 25 cm. in diameter, sordid-
pubescent, the branchlets and inflorescence and the lower surfaces of young leaves
rather densely beset with appressed (simple) coarse stiff hairs up to 0.5 mm. long,
and also with shorter suberect antrorse bristle-like hairs; leaves broadly elliptic-
ovate, often about equally narrowed to the abruptly acuminate tip and to the
rounded or acute base, 2-4.5 cm. wide, 4-9 cm. long, 1.7-2.5 times as long as wide,
the margins at base cuneately decurrent on the channeled petiole 5-6 mm. long,
1 mm. thick; midvein impressed above, the narrowly V-shaped furrow usually
persistently hispidulous; lateral veins close and parallel, about 10 pairs in addition
to rather numerous intermediate ones, apparent on both sides of the blade when
dry; marginal vein well defined and relatively straight, about equaling the lateral

McVAUGH: FLORA OF GUATEMALA 371

ones and arched between them, 1-2 mm. from the margin; leaves at maturity
somewhat strigose on both surfaces, or glabrate, lustrous above, scarcely paler
beneath, with large translucent glands on both surfaces, sometimes impressed-
punctate above; inflorescence a short axillary raceme, or the flowers sometimes
at the lowermost (bracteate) nodes of a leafy branchlet, the raceme-axis 7-20 mm.
long, bearing 2-6 opposite, decussate pairs of flowers; terminal flower abortive;
internodes angular, flattened, 0.5-0.7 mm. wide; pedicels 3-10 mm. long; bracts
ovate-deltoid to lanceolate, 1-1.8 mm. long, scarious, somewhat persistent; bracte-
oles acute, broadly deltoid-ovate, 1-1.3 mm. long, persistent, slightly coherent
by their basal margins, convex on the backs, prominently gland-dotted, essentially
glabrous, or strigose; buds 3 mm. long, pyriform, the densely pubescent hypan-
thium 1-1.5 mm. long, canescent; calyx-lobes nearly glabrous without, appressed-
pubescent within, in unequal pairs, the outer ones 1.3 mm. long, the inner 1.5-
2.5 mm. long, 2 mm. wide, the lobes in fruit subfoliaceous, 2-3.5 mm. long; disk
bristly, 2.5 mm. wide; style 3.5-4.5 mm. long; stamens about 40, up to 4 mm. long;
petals rotund or obovate, 3-3.5 mm. long; ovary bilocular, the ovules 4-6 in each
locule, collateral, pendent; fruit fleshy, purple-black, about 8-costate, 6-7 mm.
in diameter.

Material of this species is very similar to Jamaican material that
has been identified as Eugenia disticha (Sw.) DC. or as E. biflora
(L.) DC. Urban, in his paper on the West Indian Myrtaceae, inter-
preted E. biflora broadly, including in it many plants with broad,
bluntly acuminate and only obscurely glandular leaves; these are
quite different from the plants included in this Flora under the name
of E. biflora, but are indistinguishable from E. yucatanensis. It is
probable that a thorough revision of the group of E. biflora would
result in the reduction of E. yucatanensis.

Eugenia Yunckeri Standl. Field Mus. Bot. 17: 380. 31 Jan.
1938. E. siltepecana Lundell, Amer. Midi. Nat. 19: 430. Mar. 1938.

In forest and along rivers, 1300-2000 meters; to be expected in
Guatemala. Mexico (Chiapas, Matuda 1594, type of E. siltepecana).
Honduras (Comayagua, Yuncker et al. 6178, type).

A shrub or small tree up to 2.5 meters high, the branchlets, petioles and in-
florescences densely tawny-hirsutulous; lower leaf-surfaces similarly but less
densely hirsute, the erect lustrous hairs 0.5-1 mm. long; leaves elliptic, 2-4 (-5.7)
cm. wide, (5-) 7-12 (-15) cm. long, 2-3 times as long as wide, obtuse or usually
broadly and bluntly acuminate at tip, rounded to the acute or cuneate base, the
somewhat revolute margins shortly decurrent on the stout nearly terete petiole
7-12 (-15) mm. long, 1.5-2.5 mm. thick including the hairs; midvein somewhat
impressed above, hairy; lateral veins 8-10 on each side, ascending, inconspicuous
above, apparent beneath; marginal vein about equaling the lateral ones and arched
between them, 2-4 mm. from the margin; blades usually drying brown to black,
probably lustrous above when fresh, paler beneath, the glands inconspicuous; in-
florescence a crowded axillary raceme, the axis 4 mm. long or less, bearing 3-4
pairs of flowers on pedicels 1-2 mm. long or less; bracteoles oblong, distinct, gla-

372 FIELDIANA: BOTANY, VOLUME 24

brous on the inner surface, 0.7 mm. wide, 1.5-1.7 mm. long; buds pyriform,
3-4 mm. long; hypanthium campanulate, densely hirsute, 1.7-2.5 mm. long;
calyx-lobes broadly rounded, hirsute without, finely pubescent within, up to 2.5-
3 mm. wide, 2.5-4 mm. long, the smaller ones about 2 mm. long and wide; disk
2.5-3.5 mm. wide, often quadrate, hirsute; style about 7 mm. long; stamens about
125; ovary bilocular, the ovules about 8-10 in a compact group in each locule;
fruit globose or oval, sparingly short-hirsutulous, 1.2-1.5 cm. in diameter, 1.3-
1.5 cm. long, crowned by the subpersistent calyx-lobes.

SPECIES INSUFFICIENTLY KNOWN

Eugenia ardisioides Lundell, Amer. Midi. Nat. 29: 479. 1943.

British Honduras, Toledo District, Monkey River, Jenkins Creek,
in hammock in pine ridge, P. H. Gentle 4081, the type. Unknown
except from this collection.

A small tree with thin broadly elliptic and somewhat acuminate leaves 6-
13 cm. long, the racemes minutely puberulent, slender and short, the pedicels
2.5-5 mm. long, the flowers small. The type bears somewhat immature fruit
5-6 mm. in diameter. The leaves suggest in appearance those of E. acapulcensis,
but the midvein is scarcely impressed; the racemes are more open than those of
E. acapulcensis.

Eugenia itzana Lundell, Bull. Torrey Club 69: 395. 1942.

Doubtfully an independent species, E. itzana seems to fall be-
tween E. acapulcensis and E. axillaris. All four specimens cited in
the protologue of E. itzana are small-flowered for acapulcensis, re-
sembling in this respect E. axillaris. One paratype, Steere 1373, is
scarcely distinguishable from axillaris. The type (Yucatan, Chichen
Itza, Lundell 7589) is an immature flowering specimen, suggesting
superficially a small-leaved representative of acapulcensis. Similar
specimens may yet be found in Pete"n.

Eugenia oblanceifolia Lundell, Amer. Midi. Nat. 29: 480. 1943.

British Honduras, Toledo District, Monkey River, Jenkins Creek,
in hammock in pine ridge, P. H. Gentle 4078, the type. Unknown
except from this collection.

A small tree with thin narrowly elliptic and gradually acuminate leaves 4-
8.5 cm. long, the racemes minutely puberulent, 1 cm. long or less, the pedicels
2.5-6 mm. long, the flowers small. The type bears mature or nearly mature fruit
about 6 mm. in diameter.

This species, as originally suggested by Dr. Lundell, is very sim-
ilar to E. ardisioides, from which it hardly differs except in its some-
what narrower leaves. Like E. ardisioides it somewhat resembles

McVAUGH: FLORA OF GUATEMALA 373

E. acapulcensis, but the racemes are more like those of, e.g., Eugenia
riograndis.

Eugenia peroblata Lundell, Wrightia 2: 124. 1961.

British Honduras, El Cayo District, high ridge, hill slope, Gorge
Creek section, Humming Bird Highway, P. H. Gentle 8845, the type.
Known only from this collection.

A tree "2 inches in diameter," the young branches and the petioles and prob-
ably the inflorescence hispidulous; leaves elliptic, 2.5-3 cm. wide, 8-11 cm. long,
about 3 times as long as wide, about equally narrowed to the narrowly acuminate
tip and the gradually rounded base, the margins passing abruptly into the nearly
terete petiole 1 mm. thick, 5-6 mm. long; midvein convex or nearly flat above,
pubescent most of its length; lateral veins slender and nearly parallel, 10-12 pairs
including some intermediates; marginal vein about equaling the lateral ones, rather
well-defined and straight, 1.5-3 mm. from the margin; leaves nearly concolorous
when dry, the glands obscure on both surfaces; vegetative buds prominently im-
bricate-scaly with keeled scales; inflorescence and flowers not seen; fruits seen
detached, the flowers and fruits probably borne in sessile or nearly sessile clusters
at defoliated nodes; ovary and prolonged rim of the hypanthium apparently
coarsely hairy, the fruit glabrate, strongly oblate, 1.5 cm. high, 2-2.5 cm. in
diameter, surmounted by the irregularly rupturing remains of the hypanthium
and calyx; seed 1, the cotyledons free, fleshy, plano-convex, one in the upper half
and the other in the lower half of the fruit; radicle (ex Lundell) short but con-
spicuous.

This plant bears a considerable superficial resemblance to the
little-known South American and West Indian species that has been
variously known as Myrciaria involucrata Berg, Marlierea glomerata
Berg, and Plinia pinnata L. (sensu Urban, Repert. Sp. Nov. 15:
412-413. 1919, et al.). It almost certainly is not that species, but
the two species are sufficiently alike in vegetative characters to sug-
gest they are congeneric. Certainly the characters of the fruit of
E. peroblata exclude it from Eugenia and suggest that it belongs with
Plinia as that genus was interpreted by Amshoff (Fl. Suriname 3:
97-99. 1951) and McVaugh (Fl. Peru 775-780. 1958). I prefer not
to transfer it formally to Plinia in the absence of knowledge about
its flowers and the details of its ovarian structure.

Eugenia uliginosa Lundell, Wrightia 3: 20. 1961.

Known only from the type, an over-mature fruiting specimen;
Chiapas, La Grandeza, alt. 2016 meters, E. Matuda 5546.

A tree 3-4 meters high, glabrous; leaves coriaceous, elliptic-ovate, about 4 cm.
wide, 9 cm. long, rounded to a short blunt acumen and to a cuneate base, the
revolute margins strongly decurrent on the dilated summit of the channeled peti-

374 FIELDIANA: BOTANY, VOLUME 24

ole about 1 cm. long; midvein flat or concave above; lateral veins 10-15 on each
side including some intermediate ones; marginal vein about equaling the lateral
ones and but slightly arched between them, 1-2 mm. from the margin; blades
probably glossy above, paler and finely gland-dotted beneath, the upper surface
obscurely glandular- verruculose; inflorescence a very short axillary raceme or 2
racemes superposed, the axis up to 3 mm. long, quadrangular and 2 mm. thick
in fruit, bearing up to 3 pairs of flowers; pedicels in fruit 5 mm. long, 1.5 mm.
thick; bracteoles indurate-persistent, very short; fruit 1.5-1.8 cm. in diameter,
with nearly globose body and stout stipe, the whole 2.3 cm. long; calyx-lobes
apparently broadly rounded, 3 mm. wide; disk 3 mm. wide in fruit; seed one,
homogeneous.

Superficially this specimen suggests E. Percivalii Lundell, but
that species is pubescent in the flowering condition, and is a plant
of the lowlands of British Honduras.

MYRCIA DC.

References: Berg, Linnaea 27: 35-80, 82-129. 1855-1856; and in
Mart. Fl. Bras. 14, pt. 1: 59-143, 150-210. 1857.

Calyx-lobes usually 5, distinct and imbricated in bud and in flower, and usually
persistent on the fruit; hypanthium variously or not at all prolonged beyond the
summit of the ovary; petals usually 5, often half as long as the stamens or longer,
and conspicuous. Inflorescence a many-flowered panicle [i.e. several times com-
pound; the central axis, and those of the principal branches of the panicle, pro-
longed and bearing several pairs of lateral branches]. Ovules 2. Embryo with
long radicle, and thin, foliaceous, much-folded cotyledons. Fruit thin-fleshed,
1- or 2-seeded.

The term "panicle," as used below in the descriptions of species of
Myrcia, refers to the entire fertile branch that arises in the axil of a
leaf; in M. leptoclada the two panicles at a node may arise from the
two foliar axils; more commonly they arise from the opposite sides
of a nearly naked node [i.e. merely subtended by small bractlets],
and this pair of panicles may appear to be terminal because the
branchlet bearing them fails to develop beyond the bracteate node
at which they arise.

A large genus of tropical America, including probably several
hundred species. A few species not treated below occur in Costa
Rica and Panama.

Summit of the ovary and interior of the hypanthium glabrous; flower-buds gla-
brous or nearly so; plants blackening in drying, nearly glabrous, the young
branchlets and the inflorescence with somewhat appressed coppery partly
dibrachiate hairs; fruit depressed-globose M. leptoclada.

Summit of the ovary and interior of the hypanthium densely hairy; flower-buds
densely pale strigose or -silky toward the base; plants green in drying, the

McVAUGH: FLORA OF GUATEMALA 375

branchlets and the inflorescence abundantly pubescent with yellowish or pale
reddish ascending simple hairs; fruit oblong or oval to nearly globose.

M. splendens.

Myrcia leptoclada DC. Prodr. 3: 244. 1828. Aulomyrcia lepto-
clada (DC.) Berg, Linnaea 27: 40. 1855. Myrcia Gentlei Lundell,
Wrightia 2: 214. 1961.

Wet forests near sea level; Izabal (Puerto Barrios, Standley
73072). British Honduras (Toledo Dist., P. H. Gentle 4378, type
of M. Gentlei}; Honduras; Greater Antilles to Trinidad; Suriname.

A shrub or a tree up to 15 meters high, blackening in drying, the branchlets
and inflorescence sparingly pubescent with partly appressed coppery partly di-
brachiate hairs 0.2-0.4 mm. long; leaves coriaceous, ovate to elliptic or elliptic-
lanceolate, 2-5 cm. wide, 6-12 cm. long, 2-3 times as long as wide, gradually and
bluntly acuminate at tip, rounded to the base, rather abruptly contracted to the
very base but the margins shortly decurrent on the inner angles of the channeled
petiole 2.5-4 mm. long; midvein impressed above; lateral veins close and parallel,
12-15 on each side in addition to some intermediate ones, nearly straight, passing
directly into the marginal vein which is arched between them, 2-3 mm. from the
margin; blades nearly concolorous when dry, coffee-brown to black, lustrous, the
glands inconspicuous; paired panicles arising from the lowermost (bracteate) nodes
of leafy branchlets or (usually) paired at the summit of an abortive axillary axis
2-4 mm. long, the individual panicles 5-9 cm. long, many-flowered, 3 times com-
pound, very broadly pyramidal, the peduncle 1-3 cm. long, the lowest lateral
branchlets much longer, irregularly and often alternately branched, their elongate
tips often raceme-like with 2-3 pairs of flowers each, on pedicel-like branchlets
1-2 mm. long; buds turbinate, rounded at apex, 2 mm. long including a short-
stipitate base, the hypanthium glabrous; calyx-lobes 5, strigose without, appressed-
pubescent within near the tips, very unequal, the three outer ovate-deltoid, thick-
ened at base, 0.5-1 mm. long, the two inner ones orbicular, membranous, almost
petaloid, 1.5 mm. long and 1.5-1.8 mm. wide; disk bowl-shaped, 1.5 mm. wide,
glabrous; style 3-4 mm. long; stamens about 75; petals white, 1.5 mm. long; fruit
black, glabrous, depressed-globose or transversely oval, 1- to 2-seeded, 7 mm.
high, 7-10 mm. broad.

Called "parrot plum" in British Honduras.

Myrcia splendens (Sw.) DC. Prodr. 3: 244. 1828. Myrtus
splendens Sw. Prodr. 79. 1788 (type from Hispaniola). Myrcia rufi-
dula Schlecht. Linnaea 13: 416. 1839 (type from Veracruz, Schiede &
Deppe). M. costa-ricensis Berg, Linnaea 27: 104. 1855 (type from
Costa Rica, Oersted). M. discolor Berg, I.e. Ill (type from Costa
Rica, Oersted). M. Oerstediana Berg, I.e. 112 (type from Turrialba,
Costa Rica, Oersted 28). M. melanoclada Berg, I.e. 113 (type from
Cartago, Costa Rica, Oersted 7). M. plicato-costata Berg, I.e. 114
(type from Turrialba, Costa Rica, Oersted). M. Sartoriana Berg,

376 FIELDIANA: BOTANY, VOLUME 24

Linnaea 29: 220. 1858 (type from Veracruz, C. Sartorius). M. longi-
caudata Lundell, Amer. Midi. Nat. 29: 481. 1943 (type from Lance-
tilla Valley, Honduras, A. M. Chickering 153). M. Schippii Lun-
dell, I.e. 482 (type from All Pines, Stann Creek District, British
Honduras, W. A. Schipp 797). Agal (Huehuetenango).

Moist or wet thickets, or dense or open, mixed or pine forest,
sometimes on limestone, 1650 meters or less; Pete"n; Alta Verapaz;
Huehuetenango; San Marcos. Southern Mexico (Veracruz) ; British
Honduras; Honduras; West Indies; South America.

A shrub or small tree 2-5 or sometimes 10 meters high, up to 15 cm. in diam-
eter, abundantly pubescent with ascending lustrous yellowish- or brownish-white
or sometimes rufous hairs up to 1 mm. long (often 0.5 mm. long); hypanthium,
branchlets and inflorescence, and the dorsal surface of the midvein closely silky-
strigose or somewhat shaggy; calyx-lobes and lower leaf -surf ace sparingly or closely
strigose; leaves elliptic-lanceolate, elliptic, or ovate-lanceolate, 2-4 (-4.5) cm. wide,
6-12 cm. long (up to 16.5 cm. long on shoots), (2.2-) 2.5-3 (-4) times as long as
wide; blades abruptly narrowed at tip to a blunt point or, usually, rather narrowly
acuminate, often with slender tip 0.8-2 cm. long; base cuneate or, usually, rounded
with the margins cuneately decurrent on the stout channeled petiole 3-4 mm. long;
midvein impressed on the upper surface, usually bearing a line of stiff, often erect
hairs for most of its length; lateral veins rather close together and parallel, 12-20
pairs in addition to some intermediates, elevated on both surfaces in dried leaves
but not very conspicuous, the small connecting veins forming a conspicuous net-
work; marginal vein about equaling the laterals and but slightly arched between
them, 1 (-2) mm. from the margin; upper surface of the blade smooth and lus-
trous, bright green, the lower paler; convex glandular dots sometimes apparent
on new leaves, rarely on mature leaves, the glands never impressed-punctate; in-
florescence axillary and subterminal, 4-6 (-7.5) cm. long, with up to about 50
flowers (often fewer) in each panicle, 3 times compound, the branches opposite
or nearly so, terete or but slightly flattened below the nodes (1-1.5 mm. wide below
the first node); peduncle usually 1.5-2.5 cm. long, often %-% as long as the en-
tire panicle; flowers aggregated toward the tips of the branches; buds 2-2.5 mm.
long before anthesis, the hypanthium narrowly turbinate; calyx-lobes 5, drying
dark, glabrous or very sparingly hairy on the inner surface, bluntly deltoid or
broadly rounded, 1-1.2 mm. long, about 1.5 mm. wide, often reflexed in post-
anthesis; disk about 2 mm. wide, densely hairy, the center slightly or markedly
depressed; style 3.5-5 mm. long, hairy near base or below the middle; petals
white, about 2-3 mm. long; stamens white, about 75-85, nearly as long as the
style; fruit oblong or oval, or sometimes nearly globose, at first red, turning black,
at maturity about 1 cm. long.

This species belongs to a complex that is taxonomically poorly
known (cf. Fieldiana, Bot. 29: 470-473. 1963). It is closely related
to several South American species, viz. M. acuminata (HBK.) DC.,
M. fallax (Rich.) DC., and M. mollis (HBK.) DC. In M. splendens
and M. fallax the inner face of the calyx-lobes is entirely glabrous
or very sparingly pubescent. In M. mollis, a Colombian species

McVAUGH: FLORA OF GUATEMALA 377

extending northward as far as Costa Rica, the calyx-lobes are about
equally soft-hairy on both sides; the inflorescence is stouter than
that of M. splendens, the peduncle usually 2-2.5 mm. wide below the
first node. Calyx-lobes that are pubescent on the inner surface may
be found in an occasional Central American specimen (e.g. Schipp
797, the type of M. SchippH), but these specimens do not seem to
differ otherwise from the less pubescent ones, more typical of M.
splendens, found in the same localities.

A species of somewhat doubtful status is Myrcia belizensis Lun-
dell, Wrightia 2: 213. 1961, described by the collector as a woody
vine. This is known only from one locality in British Honduras
(near San Antonio, Toledo District, Gentle 6571, the type). The
specimens resemble M. splendens but the petioles are 5-9 mm. long,
resembling in this respect those of M. fallax. The latter, not yet
reported from Central America except from Panama, has a relatively
stout inflorescence, the peduncle usually 2-2.5 mm. wide just below
the first node.

MYRCIANTHES Berg
Reference: McVaugh, R., Fieldiana, Bot. 29:473-497. 1963.

Calyx-lobes 4, rarely 5, distinct, imbricated, persistent. Hypanthium not
prolonged beyond the summit of the ovary. Petals as many as the calyx-lobes,
conspicuous. Ovary usually bilocular, the ovules several (5-20, usually 10-15)
in each locule, in a subcapitate group attached near the middle of the central
dissepiment. Seeds one or two, the cotyledons distinct, accumbent, large, plano-
convex; radicle often hairy, stout, usually about one-third as long as the cotyle-
dons or less; plumule often silky-hairy, much shorter than the radicle. Inflores-
cence an axillary dichasium, the terminal flowers usually sessile in the forks.

An American genus of about 40 species, from peninsular Florida,
the West Indies and Mexico to Argentina and Bolivia. Only the
following species is known from northern Central America; one or
two additional species occur in Costa Rica and Panama.

Myrcianthes fragrans (Sw.) McVaugh, Fieldiana, Bot. 29: 485.
1963. Myrtus fragrans Sw. Prodr. 79. 1788. Eugenia fragrans (Sw.)
Willd. Sp. PI. 2: 964. 1800. Anamomis fragrans (Sw.) Griseb. Fl.
Brit. W. Ind. 240. 1860. E. triflora Sesse" & Moc. Naturaleza II. 1:
app. 83. 1888 (type from Cuernavaca, Morelos). Myrtus biflora
sensu Sess<§ & Moc. Fl. Mex. ed. 2. 125. 1894 (type from Cordoba,
Veracruz). Myrcia Seleriana Bonn. Sm. Bot. Gaz. 27: 332. 1899
(type from Chacula, Guatemala, Seler 3169). E. Steyermarkii Standl.
Field Mus. Bot. 22: 360. 1940 (type from Guatemala near Volcan

378

FIELDIANA: BOTANY, VOLUME 24

FIG. 52. Dichasium and embryo in Myrcianthes. A, The inner face of one
cotyledon showing position of hypocotyl and radicle and small plumule at right;
X ca. 17. B, Seed (somewhat generalized) with outer coats removed to show plano-
convex cotyledons and projecting radicle; X ca. 7. C, Seven-flowered dichasium
of M.fragrans (Steere 2976) ; X %.

de Tacana, Steyermark 36210). E. Lopeziana A. Molina, Ceiba 3:
170. 1953 (type from Honduras, Molina 3007).

Wet or moist, mixed forest, on limestone, or in pine-oak forest,
800-2000 meters; Baja Verapaz; Quezaltenango; Sacatepe"quez; San
Marcos; Huehuetenango. Southern Florida; eastern and southern
Mexico; British Honduras; Honduras; Nicaragua; Costa Rica; West
Indies; northern South America.

A shrub or small tree up to about 15 meters high, the brown outer bark peel-
ing off in plates and exposing the paler or cream-colored inner bark; herbage and

McVAUGH: FLORA OF GUATEMALA 379

branchlets more or less pubescent with pale appressed or ascending hairs, the hy-
panthium whitened by a dense covering of gray-white hairs, contrasting with the
green glabrous calyx-lobes; petals and calyx-lobes ciliate, the latter also appressed-
hairy within; leaves glabrate, elliptic to ovate or obovate, 1.5-3.5 cm. wide, 3-
7 cm. long, 1.8-2.6 times as long as wide, obtusely pointed or obtusely acuminate
or occasionally rounded at apex, acute at base or the margins at right angles,
usually cuneate at very base and the margins decurrent on the channeled petiole
(2.5-) 4-6 mm. long; mid vein impressed above, prominent beneath; lateral veins
about 8-12 on each side, indistinct and often with smaller intermediate veins,
slightly elevated on both sides when dry; marginal vein about equaling the laterals
and arched between them; blades somewhat lustrous above, paler beneath, copi-
ously dotted with minute glands; inflorescence a 3- or often 7-flowered dichasium
(sometimes reduced to a single flower on a pedicel 1 cm. long), the central flower
usually sessile, the 2 lateral flowers sessile or on branches 3-5 mm. long; peduncle
2-4 (-6) cm. long, (0.7-) 1.2-1.5 mm. wide at apex, somewhat compressed; bracts
and bracteoles linear, 2-3 mm. long, falling at anthesis; calyx-lobes usually 4,
(rarely 5 in some flowers), broadly rounded, 2.5-3.5 mm. wide at base, all alike
or in 2 unequal pairs, the outer pair thicker and somewhat obtusely deltoid; disk
usually quadrate, 3-3.5 (-4) mm. wide, the staminal ring hairy, the flat central
portion glabrous; style 6.5-7.5 mm. long; stamens about 150, the longer ones
7-9 mm. long; petals white or yellowish white, concave, oblong-obovate, 4-6 mm.
long; ovary bilocular; ovules 15-20 in each locule, radiate in a subcapitate group
affixed to the central dissepiment; fruit dark purple-black, subglobose, smooth or
roughened by the numerous superficial glands, about 1 cm. in diameter; seed
bean-shaped, usually only one maturing; cotyledons 2, plano-convex; radicle short
less than half as long as the cotyledons; plumule silky, often very short.

This variable species ranges widely around the Caribbean, and
perhaps may ultimately be divided into several infraspecific taxa.
Most specimens from the continent of North America, and a few
from northern South America, are similar to the nomenclaturally
typical plant of Jamaica (M . fragrans var. fragrans) except that the
petioles are often longer, the leaf-blades narrower and the dichasia
often 7-flowered instead of 3-flowered. The few Guatemalan speci-
mens I have seen are of this type. A specimen from Quintana Roo,
however (Cozumel Island, Steer e 2976), resembles rather the strongly
glandular plant with obovate-oblong leaves that is common in Cuba
and Hispaniola. The var. hispidula McVaugh, a plant of Costa
Rica and Panama, has not been found in northern Central America.

MYRCIARIA Berg

Reference: Berg, Linnaea 27: 320-338. 1856; and in Mart Fl. Bras.
14, pt. 1: 358-376. 1857; McVaugh, R., Fieldiana, Bot. 29: 499-504.
1963.

Calyx-lobes 4, distinct and imbricated. Hypanthium markedly prolonged
above the summit of the ovary, circumscissile at base, and cleanly deciduous,

380

FIELDIANA: BOTANY, VOLUME 24

with the perianth and androecium, about the time of anthesis. Flowers sub-
sessile, in racemosely arranged clusters of (usually) 4 each. Ovules 2. Embryo
(according to Amshoff) undivided. Fruit fleshy, subglobose, about 2-seeded,
naked and umbilicate at apex.

FIG. 53. Myrciaria floribunda. A, Open flower somewhat past anthesis, from
below, showing the base of the hypanthium after circumscission. B, Lateral
(i.e. dorsal) view of bud, showing partly enveloping bracteoles at base. Both X 15.

A well-marked genus, probably of about 25 species, in tropical
America from Mexico and the West Indies southward, particularly
at low elevations. One additional species occurs in Costa Rica and
Panama.

Leaves of an ovate type, mostly 2-2.5 times as long as wide, sharply acuminate
and often cuspidate, broadly rounded at base, the margins little if at all de-
current on the petiole; branchlets and petioles coarsely and rather sparsely
hispidulous; flowers sessile; bracteoles very broadly rounded and imbricate,
connate at very base only, 1.5-2 mm. long and wide, much exceeding the
ovary; Guatemala (Peten) M. Ibarrae.

Leaves of a lanceolate type, mostly 2.5-3 times as long as wide, variously acumi-
nate but the very tip usually blunt or merely acute, the base of the blade
somewhat rounded below the middle, usually acute or cuneate with the mar-
gins cuneately decurrent on the splayed summit of the petiole; branchlets
and petioles finely pubescent or rarely glabrous; pedicels usually 0.5-1 mm.
long; bracteoles not imbricate (or very slightly so in the bud stage), united
by their proximal margins into an oval bilobed or finally explanate involucre
2 mm. long; West Indies; eastern Mexico and Central America; Colombia;
lowlands of the Guianas; Orinoco and Amazon lowlands from southern Vene-
zuela to Peru and eastern Brazil (Maranhao) M. floribunda.

Myrciaria floribunda (Willd.) Berg, Linnaea 27: 330. 1856.
Eugenia floribunda West ex Willd. Sp. PI. 2: 960. 1800. E. 0' Neillii
Lundell, Bull. Torrey Club 64: 555. 1937 (type from British Hon-
duras, Hugh O'Neill 8767). M. 0' Neillii (Lundell) I. M. Johnst.
Sargentia 8: 228. 1949.

McVAUGH: FLORA OF GUATEMALA 381

Wet thickets, mangrove swamps or mixed forest, 600 meters or
less; Pete"n; Alta Verapaz; Izabal. Southern Mexico (Oaxaca, Ta-
basco, Chiapas); British Honduras; El Salvador; Costa Rica to
northern Colombia; West Indies; Guianas; eastern Brazil.

Shrub or small tree up to 10 meters high, 15 cm. in diameter; branchlets red-
dish brown, the bark checking longitudinally and peeling in small plates; branchlets
and young growth finely soft-pubescent, the leaves glabrate, the inflorescence
somewhat pubescent; leaves ovate or lanceolate, 0.7-2 (-3) cm. wide, 2.5-6 (-8)
cm. long, 2.5-3 (-4) times as long as wide, caudate-acuminate (the tip sometimes
2 cm. long, 2.5-3 mm. wide at base), or less often short-acuminate or merely acute,
the very tip usually blunt; blade rounded in the proximal third to the mostly
acute or cuneate base, the margins somewhat decurrent on the broadened summit
of the flat or sometimes sulcate petiole 3-6 mm. long; midvein flat or slightly con-
cave above, often persistently pubescent like the petiole; lateral veins rather
numerous, nearly straight and parallel, 10-20 pairs, passing directly into a mar-
ginal vein, the latter nearly straight, parallel to the leaf-margin and about 0.5-
1 mm. from it; blades rather thin, the two surfaces nearly concolorous, the upper
somewhat lustrous, light green and nearly featureless, the lower usually with some
convex glands when dry; inflorescence axillary, or apparently somewhat supra-
axillary; axis less than 1 mm. long, bearing 1 or 2 decussate pairs of flowers on
stout pedicels up to 0.5 mm. thick, 0.5-1 mm. long; bracts dark brown, broadly
rounded, ciliate and pubescent, 0.8 mm. wide; bracteoles persistent, broadly
rounded, ciliate and pubescent without, united by their proximal margins into a
cuplike involucre 0.5-0.6 mm. high, this finally explanate, 2 mm. long, 1.5 mm.
wide; hypanthium sessile, broadly conic or cup-shaped, about 1.5 mm. high and
1.5 mm. wide at the mouth, deciduous after anthesis, glabrous within and without;
calyx-lobes broadly rounded, densely silky-pubescent within, and near the tips
on the outer surface, the free parts broadly rounded, 1-1.5 mm. wide, imbricate
in the bud, at anthesis separating by irregular splits in the distal half of the hypan-
thium; bracteoles, hypanthium, calyx-lobes and corolla with prominent large
glands; style 4-5 mm. long, pilose below with long pale hairs like those of the
summit of the ovary; stamens about 75, up to 5 mm. long, the anthers 0.5 mm.
long; petals irregularly ovate, ciliate, 1.5 mm. long; ovary bilocular, the ovules 2
in each locule, collateral; fruit soft, globose or oblate, about 7 mm. in diameter,
naked at apex, the circular scar there about 1 mm. across.

Myrciaria Ibarrae Lundell, Wrightia 2: 213. 1961. Guayabillo.

In low forest (zapotal), in lowlands; Pete"n, the type from Dos
Lagunas, E. Contreras 1645. Endemic.

A tree 8-10 meters high, 7-10 cm. in diameter, the branchlets and petioles
uniformly hispidulous-pubescent with coarse bristle-like erect or often antrorsely
crisped pale hairs 0.3-0.5 mm. long; leaves glabrous or the midvein retaining
some hairs, the blades ovate or lance-ovate to elliptic, 1-2 (-2.8) cm. wide, 4-6
(-8) cm. long, (1.7-) 2-2.5 (-3) times as long as wide, the apex gradually and
sharply acuminate, often cuspidate, the base broadly rounded, the margins not
at all or scarcely decurrent, passing abruptly onto the inner angles of the chan-

382 FIELDIANA: BOTANY, VOLUME 24

neled petiole 4-7 mm. long; midvein flat above or concave toward the base, prom-
inent as a pale line beneath; lateral veins obscure, close and parallel, very slender,
15-20 or more on each side including the intermediate ones; marginal vein about
equaling the lateral ones, nearly straight, 0.3-0.7 mm. from the revolute carti-
laginous margin; leaves nearly concolorous, with scattered small convex glands
beneath; flowers sessile, nearly glabrous, in sessile clusters of 2-6 (usually 2) at
a node, the racemose axis about 1 mm. long and 1 mm. thick, with up to 4 approx-
imate nodes; bracts indurate, ovate-deltoid to reniform, strigose, persistent;
bracteoles broadly rounded, pale, ciliate, concave, coriaceous, up to 1.5 (-2) mm.
long and wide, strongly imbricate at least in the young flower and half-enveloping
the oval buds which are 2.5-3 mm. long, contracted toward the mouth; hypan-
thium prolonged 1.5 mm. beyond the summit of the ovary, glabrous without
and within, its base 1-1.3 mm. in diameter at time of abscission; disk white-
tomentose, 1-1.5 mm. wide; style 5-6 mm. long; calyx-lobes in unequal pairs, short
and broadly rounded, appressed-pilose within, the larger ones 1-1.5 mm. wide,
less than 1 mm. long; stamens about 75, up to 5 mm. long, the anthers 0.4 mm.
long; petals 4, ciliate, ovate to subrotund, white, 1.5-2 mm. long and wide; locules
2, the ovules 2 in each locule; fruit (according to Lundell) globose, 1- or 2-seeded,
up to 2.5 cm. in diameter, the embryo homogeneous.

According to Lundell the large fruit is edible and was undoubtedly
utilized by the ancient Maya. The species is extremely well marked
for a member of this genus, but seems to be an endemic of very lim-
ited distribution, never having been reported except through the
specimens cited in the protologue.

PIMENTA Lindley. Allspice.

Calyx-lobes 4, distinct, imbricated in bud and in flower, persistent on the
fruit; hypanthium not or scarcely produced beyond the summit of the ovary;
petals 4. Inflorescence a many-flowered panicle [i.e. several times compound;
the central axis, and those of the principal branches of the panicle, prolonged
and bearing several pairs of lateral branches]. Ovary bilocular, the ovules 1 (-2)
in each locule. Embryo involute-cyclic, forming a double spiral. Fruit fleshy,
verrucose, usually 2-seeded. Whole plant very fragrant.

The genus consists of a single species.

Pimenta dioica (L.) Merrill, Contr. Gray Herb. 165: 37, /. 1.
1947. Myrtus Pimenta L. Sp. PL 472. 1753. M. dioica L. Syst. Nat.
ed. 10. 1056. 1759. Pimenta officinalis Lindl. Collect. Bot. sub 1. 19.
1821. Myrtus Tabasco Schlecht. & Cham. Linnaea 5: 559. 1830 [as
to Mexican specimens only, not as to type]. Eugenia micrantha
Bertol. Nov. Comm. Acad. Bonon. 4: 422. 1840 (type from some-
where in Guatemala, J. Velasquez). Pimenta officinalis [Berg], c
Tabasco sensu Berg, Linnaea 27: 425. 1856 [not Myrtus Tabasco
Schlecht. & Cham.]. P. Pimenta (L.) Karst. Deut. Fl. Pharm. Med.

McVAUGH: FLORA OF GUATEMALA 383

Bot. 790. 1882. Myrtus piperita Sesse* & Moc. Fl. Mex. ed. 2. 124.
1894 (the type from Papantla, Veracruz, Sesse et al. 2043) . Pimenta
dioica var. Tabasco sensu Standl. Ceiba 3: 172. 1953 [not Myrtus
Tabasco Schlecht. & Cham.]. P. Tabasco sensu Lundell, Wrightia 2:
58. 1960 [not Myrtus Tabasco Schlecht. & Cham.]. Pimiento; Pimi-
enta gorda; Pimienta; Peensia (Coban, Quecchi?) ; Pimienta de Chi-
apas; Pens (Quecchi) ; Ixnabacuc (Pete*n, Maya, fide Lundell) ; Pimi-
enta de Jamaica.

Common in moist or wet climax forest, usually on limestone,
350 meters or less; Pete*n; Alta Verapaz; cultivated commonly in
fincas of Guatemala, or sometimes for ornament or for its fruit in
parks, along streets, or about dwellings, at low and middle elevations.
Southern Mexico, including the Yucatan Peninsula of Mexico; Brit-
ish Honduras; Central America; West Indies; perhaps northern
South America.

A tree up to 20 meters high, 30 cm. in diameter, with pale brown bark peel-
ing off in thin scales or rather large sheets; vigorously growing branchlets flat-
tened and 4-angled, the angles terminating distally in the position of stipules;
branchlets, inflorescence and young foliage closely appressed-pubescent with
sordid or yellowish- white hairs, the hypanthium conspicuously canescent; leaves
coriaceous, soon glabrate, ovate or elliptic, 3-9 cm. wide, 9-20 cm. long, mostly
2-3 times as long as wide; blades variable in shape, generally acute but the very
tip obtuse or subacuminate, the base acute to rounded or even cuneate, the margins
at base even on the broader leaves cuneately long-decurrent on the stout broadly
channeled petioles 1.5-2 (-3) cm. long, 1.5-2.3 mm. in diameter; mid vein deeply
sulcate above, elevated its whole diameter beneath; lateral veins 9-12 pairs,
distant, rather prominent beneath, stronger than the few intermediate ones,
somewhat diminished distally and forming a connecting series of high irregular
angular arches the bases of which may be as much as 1 cm. from the margin;
minor veins forming a pale elevated network on the lower surface, the numerous
glands conspicuous among them; inflorescence a myrcioid panicle 6-12 cm. long,
many-flowered, 3 to 4 times compound, the peduncle 2-4.5 (-7) cm. long, up
to 2.5 mm. wide below the first node; flowers mostly clustered near the tips,
sessile but the lateral ones on short branches and so appearing pedicellate; bracts
subtending the flower-clusters 1-2 mm. long, deciduous at anthesis; buds turbinate,
2.5-3.5 mm. long; calyx lobes 4, nearly equal, broadly rounded, concave, thinly
pubescent without, densely canescent-tomentulose within, about 2 mm. wide, 1.5
mm. long; disk cup-shaped in flower, 2.5 mm. wide, the stamens interspersed with
white hairs; style 4-5 mm. long, prominently capitate, the stigma twice as thick
as the style; stamens about 150, up to 5 mm. long; petals white, suborbicular,
about 5 mm. long; ovary bilocular, the ovules 1 (-2) in each locule; fruit sub-
globose or oblate to somewhat pyriform, (4-) 6-8 mm. in diameter, prominently
verrucose with convex oil-bearing glands; seeds usually 2, laterally compressed,
suborbicular, the embryo forming a double spiral.

In Jamaica the production of allspice has been an important in-
dustry. Those who regard the plant of Mexico and Central America

384 FIELDIANA: BOTANY, VOLUME 24

as a species distinct from the West Indian one point out that the
commercial product from Mexico is held to be inferior to that of
Jamaica; that the dried fruits of the mainland plants are larger than,
and contain less than half as much volatile oil as, the Jamaican
berries. It is apparently true that the West Indian plants have
somewhat smaller flowers and fruits, and more slender panicles, than
those of Central America, but in other respects they are scarcely
distinguishable. The Central American populations have passed
under a series of names based nomenclaturally on Myrtus Tabasco
Schlecht. & Cham. Linnaea 5: 559. 1830, but that name applied to
a Venezuelan plant which is probably not the same as the Central
American ones. For a note on the nomenclature of Myrtus Tabasco,
see Fieldiana, Bot. 29: 511-512. 1963.

It is interesting to note that this species, as suggested by its name,
"dioica," is popularly supposed to produce male and female flowers
upon different trees. This is, in fact, the origin of the Linnaean name
"dioica," Linnaeus in 1754 having received from Philip Miller Ja-
maican specimens and a letter asserting the existence of the two
types of flowers. Some botanists have denied vigorously the exist-
ence of two types of flowers in the tree, whereas others have stated
categorically that the trees were polygamous or dioecious. Merrill
(Contr. Gray Herb. 165: 30-38. 1947), in an article on the nomen-
clature of Pimenta dioica, reviewed a series of statements by horti-
culturists and others who knew the tree in the living condition; the
evidence was somewhat contradictory, but the consensus was that
some individual trees definitely are barren at least in certain years
and perhaps always; that barrenness may result from damage to the
plant because of wasteful methods of harvesting; that even the so-
called "male" flowers are usually perfect, with well-developed sta-
mens and pistil. Merrill suggested that barrenness in at least some
individuals might result from a combination of genetically controlled
factors producing seasonally late flowering and at the same time
self -sterility.

The notes in the following paragraph are those of Mr. Standley:
This is one of the most delightful of Central American trees because
of the intense and highly agreeable odor that it exhales, whether fresh
or dry. Most plants with characteristic odor lose it upon drying, but
allspice specimens in the herbarium retain their fragrance indefinitely,
and to such a degree that it is very apparent whenever the case con-
taining the specimens is opened. The name allspice is derived from
the fact that the spice furnished by the tree, consisting of the dried

McVAUGH: FLORA OF GUATEMALA 385

unripe berries, is supposed to combine the flavor of cloves, cinnamon,
and nutmeg. This fruit is much used as a condiment for flavoring
food in most American and European countries, and it is sold com-
monly in Guatemalan markets. It is also employed to some extent
in domestic medicine. In Guatemala the Indians often apply the
powdered seeds to the corpses of children, saying that thus they are
preserved indefinitely, a practice that probably is of very ancient
origin. The trees may be seen in most of the small fincas of the
country, easily recognizable even when sterile by the characteristic
odor and by the distinctive bark, much like that of the guava. It is
said that the bark is shed more or less regularly twice a year, but it
always is peeling from the trunks, giving them a somewhat mottled
appearance. When covered with its abundance of white flowers the
tree is very handsome, conspicuous from a long distance, but the
flowers last only a few days at most. The wood is tough and close-
grained; heartwood reddish brown, the sapwood of a lighter color;
of medium luster, very hard and heavy, fine-textured; finishes very
smoothly. Probably no use is made of it locally.

PSIDIUM L.

References: Berg, Linnaea 27: 352-378. 1856; and in Mart. Fl.
Bras. 14, pt. 1: 381-411. 1857; McVaugh, R., Fieldiana, Bot. 29:
512-529. 1963.

Calyx various, usually splitting irregularly down to the ovary in anthesis,
its tube (hypanthium) prolonged beyond the summit of the ovary. Calyx-limb
of 4-5 distinct teeth or lobes, or completely closed in the bud, or with an apical
pore. Ovary 2- to 7-locular, the sporophores appearing to arise from the central
axis of the ovary, mostly connate in pairs and simulating stalked axile bilamellate
placentae. Petals often showy. Fruit often fleshy, many-seeded, sometimes
large and edible. Seed hippocrepiform or reniform, with bony testa and uncinate
or curved embryo. Inflorescence (in Guatemalan species) a 3-flowered dichasium,
or the flowers solitary.

An American genus which according to Berg includes about 100
species. The type species, P. Guajava L., the guava (Sp. guayabo),
is now widely naturalized throughout the tropics. A few species of
somewhat doubtful status, not treated below, are reported from
Mexico and from eastern Central America (Costa Rica).

Flowers at leafless nodes on old wood, sessile or slender-pedicellate, solitary or
several at the same node in sessile glomerules or irregular groups; leaves sub-
sessile or the petioles very short and stout, 1.5-2 mm. thick, 6 mm. long or
less; leaves glabrous or essentially so at maturity, with rather numerous
(10-15 or more) and parallel lateral veins on each side of the midvein; mid-

386 FIELDIANA: BOTANY, VOLUME 24

vein narrowly impressed on the upper surface; lowlands of the Atlantic Coast,
Honduras to British Honduras.

Branchlets acutely 4-angled, the angles becoming winglike in the distal parts
of the internodes and terminating in the stipular positions at the sides of
the petioles; flowers sessile or nearly so, the pedicels 1 mm. long or less;
ovary bilocular.
Plants, including the inflorescence, glabrous or essentially so.

Leaves subsessile, rounded to acute at base, 3-7 (-12) cm. long, mostly
1.6-2.3 times as long as wide; style 6-8 mm. long; stamens 60-75.

P. biloculare.

Leaves on petioles 3-5 mm. long, rounded or subauriculate at base, 12-
20 cm. long, 3-3.5 times as long as wide; style 9-10 mm. long; stamens

about 200 P. musarum.

Young leaves and branchlets, and the inflorescence, coarsely silky- or veluti-
nous-tomentose; leaves subsessile, abruptly rounded or subcordate at
base, 6-10 cm. long, 2-3 times as long as wide; style 9-10 mm. long;
stamens about 125 P. apodanthum.

Branchlets terete or compressed, the nodes enlarged; plants glabrous; flowers
pedicellate; calyx closed in the bud or with a small apical opening; style
5.5-7 mm. long.

Petioles 3-6 mm. long; pedicels 10-18 mm. long; buds 5-6 mm. long, apicu-
late, closed except a minute subapical pore; locules of the ovary 3 (-2),
the ovules about 20 (-45) in each locule P. anglohondurense,

Petioles 1-3 mm. long; pedicels 3-4 mm. long; buds 3-4.5 mm. long, sub-
globose or pyriform, the calyx-lobes united their whole length but leav-
ing an apical opening 1.5-2 mm. across; locules 2 (-1), the ovules about
50 in each locule P. mouririoides.

Flowers solitary or the peduncles 3-flowered, from the axils of foliage leaves on
developing branchlets, or sometimes in the axils of small bracts at the lower
nodes of developing leafy branchlets; lateral veins usually fewer than 10 on
each side, often 5-8; plants of middle or sometimes low elevations, often cul-
tivated or in disturbed habitats.

Lateral veins 12-20 on each side of the midvein, usually impressed above, prom-
inent beneath and well differentiated from the small intermediate veins;
herbage heavily pubescent especially when young with soft silvery-gray or
pale reddish hairs; branchlets somewhat 4-angled or sometimes terete; buds
closed, 10-16 mm. long; peduncles usually 1-flowered P. Guajava.

Lateral veins 5-10 on each side of the midvein, usually narrow and convex
above, if more than 10 then very slender and scarcely differentiated (on
the lower surface) from the intermediate veins; pubescence, branchlets and
flowers various.

Lower leaf-surface hirsutulous or tomentose, the numerous hairs 0.5-1 mm.

long, often obscuring the surface; inflorescence often 3-flowered.
Branchlets strongly 4-angled; leaves gray-tomentose beneath; lateral veins
5-9 on each side, ascending and parallel; flowers and buds unknown.

P. Xhypoglaucum.

Branchlets terete to compressed; herbage pale- or rufous-tomentose; lateral
veins 6-10 on each side, arcuate-ascending, incurving toward the tips;
buds 10-12 mm. long, closed or very nearly so at the tips.

P. guineense.

Lower leaf-surface glabrous or sparingly pubescent or strigose; inflorescence
usually 1-flowered.

McVAUGH: FLORA OF GUATEMALA 387

Leaves cuneate-obovate, mostly 2-5 cm. wide and 5-10 cm. long, obtuse
or rounded or abruptly deltoid-acuminate at apex.

Branchlets compressed or terete; flower-buds always glabrous, about
8 mm. long, the calyx very shortly 4-lobed; pedicels stout, 1 mm.
thick, 4 mm. long (up to 1.5 cm. long in fruit); style 6 mm. long,
the broad peltate stigma 1.3-1.5 mm. wide; lateral veins 8-10 on
each side; midvein flat or slightly impressed above; cultivated.

P. Cattleianum.

Branchlets strongly 4-angled; flowers unknown; peduncles 1- or 3-flow-
ered, 1.5-2 cm. long; lateral veins 4-6 on each side; small veinlets
forming a prominent reticulum; midvein convex above; probably a
hybrid of unknown ancestry; see P. chrysobalanoides (p. 396), in
discussion of P. guineense.

Leaves elliptic to ovate or somewhat obovate, rounded or acute at base.

Leaves mostly elliptic, about equally narrowed to both ends, the blades
5-12 cm. long, the petioles 5-8 mm. long; midvein impressed or
sometimes merely concave above; buds 7-12 (-20) mm. long, closed,
or the tips of the calyx-lobes minute, pressed together and simulating
an apical pore; anthers 1.5-2 mm. long.

Plant completely glabrous, a shrub 60 cm. high or less; branchlets
terete or compressed; leaves acute, usually cuspidate; buds 7-10
mm. long, the free tips of the calyx-lobes less than 0.5 mm. long;
style 8-9 mm. long; fruit 1.5 cm. in diameter P. Popenoei.

Branchlets and inflorescence sparingly appressed-pubescent; branchlets
often 4-angled; leaves acuminate; buds 8-12 (-20) mm. long, com-
pletely closed; style 13-15 mm. long; fruit up to 3-5 cm. long and
wide P. Friedrichsthalianum.

Leaves ovate-lanceolate to ovate or elliptic, mostly 3-6.5 cm. long, the
petioles 1-4 mm. long; midvein flat or somewhat convex above;
buds 4-7 mm. long, closed or the calyx 4- or 5-lobed; anthers 0.4-
0.8 mm. long.

Calyx openly 5-lobed, the lobes in mature buds broadly rounded or
deltoid, 2.5-3 mm. wide, about half as long as the globe of the
unopened petals; style 7-8 mm. long; leaves nearly sessile, the
stout subpetiolar base 1-2 mm. long or less; a shrub or subshrub
mostly less than 1 meter high P. salutare.

Calyx completely closed in the bud and then calyptrate and circum-
scissile, or with 4 minute free lobes surrounding an apical pore,
then opening by longitudinal fissures; style 4-5 mm. long; leaves
with definite slender petiole 1.5-4 mm. long; a tree up to 12-15
meters high P. Sartorianum.

Psidium anglohondurense (Lundell) McVaugh, Fieldiana, Bot.
29: 521. 1963. Eugenia Schippii Standl. Field Mus. Bot. 11: 137.
1932, not Psidium Schippii Standl., 1931. E. anglohondurensis Lun-
dell, Wrightia 2: 123. 1961.

Wet mixed forest or open pine forest, mostly near sea level; Brit-
ish Honduras, chiefly in pine forest, the type from Commerce Bight,
Stann Creek District, P. H. Gentle 8354.

388

FIELDIANA: BOTANY, VOLUME 24

FIG. 54. Floral morphology in Psidium. A, Bud of P. mouririoides (Gentle
4142, type); X 5. B, Axillary flower of P. salutare, showing open lobed calyx
(Haught 2602); X 2.5. C, Diagrammatic section of seed of P. guineense, showing
curved embryo unshaded in center; X 10. D, Bud and old flower of P. anglo-
hondurense (Bartlett 13060); X 5. E, Lateral and terminal views of bud and old
flower of P. biloculare (Gentle 3382); X 5.

A tree, up to 7-8 meters high, 25 cm. in diameter, nearly glabrous, the vege-
tative buds sparingly reddish-strigose; bracts, bracteoles and petals ciliate, and
the petals somewhat appressed-pubescent; staminal ring densely pubescent about
the bases of the stamens with short red hairs; branchlets terete or compressed,
with enlarged nodes; leaves elliptic, 3-5.5 cm. wide, 9-14 (-20) cm. long, 2.5-3
(-4.2) times as long as wide, gradually acuminate, the basal margins forming
an acute angle or cuneately narrowed, usually decurrent on the very stout adaxially
concave petioles 1.5-2 mm. thick, 3-6 mm. long; petioles in age transversely
rimose, the papery outer layers flaking off; midvein impressed above; lateral
veins rather numerous and parallel, 10-15 principal pairs in addition to some

McVAUGH: FLORA OF GUATEMALA 389

intermediate ones, visible in dried leaves as fine lines on both surfaces, often
slightly etched in on the upper surface; marginal vein evident, 2-3 mm. from
the margins, about equaling the laterals but scarcely arched between them;
blades dull green above, paler beneath, the minute glands scarcely visible above,
less abundant but evident beneath; flowers at leafless nodes of 2- to 3-year-old
twigs, 1-4 at a node, the pedicels 10-18 mm. long, 0.6 mm. in diameter (to 1 mm.
in fruit), seeming to arise directly from the axillary region above the leaf-scar;
buds 5-6 mm. long, apiculate, obovoid or constricted at the middle, closed except
for a small ciliate subapical pore, the calyx splitting irregularly into 3-4 ovate
or strap-like segments 3-5 mm. long, these up to 4 mm. wide, appressed-pubescent
within near base; bracteoles scarious, broadly ovate-triangular, 0.6-1 mm. long
and wide, deciduous about as the flower opens or somewhat persistent; disk 4-5
mm. wide, the center glabrous; style 6-7 mm. long, glabrous, subcapitate; stamens
ca. 175, 6 mm. long, the anthers nearly 1 mm. long; petals white, 6 mm. wide,
8 mm. long; ovary 2- or 3-locular, the ovules about 20-45 in each locule, in a
subcapitate group from an axillary bilamellate placenta; fruit "blue-black"
(Schipp), globose, 1-1.5 cm. in diameter, the hard flesh 2-2.5 mm. thick when
dry; seeds chocolate-brown, slightly rough, often 2 developing in a locule and
then plano-convex, 7-8 mm. wide, curved into a nearly complete ring, the outer
layer simulating a membranaceous testa but not separating from the bony inner
layer surrounding the ring-like embryo more than 10 mm. long, 0.5 mm. in di-
ameter.

Psidium apodanthum (Standl.) McVaugh, Fieldiana, Bot. 29:
520. 1963. Eugenia apodantha Standl. Field Mus. Bot. 17: 380. 1938.

Lowlands, forests, probably along rivers, to be sought in eastern
Guatemala. Honduras (Dept. Comayagua, near Siguatepeque,
Yuncker et al. 5766, the type).

A tree to 5 meters high, 7.5 cm. in diameter, the branchlets sharply 4-angled,
the distal portions of the internodes bearing corky wings 'in the stipular positions;
bark of older branchlets pale, checking in thin longitudinal small plates; young
branchlets and petioles, and lower surfaces of young leaves, coarsely velutinous-
tomentose with pale brown nearly erect sharp hairs up to 0.5-0.7 mm. long;
flowers silky-tomentose with similar hairs; leaves subsessile, elliptic or elliptic-
obovate, 2-4 cm. wide, 6-10 cm. long, 2-3 times as long as wide, the tips bluntly
pointed or obscurely acuminate, the blades narrowed toward the base from the
middle or below, then abruptly rounded or subcordate, the margins very shortly
decurrent on the inner angles of the corky-rimose petiole 1.5-2 mm. thick, 2-2.5
mm. long; midvein deeply and narrowly impressed above; lateral veins 10-12
pairs in addition to some intermediate ones, inconspicuous above, rather prom-
inent beneath; marginal veins about equaling the lateral ones and slightly arched
between them, 1.5-2.5 mm. from the margin; blades obscurely glandular, some-
times punctate above; flowers sessile, 1-4 together in axillary clusters at leafless
nodes, subtended by flat scarious bracts; buds obovoid, about 5 mm. long, broadly
rounded at apex, the conic densely hairy canescent hypanthium 1.5 mm. long;
calyx-lobes free from the first, bluntly ovate or linguiform, 3 mm. long, 2.5 mm.
wide at base, tomentulose without, glabrous or sparingly tomentulose within,
at anthesis separating down to the disk and then 4-6 mm. long, 2.5-4.5 mm.

390 FIELDIANA: BOTANY, VOLUME 24

wide, flat; disk 4-5 mm. wide, hairy in the staminal ring and about the base of
the style; style (6.5-) 9-10 mm. long; stamens about 125, about as long as the
style; petals white, obovate, 7 mm. long, delicately ciliate; ovary bilocular, the
ovules about 20 in each locule, attached in a sessile subcapitate group above
the middle of the central dissepiment; immature fruit crowned by the four some-
what irregularly shaped but nearly equal calyx-lobes; immature seeds suborbicular-
reniform, laterally compressed, about 3 mm. wide, more than 1 mm. thick; testa
bony; embryo C-shaped.

Psidium biloculare McVaugh, Fieldiana, Bot. 29: 520. 1963.
Eugenia Gentlei Lundell, Carneg. Inst. Wash. Publ. 478: 216. 1937.
Not Psidium Gentlei Lundell, 1943.

River banks, at low elevations; Izabal (Rio Chacon, H. Johnson
1191). British Honduras, in pinelands (the type from Gracie Rock,
Sibun River, Belize District, P. H. Gentle 1684).

Shrub or small tree to 20 cm. in diameter, glabrous except for a few coppery
stiff hairs up to 0.4 mm. long on the inflorescence, on the vegetative buds and
at the youngest growing nodes; branchlets, at least on the distal parts of the inter-
nodes, sharply wing-angled, the 4 corky wings terminating in the stipular positions
at the bases of the petioles; leaves coriaceous, elliptic or especially on young shoots
ovate, (1-) 1.5-2.5 (-4) cm. wide, (2-) 3-7 (-12) cm. long, 1.6-2.3 (-3) times
as long as wide, broadly acuminate or merely obtuse at tip, gradually rounded
to an acute base (abruptly rounded on shoots) ; blades almost sessile, the margins
decurrent on the channeled petiolar base and often apparently extending into
the axil, the base itself becoming corky, as seen from the abaxial side, 2-4 mm.
long, up to 2 mm. thick; midvein impressed above; lateral veins about 10 pairs
in addition to some intermediate ones, obscure on both surfaces; marginal vein
obscure, about equaling the laterals and but slightly arched between them, ca.
1 mm. from the margin; blades dark green and probably lustrous above, paler
beneath, the upper surface with numerous minute impressed glandular dots,
the lower sparingly but evidently dark-dotted; flowers closely sessile, 1-2 together
at leafy or leafless nodes, subtended by a pair of scarious rounded ciliate bracteoles
about 0.8 mm. long and wide; bracts about the same size, scarious; buds pyriform,
4-5 mm. long, the small hypanthium conic; calyx-lobes 4, the free terminal parts
rounded or ovate, 1 mm. long, 1-1.5 mm. wide at base, the calyx as a whole splitting
at an thesis down to the margins of the disk and the lobes becoming 2-2.5 (^4)
mm. wide at base, 2.5-4 (-5.5) mm. long; disk 2.5-4 mm. wide, glabrous or
the staminal ring hairy; style 6-8 mm. long, subcapitate; stamens ca. 60-75,
up to 8 mm. long; anthers 1 mm. long; petals rather narrowly obovate, ciliate,
5 mm. wide, 7 mm. long; ovary bilocular, the ovules about 20-30 in each locule,
in a subcapitate group arising from the central dissepiment; fruit globose, oblate
or somewhat pyriform, up to at least 1-1.2 cm. long and wide; seeds 1-2 in each
locule, laterally compressed and hippocrepiform, 4-6 mm. wide, 1.5-2.5 mm.
thick, notched at one side, the outer layer of the seed coat membranous, the inner
bony; embryo C-shaped.

This species is difficult to distinguish from P. apodanthum except
by the presence or absence of pubescence. It may be that additional

McVAUGH: FLORA OF GUATEMALA 391

distinguishing features reside in the buds or flowers, but in P. apo-
danthum neither of these is known. Certain specimens from British
Honduras (Gentle 5358), and from Izabal (Steyermark 38544) seem
to be intermediate between the two species in having the calyx
tomentulose but the plants otherwise essentially glabrous.

Psidium Cattleianum Sabine, Trans. Roy. Hort. Soc. 4: 315,
1. 11. 1821. P. Morale Raddi, Opusc. Sci. Bologna 4: 254, t. 7, /. 2.
71821. Guayaba; guayaba japonesa. Strawberry guava.

Cultivated around Coban (about 1300 meters) and doubtless
elsewhere in Guatemala; planted infrequently in Central America.
Native probably of the lowlands of eastern Brazil.

A shrub or small tree, quite glabrous even to the vegetative buds, or the
branchlets, leaves and pedicels minutely hispidulous-puberulent, the flower buds
always glabrous; leaves cuneate-obovate, 1.5-6 cm. wide, 4-12 cm. long, 1.75-3
times as long as wide, the tip obtuse or rounded or sometimes obscurely acuminate,
the base cuneate or sometimes acute, the margins decurrent on the rather short
petioles 3-10 (-20) mm. long; midvein flat or slightly impressed above; lateral
veins 8-10 on each side, inconspicuous, ascending and parallel, connected by a
series of asymmetric angular arches forming a somewhat poorly defined marginal
vein 1-3 mm. from the margin; flowers solitary and axillary, or at the lowermost
leafless (bracteate) nodes of otherwise leafy branchlets, or rarely 2-3 pairs in
raceme-like groups on short leafless branchlets; pedicels 4 mm. long (up to 8-15
mm. long in fruit), 1 mm. thick; bracteoles lance-ovate, 2.5 mm. long, ?deciduous;
buds pyriform, about 8 mm. long, open at the end, the calyx with about 4 broad
low rounded lobes, splitting in the sinuses into irregular lobe-like segments 5-6
mm. long and wide; style 6 mm. long, much narrower than the capitate eroded
stigma 1.3-1.5 mm. wide; petals white, obovate; fruit obovoid or globose, 2.5-
3.5 cm. long, purplish red or sometimes yellow, with a thin skin, the pulp white,
sweet and aromatic, containing numerous hard compressed-triangular seeds 2-2.5
mm. long.

Called "cas dulce" in Costa Rica. For an account of the nomen-
clature of this species see F. R. Fosberg, Proc. Biol. Soc. Wash. 54:
179-180. 1941; C. A. Schroeder, Journ. Arnold Arb. 27: 314-315.
1946.

Apparently native plants from eastern Brazil, especially near the
coast from Parana to Sao Paulo, are usually glabrous; cultivated
specimens from northern South America and Central America are
often pubescent, and the leaves sometimes suggest by their size
and shape a possible admixture of genes from P. Sartorianum.

Psidium Friedrichsthalianum (Berg) Ndzu. in Engl. & Prantl,
Nat. Pflanzenfam. 3, Abt. 7: 69. 1893. Calyptropsidmm Friedrichs-
thalianum Berg, Linnaea 27: 350. 1856.

392 FIELDIANA: BOTANY, VOLUME 24

Along the edges of tidal streams (Honduras), in forests, or in up-
land pastures or clearings, sometimes in cultivation; Santa Rosa
(Santa Rosa, 900 m., Heyde & Lux 2984). Southern Mexico (Chi-
nantla, Liebmann 3993) ; Central America to Panama (the type col-
lected at Granada, Nicaragua — not Guatemala as stated in the
original description — by Friedrichsthal) ; Colombia.

A tree 6-10 meters high, the branchlets and inflorescence sparingly appressed-
pubescent with sordid hairs; leaves glabrate; branchlets often 4-winged, the wings
terminating distally in the stipular positions; leaves 2.5-5 cm. wide, 5-12 cm.
long, 1.6-2 (-3) times as long as wide, mostly elliptic, about equally narrowed
to the acuminate tip and the broadly acute or rounded base, the margins cuneately
decurrent on the thin inner angles of the deeply channeled petiole 5-8 mm. long;
midvein deeply and narrowly impressed; lateral veins 5-8 on each side in addition
to some intermediate ones, ascending, diminishing distally, the marginal vein
inconspicuous but well defined, arching, 1.5-4 mm. from the margin; blades when
young membranaceous and resinous-dotted, at maturity sometimes gland-dotted,
the midvein on the lower surface bearing prominent convex glands; flowers solitary
(rarely with sessile central flower and a pair of pedicellate lateral flowers forming
a dichasium), axillary or in 1-2 bracteate pairs at the lower (leafless) nodes of
leafy branchlets; bracteoles narrow, deciduous, 1.5 mm. long; pedicels 10-20 mm.
long, 1-1.3 mm. thick; buds closed, mostly ovate or obovate, 8-12 mm. long,
acute at the base, broadly rounded at the apex and blunt or apiculate, breaking
irregularly through the apex or near it into 1-3 broad lobe-like or hooded seg-
ments [buds sometimes pyriform, 15-16 (-20) mm. long including the narrowly
obconic hypanthium 5 mm. long, the subglobose body 8-9 mm. long and the
subulate apiculum 2 mm. long; pedicels in these buds up to 3-4 cm. long]; style
13-15 mm. long, the stigma much wider than the style, 1.5 mm. wide, flat or
concave; stamens very numerous, probably about 300, up to 12 mm. long; anthers
linear-oblong, 1.6-2.1 mm. long; petals about 15 mm. long; fruit globose or oval,
up to 3-5 cm. long and wide, yellow, acid; seeds laterally compressed, 5-6 mm. long.

Called "arrayan" (El Salvador) ; "guayaba de danto" or "guay-
aba agria" (Honduras); "cas" (Costa Rica); "guayaba de agua"
(Panama). Mr. Standley states that although the fruit is edible it
is so sour that it is not agreeable to most people.

Psidium Guajava L. Sp. PI. 470. 1753. Guayaba (fruit) ; guayabo
(plant) ; pataj, paid (Quecchi) ; cac (Poconchi) ; ch'amxuy (Ixil) ; piac
(Cacchiquel, Antigua) ; ikiec (Cacchiquel, Tecpam) .

Mostly in moist or dry thickets, especially in pastures, frequently
forming almost pure stands of considerable extent, 1800 meters or
less, most common at 1000 meters or lower; planted generally; prob-
ably found in every Department of Guatemala. Florida; Mexico;
Central America; Costa Rica and Panama; West Indies; tropical
South America; naturalized in the tropics of the Old World.

McVAUGH: FLORA OF GUATEMALA 393

A shrub or tree up to 10 meters high, pubescent on the young growth, and
the inflorescence and at least the lower surface of the leaves sparingly or densely
pubescent with mostly appressed, soft silvery-gray or pale reddish hairs up to
about 0.5 mm. long; branchlets terete, or quadrangular with pronounced angles
or low wings below the nodes; leaves elliptic or oblong, 3-6 cm. wide, (4.7-) 8-14
cm. long, mostly 2-3 times as long as wide, rather abruptly rounded or occasionally
narrowed to the obtusely pointed or rounded tip, the base abruptly rounded or
subcuneately narrowed, the margins decurrent on the stout petiole 1-2 mm.
thick, 4-7 (-10) mm. long; veins prominent beneath, usually markedly impressed
above; lateral veins 12-20 pairs, often conspicuous and parallel, not forming a
marginal vein but diminishing distally, each approaching to within 1-2 mm.
from the margin, then incurving and joined to the next succeeding vein by a
strong straight perpendicular vein; upper surface of blade inconspicuously gland-
dotted, glabrous or thinly pubescent, the small veins little or not at all elevated
and scarcely apparent; lower surface finely appressed-pubescent or glabrous,
thickly gland-dotted; inflorescence axillary, 1-flowered (very rarely 3-flowered),
the peduncle (7-) 10-20 (-25) mm. long, 1 mm. or more thick, up to 2.5 mm.
thick in fruit; bracteoles subulate, often glabrous, 3-4 mm. long; buds at maturity
13-16 mm. long, completely closed at tip, the hypanthium 5-7.5 mm. long, con-
stricted at summit and thence enlarged into the ovoid pointed calyx; calyx at ma-
turity splitting irregularly into 4-5 lobes which are appressed-pubescent distally;
proximal third of the calyx, and the summit of the ovary, glabrous; style 10-12
mm. long, the flat peltate stigma 0.5 mm. wide; petals white, elliptic, cucullate,
10-12 mm. long; stamens 1507-275, the longest as long as the style; anthers
0.8-1.2 mm. long; fruit globose or pyriform, 2-6 cm. long, yellow or pinkish.

The following notes were supplied by Mr. Standley: The Maya
name of Yucatan is "pichi"; sometimes called "coloc" and "putah"
in British Honduras, and the name "pichi" also is used there by
Maya-speaking people. The name "guayaba" is of Antillean origin.
It occurs in many place names of Guatemala, such as El Guayabo,
Los Guayabales, etc. The tree is a very common one in the low-
lands of Guatemala, often forming pure stands called gua/y abates.
The fruits are eaten by many animals, and it is probably on this
account that the seedlings are spread so widely, the hard seeds being
indigestible. It is noteworthy that the young plants spring up in
perhaps greatest abundance in cattle pastures. The wood is brown
or light brown. It is little used except for firewood, but in Guate-
mala spinning tops sometimes are made from it. In El Salvador it
is utilized for fine-toothed combs, but these, when wet, are inclined,
even when well polished, to disintegrate or shed their fibers; hence
the wood of Salamo (Calycophyllum) is preferred for the purpose.
The bark is employed in some parts of Mexico, and perhaps also of
Central America, for tanning leather. A decoction of the leaves is
astringent, and is administered as a remedy for diarrhea. The fruit
of the guava is highly esteemed in Guatemala, and in most other

394 FIELDIANA: BOTANY, VOLUME 24

regions where it grows, in spite of its very strong and not altogether
pleasant odor. Vast quantities of the fruits are eaten raw by chil-
dren and older people, but enough is left, so great is the production,
for a great variety of mammals and birds. In Guatemala the fruits
are not common in most of the markets, probably because most
people have an ample supply at or near home. The fruits are stewed
and often served as a dessert. Their most important use is in prep-
aration of the thick jelly or guava paste which, cut in thin squares,
is one of the favorite and most common desserts of all Central Amer-
ica. The fruits vary greatly in shape, size, and flavor, some being
rather acid.

Psidium guineense Sw. Prodr. 77. 1788. P. Araca Raddi,
Mem. 5, 1. 1 [Op. Sci. 4: 854?]. 1821. P. molle Bertol. Nov. Comm.
Acad. Bonon. 4: 422. 1840 (type collected in Guatemala by J. Velas-
quez). P. costa-ricense Berg, Linnaea 27: 368. 1856 (type from Irazu
["laru"], Costa Rica, Oersted 17). ?P. Schiedeanum Berg, I.e. (type
from Mexico, Schiede 541). P. laurifolium Berg, Linnaea 27: 364.
1856 (type from Masaya, Costa Rica, Oersted). ?P. monticola Berg,
in Oerst. Vid. Medd. Kjoeb. 1855: 11. 1856-57. Guajava mollis
(Bertol.) 0. Ktze. Rev. Gen. 1: 240. 1891. P. rotundifolium Standl.
Field Mus. Bot. 8: 318. 1931 (type from All Pines, British Honduras,
W. A. Schipp S85) . P. Schippii Standl. I.e. 319 (type from All Pines,
British Honduras, Schipp 595). Guayaba agria; guayaba acida;
guayaba hedionda; chamach or chamacch (Quecchi); pataj (Coban,
Quecchi) ; guayaba; arrayana; pichippul (Pete*n, Maya, fide Lundell).

Wet to dry thickets or open forest, often in oak or pine forest,
frequently on rocky open hillsides or plains, 2400 meters or less,
mostly at middle elevations; Pete"n; Alta Verapaz; Baja Verapaz;
El Progreso; Izabal; Zacapa; Chiquimula; Jalapa; Jutiapa; Santa
Rosa; Guatemala; Sacatepe"quez ; Chimaltenango; Quiche"; Huehuete-
nango; Quezaltenango. Mexico (Sinaloa to Oaxaca; San Luis Potosi
to Chiapas; Quintana Roo); British Honduras; Central America to
Panama; West Indies (Jamaica; Martinique; Trinidad); throughout
tropical South America to northern Argentina.

A shrub 1-2.5 meters tall or occasionally a small tree up to 7 meters high,
usually heavily pubescent on the young growth, the inflorescence, and at least
the lower surface of the leaves with erect, soft, flexuous, pale- or coppery-red hairs
up to 1 mm. long; branchlets nearly terete to moderately compressed, rarely with
definite angles below the leaf-bases; leaves broadly elliptic or sometimes obovate,
(2.7-) 4-6 (-8) cm. wide, (3.5-) 6-10 (-14) cm. long, (1-) 1.6-2.3 (-3) times as
long as wide, abruptly narrowed about equally to the rounded base and to the

McVAUGH: FLORA OF GUATEMALA 395

rounded or obtusely pointed apex, or narrowed gradually from above the middle
toward the often asymmetrical rounded or rarely subcuneate base; margins de-
current on the stout petiole 1-2 mm. thick, 5-10 mm. long; midvein slightly im-
pressed above; lateral veins 6-10 on each side, when dry convex on the upper
surface like the small veinlets, prominent beneath, strongly ascending, not form-
ing a marginal vein but each lateral decreasing distally, curving toward the tip
of the leaf, and inward, from a point 1-2 mm. from the margin, sometimes con-
nected to the next lateral by a strong straight perpendicular vein near the tip;
upper surface of leaf inconspicuously gland-dotted, usually somewhat pubescent
(at least with a line of reddish hairs along the midvein), often with numerous evi-
dent small veins which are slightly elevated when dry and form a pattern (readily
visible with the unaided eye) at right angles to the midvein and at an acute angle
to the lateral veins; lower surface persistently pubescent, thickly and conspicu-
ously gland-dotted; inflorescence an axillary dichasium, often 3-flowered but
1-flowered in some axils, or in some plants all 1-flowered; peduncle (12-) 17-32 mm.
long, 1-2 mm. wide near summit and there somewhat compressed; central flower
sessile or nearly so, the two lateral ones on pedicels 8-12 mm. long; bracts and
bracteoles linear or subulate, about 3 mm. long, or the bracts leafy and up to
1.5 cm. long; bracteoles often with abortive axillary buds; buds at maturity 10-
12 mm. long, incompletely closed at tip, the hypanthium 4-5 mm. long, constricted
at summit and thence enlarged into the obtusely pointed or nearly globose calyx;
calyx at maturity splitting irregularly into 4-5 lobes which are appressed-pubes-
cent distally; proximal third of the calyx, and the summit of the ovary, gla-
brous; style glabrous, 7-11 mm. long, the flat peltate stigma 0.5 mm. wide; petals
white, elliptic, cucullate, 10-14 mm. long; stamens about 150-200, as long as the
style or shorter; anthers 1.2-2.2 mm. long; fruit globose or pyriform, 1-2 cm. in
diameter, yellowish-green or yellow, barely edible.

A most variable species, usually a shrub but sometimes a small
tree, often confused in herbaria with the common and much culti-
vated guava, Psidium Guajava. It is probable that these two species
hybridize in nature, with the production of individuals intermedi-
ate in various ways. Ordinarily the two may be distinguished,
as noted in the key, by the average number of veins in the leaves,
by the degree to which the lateral veins are impressed, and by the
degree of prominence of the small veins of the leaves in dried speci-
mens. Additional useful distinguishing features may be found in
the number of flowers (usually 1 only in P. Guajava, often 3 in
P. guineense), the larger and more pointed and completely closed
buds of P. Guajava, the longer peduncles of P. guineense. A detailed
revisionary analysis in the field and in the herbarium, of the complex
centering around these two species would be a valuable contribution
to the taxonomy of tropical cultivated plants.

It is probable that the following species, Psidium Xhypoglaucum
Standl., has resulted from hybridizations between P. guineense and
P. Guajava. For notes on this and the other three species described

396 FIELDIANA: BOTANY, VOLUME 24

by Standley in 1931, from a single locality in British Honduras, see
Fieldiana, Bot. 29: 522-524. 1963. These four species were all based
on fruiting specimens; Standley in differentiating them emphasized
the differences in leaf-form but did not compare them to P. guin-
eense, which they all strongly resemble. The most distinctive is
P. chrysobalanoides Standl. (Field Mus. Bot. 8: 319. 1931), which is
nearly glabrous, with 4-angled branchlets and obovate-cuneate leaves
2.5-5 cm. wide, 4-7.5 cm. long. The proper taxonomic disposition
of this plant can only be determined by additional specimens.

According to unpublished notes by Mr. Standley, P. guineense
is a very common shrub in many parts of Guatemala, and is native
through most of the country. It is rarely if ever cultivated ; the fruits
are smaller than those of the edible guava, and too bitter or resinous
to be palatable. The specific name is an unfortunate one, given by
Swartz under the erroneous belief that the plant had been imported
into Santo Domingo from West Africa.

Psidium Xhypoglaucum Standl. Field Mus. Bot. 8: 320, pro
spec. 1931. P. guineense Sw. X P. Guajava L.

Open rocky slopes, brushy hillsides, meadows and pastures, some-
times in pine-oak forest, often in disturbed grounds, 1000 meters or
less; Jutiapa (Cerro Colorado, just west of Jutiapa, Standley 76173,
76190). Mexico (Chiapas); British Honduras, the type from All
Pines, W. A. Schipp S99; Honduras.

A shrub 40 cm. to 1 meter high, the branchlets thinly pale-pubescent and
strongly 4-angled; leaves densely and softly gray-tomentose beneath, the surface
obscured by the matted hairs; blades elliptic-obovate on shoots, often obovate
and cuneate at base; on slowly growing flowering branchlets often elliptic and
somewhat rounded at base; blades 2-5 cm. wide, 3.5-9 cm. long, 1.5-2 times as
long as wide, the petioles 3-6 mm. long; midvein impressed above; lateral veins
5-9 on each side, strongly ascending and parallel, forming with the veinlets, on
the upper surface of dried leaves, a reticulum very like that in P. guineense; in-
florescence 3-flowered, on a peduncle up to 3.5-5 cm. long, or in some plants
1-flowered, the peduncle 1-1.5 cm. long; flowers and buds unknown; fruit globose,
2-2.5 cm. in diameter, said to be sour.

Most individuals of this so-called species bear a strong super-
ficial resemblance to plants of P. guineense, but usually exhibit also
some features of P. Guajava, especially the gray pubescence of the
leaves and the 4-angled branchlets. The lateral veins of the leaves
suggest those of P. Guajava in being almost parallel, but are 5-9 in
number, suggesting in this respect P. guineense. The hybrid nature
of this plant is suggested by its morphological intermediacy and by

McVAUGH: FLORA OF GUATEMALA 397

its common occurrence in disturbed habitats where both of the sup-
posed parent species grow. Experimental confirmation of its hybrid
nature is much to be desired.

Psidium mouririoides (Lundell) McVaugh, Fieldiana, Bot. 29:
521. 1963. Eugenia mouririoides Lundell, Amer. Midi. Nat. 29: 480.
1943.

British Honduras, known only from the type, collected in ham-
mock on pine ridge, Toledo District, Monkey River, Jenkins Creek,
P. H. Gentle 4142.

A small tree, 12 cm. in diameter, glabrous, the branchlets terete or com-
pressed, with enlarged nodes; leaves coriaceous, broadly elliptic, 2-5 cm. wide,
4.5-10 cm. long, 1.6-2.5 times as long as wide, acuminate with depressed tip,
rounded at base, the margins decurrent on the very stout adaxially concave peti-
ole 1.5-2 mm. thick, 1-3 mm. long; midvein impressed above; lateral veins 10-15
pairs in addition to some intermediates, inconspicuous on both surfaces; marginal
vein about equaling the laterals, scarcely arched between them, 1-2 mm. from the
margin; blades dull green above, paler beneath, the upper surface with many
minute impressed glandular dots, the lower sparingly dark-dotted; flowers 1-3
together at leafless nodes, on slender pedicels 3-4 mm. long; expanding buds sub-
globose or short-pyriform, 4-sulcate, open at the tips, the calyx-lobes united their
whole length except at the convex, ciliate apical margins surrounding a rounded
or quadrate opening 1.5-2 mm. across; calyx at anthesis splitting longitudinally,
somewhat irregularly into 4 ovate lobes 3 mm. long and about as wide at base;
disk 4.5 mm. wide, the staminal ring hairy; style about 5.5 mm. long, scarcely
subcapitate; hypanthium broadly cup-shaped, closely invested by a pair of con-
nate-perfoliate scarious rounded bracteoles up to 1.5 mm. wide, 1-1.5 mm. long;
stamens about 125, up to 6 mm. long, the anthers 0.8 mm. long; petals suborbic-
ular, white, ciliate, about 6 mm. wide, up to 7 mm. long; ovary bilocular, the
ovules about 50 in each locule, ascending, in a subcapitate group arising from the
central dissepiment [this seen in one bud only; in 2 buds the ovary was unilocular,
with ca. 50 ovules attached laterally]; fruit not seen.

Called "wild guava" in British Honduras.

This species is presumed to belong to the genus Psidium because
of the characteristic way the opening calyx splits as the flower ex-
pands, and because the plant is extraordinarily like Psidium anglo-
hondurense, which grows in the same region. Final confirmation of
its generic status must await the discovery of fruiting specimens.
The unilocular ovary found in two out of three buds taken from the
type specimen of P. mouririoides may be an anomaly, or perhaps an
indication of evolutionary specialization in this species.

Psidium musarum (Standl. & Steyerm.) McVaugh, Fieldiana,
Bot. 29: 521. 1963. Eugenia musarum Standl. & Steyerm. Field
Mus. Bot. 22:358. 1940.

398 FIELDIANA: BOTANY, VOLUME 24

In wet forests, 350 meters or less; Izabal (Rio Juyama, about
15 miles southwest of Bananera, Steyermark 39165, the type) ; Alta
Verapaz.

Shrub 2.5 meters high, glabrous, the branchlets acutely 4-angled, the angles
terminating distally in the stipular positions; leaves elliptic or elliptic-lanceolate,
4-6.5 cm. wide, 12-20 cm. long, 3-3.5 times as long as wide, slenderly acuminate,
rounded or subauriculate at base, the cartilaginous margins abruptly decurrent
on the acute inner angles of the broadly concave petioles 3-5 mm. long; mid vein
narrowly impressed above; lateral veins 10-12 on each side in addition to some
intermediate ones, inconspicuous above, apparent beneath; marginal vein about
equaling the lateral ones and but little arched between them, 2-8 mm. from the
margin; blades dull dark green above, pale beneath; pale resinous glands some-
times apparent among the pale translucent cells of the lower surface; flowers soli-
tary or in pairs, in small sessile glomerules at leafless nodes, the pedicels less than
1 mm. long and about as thick as long; bracteoles rigidly coriaceous, obtuse, con-
nate, reddish, forming a boat-shaped cupule 2-2.5 mm. long, nearly 1 mm. wide
and high; buds not seen; hypanthium broadly campanulate or cup-shaped, in an-
thesis 2-3 mm. long and wide; calyx-lobes in bud free at the tips, the longer ones
ciliate for about 3 mm. from the tips; calyx splitting between the lobes in anthesis,
the splits extending 1-1.5 mm. below the sinuses, the lobes after anthesis rotate,
oblong-ovate, blunt, 2.5-4 mm. wide at base, 3-4 mm. long; disk 3-4.5 mm. wide,
quadrate, the petal-scars shallowly deltoid, prominent at the corners of the disk;
style 9-10 mm. long, tapering to a small capitate stigma; stamens about 200,
about as long as the style; anthers 0.7 mm. long; petals white, 4, broadly obovate
or suborbicular, 6-7 mm. long, ciliate; ovary bilocular, the ovules 5 in each locule,
pendent or radiating from a centrally located placenta, apparently flat or C-shaped;
fruit "dull blue" (Steyermark), globose or oblate, 10-12 mm. in diameter.

Psidium Popenoei Standl. Ceiba 1: 41. 1950.

Savannahs and pine forests, about 1000 meters or less; to be ex-
pected in eastern Guatemala. Honduras (Dept. Santa Barbara, on
Rio Chamelecon, Standley & Lindelie 7502; Dept. Comayagua, near
Siguatepeque, Standley & Chacdn 6369, the type) .

A shrub 50-60 cm. high, quite glabrous even to the young growth, the branch-
lets terete or compressed; leaves elliptic, about equally narrowed to the acute and
often cuspidate tip, and to the somewhat rounded or acute base, 2-4.7 cm. wide,
6-11 cm. long, about 2-3 times as long as wide; margins often rather long-decurrent
on the sharp inner angles of the broadly channeled petiole 5-8 mm. long; midvein
impressed above, sometimes merely concave; lateral veins 5-8 on each side, ascend-
ing, prominent on both sides but elevated and conspicuous beneath, much stronger
than the intermediate ones if any, diminishing distally and incurving about 1 mm.
from the margin to join the next succeeding ones; blades darker green and lus-
trous above, dotted beneath with many small resinous glands, sometimes im-
pressed-punctate above especially in age; flowers solitary in the axils of foliage
leaves, the peduncles compressed, slightly clavate in fruit, 1-2 cm. long, 1 mm.
thick; bracteoles not seen, falling before the buds mature and leaving a scar at
the base of the hypanthium; buds pyriform, 7-10 mm. long, obtuse, the calyx

McVAUGH: FLORA OF GUATEMALA 399

almost completely closed, its tip with a minute pore surrounded by the approxi-
mate tips of 4-5 incurved lobules less than 0.5 mm. long; calyx in anthesis break-
ing irregularly, longitudinally, splitting its whole length into 3-5 variously shaped
but mostly strap-like segments up to 6 mm. wide and 8 mm. long, the sinuses
finally extending to the margin of the staminal ring; disk 4 mm. wide, sometimes
5-angled, the angles marked by 5 prominent petal-scars, these lenticular or shal-
lowly triangular, 1.5 mm. long; style 8-9 mm. long, capitate; stamens very numer-
ous, about 300; petals white; ovary about 4-locular, the stalked bilamellate pla-
centae seeming to arise from the inner angles of the locules, bearing numerous
ovules; fruit globose, 1.5 cm. in diameter, the many straw-colored seeds with bony
testa, somewhat laterally compressed, 3-3.5 mm. long, the embryo C-shaped.

Called in Honduras "guayabillo," "guayabo," "arrayan," or
"huevo de gato."

Psidium salutare (HBK.) Berg, Linnaea 27: 356. 1856. Myrtus
salutaris HBK. Nov. Gen. & Sp. 6: 132 [folio ed. 105]. 1823. P. tili-
atum Benth. Hook. Jour. Bot. 2: 318, as to type [Schomburgk 365].
1840. P. salutare a stricta Berg, I.e. ?P. salutare /3 laxum Berg, I.e.
357. P. salutare 7 subalternum Berg, I.e. 357, at least as to lectotype
[Schomburgk 365]. Calycolpus parviflorus Sagot, Ann. Sci. Nat. VI.
20: 181. 1885. P. Gentlei Lundell, Amer. Midi. Nat. 29: 483. 1943.

Savannahs, pine ridges, and rocky or grassy hills, 1000 meters
or less. British Honduras, Toledo District, Monkey River, Jenkins
Creek, open pine ridge, P. H. Gentle 4062, type of P. Gentlei. Mexico
(Nayarit, Rose 1950; Chiapas); Central America; Panama; Colom-
bia to the Guianas; Isle of Pines; Hispaniola.

A small shrub or sub-shrub 30-50 (-100) cm. high from stout and often
gnarled woody roots, the whole plant glabrous (the youngest leaves sometimes
ciliate, the calyx-lobes ciliate and also appressed-pubescent within);1 leaves nearly
sessile, subcoriaceous, ovate-lanceolate to ovate or elliptic, 1-3 cm. wide, (2-) 3-
6.5 cm. long, 1.8-3.5 times as long as wide, the tip acute or acuminate, usually
appearing sharp-pointed to the unaided eye; base rounded and the margins rather
abruptly decurrent on the stout subpetiolar base 1-2 mm. long or less; midvein
flat or somewhat convex above; lateral veins inconspicuous, 5-10 pairs in addition
to some intermediate ones, arching into an ill-defined marginal vein 1-2 mm. from
the margin; blades dark green above, paler and often drying dull and brown be-
neath; glands usually evident beneath as minute dots; inflorescence axillary, rarely
3-flowered with the terminal flower sessile, the flowers usually solitary in the leaf-
axils or near the base of the branch in the axils of much reduced leaflike bracts;

1 Authors generally, including Kunth, Berg and Amshoff , state that the branch-
lets are, or may be, puberulent or pubescent. The South American and most
Central American specimens I have seen are quite glabrous, including the branch-
lets. Some North American specimens, on the other hand, are notably pubescent
on the branchlets and peduncles. Plants otherwise most like the South American
ones, e.g. Gentle 4062, have the foliage quite glabrous from the first, even when
the branchlets are pubescent.

400 FIELDIANA: BOTANY, VOLUME 24

pedicels 1.5-2.5 cm. long, compressed distally and there up to 1 mm. wide in fruit;
bracteoles linear, deciduous soon after an thesis, up to 1 mm. wide, 4-6 mm. long;
buds 6-6.5 mm. long, pyriform, abruptly expanded above the conic base; calyx
deeply 5-lobed, about half as long as the globe of the petals in mature buds, the
lobes broadly rounded, 1.5-2.5 mm. high, 2.5-3 mm. wide, somewhat separated
in anthesis by irregular longitudinal ruptures in the sinuses between them; disk
4-5 mm. wide, glabrous; style 7-8 mm. long, capitate; stamens very numerous,
probably about 250, about as long as the style and the petals; anthers broadly
oval, about 0.8 mm. long; petals white, 6-7 mm. long; ovary 3-locular, the ovules
biseriate, inwardly directed from an axile stalked placenta; fruit subglobose, when
dry about 1 cm. in diameter, said to be of good flavor.

Amshoff, in the Flora of Panama (Ann. Missouri Bot. Card. 45:
196. 1958), reduced Psidium Oerstedeanum Berg (Linnaea 27: 360.
1856) to the synonymy of P. salutare. The original specimens of
P. Oerstedeanum came from Costa Rica (not Guatemala as mis-
takenly reported by Berg; see Fieldiana, Bot. 29: 524. 1963). Most
Central American plants identified as P. Oerstedeanum seem to have
been merely pubescent specimens of P. salutare. Some plants, how-
ever, with obovate or elliptic and rather prominently veiny leaves,
strongly pubescent stems and peduncles and canescent buds, seem
to represent hybrids between P. salutare and another species of
Psidium, perhaps P. guineense. Probably P. Oerstedeanum Berg orig-
inated in this way. A recently described species from Chiapas,
Psidium chiapasensis Lundell (Trapichito, near Comitan, Matuda
5769, type), is doubtless a synonym of P. Oerstedeanum and simi-
larly of hybrid origin. The supposed hybrids resemble for the most
part small-leaved and gray-pubescent specimens of P. guineense (the
leaves 3-6 cm. long), but the open 5-lobed calyx is that of P. salutare.

Typical Psidium salutare seems to be unknown from Guatemala.
Specimens of what is probably a hybrid between P. salutare and
some other species are known from several localities. All the plants
are characterized by the collectors as depressed or procumbent shrubs
60 cm. high or less. The specimens are variable as to amount of
pubescence and as to width of leaves. The calyx is more nearly
closed than in P. salutare, but is very shortly 5-lobed from the be-
ginning. The following specimens may be cited :

Open gravelly or rocky slopes, rocky summits, and pine forests,
800-1500 meters; Huehuetenango (Steyermark 51656); Chiquimula
(Steyermark 30553); Jutiapa (Standley 76270).

Leaves elliptic-oblong, 2-6 cm. long, 2-3 times as long as wide, rather spar-
ingly pubescent at maturity, gland-dotted beneath, sharply apiculate, the petioles
1-3 mm. long; flowers solitary on slender pedicels 2-3 cm. long, the buds canescent,
7-8 mm. long, slightly open at the tip, the calyx-lobes 5, free about 1 mm. at the

McVAUGH: FLORA OF GUATEMALA 401

tips, the calyx splitting down between the lobes so as to make them apparently
2 mm. high and 3 mm. broad just before anthesis, then finally splitting irregularly
to the base; fruit unknown.

Psidium Sartorianum (Berg) Ndzu. in Engl. & Prantl, Nat.
Pflanzenfam. 3, Abt. 7: 69. 1893. Mitranthes Sartoriana Berg, Lin-
naea 29 : 248. 1858. Calycorectes protractus Griseb. Cat. PI. Cub. 284.
1866. Calyptropsidium Sartorianum (Berg) Krug & Urban ex Urban,
Bot. Jahrb. 19: 571. 1895. Calyptranthes Tonduzii Bonn. Sm. Bot.
Gaz. 23: 245. 1897 (type from near San Jose", Costa Rica, Tonduz
9822). Psidium microphyllum Britton, Sci. Surv. P. R. & Virgin Is.
6: 555. 1930 (type a cultivated plant from Puerto Rico). Mitropsid-
ium Sartorianum (Berg) Burret, Notizbl. Bot. Gart. Berlin 15: 487.
1941. M. oblanceolatum Burret, I.e. (type from Santa Marta, Co-
lombia, H. H. Smith 403). M. Pittieri Burret, I.e. 488 (type from
near Caracas, Venezuela, Pittier 9277). Psidium yucatanense Lun-
dell, Contr. Univ. Mich. Herb. 7: 35. 1942 (type from Belize District,
British Honduras, P. H. Gentle 9). P. Solisii Standl. Field Mus.
Bot. 23: 133. 1944 (type from Costa Rica, F. Solis 509). ? 'Psidium
Molinae Amshoff, Act. Bot. Ne'er!. 5: 277. 1956 (type from Dept.
Morazan, Honduras, Standley 21255). P. Sartorianum var. yucata-
nense McVaugh, Fieldiana, Bot. 29: 527. 1963.

Forests and savannahs, in a wide variety of habitats from shade
to full sun, 1500 meters or less; Chiquimula (Montana Castilla,
southeast of Quezaltepeque, Steyermark 31257). Mexico (Oaxaca to
Jalisco and Sonora; Veracruz, Sartorius, the type; Chiapas; Yuca-
tan); British Honduras; Honduras; El Salvador; Costa Rica; Pan-
ama; West Indies (Cuba); northern Colombia and Venezuela.

A tree up to 12-15 meters high and 30 cm. in diameter, with smooth mottled
bark, the branchlets and inflorescence more or less hirtellous-puberulent with pale
sharp crisped or often antrorse hairs up to 0.4 mm. long; leaves small (5 cm. long
or less), elliptic to ovate or occasionally obovate, usually narrowed about equally
to both ends, the tip from short-acuminate to acute or obtuse, the base acute to
rounded or cuneate, the margins cuneately or somewhat abruptly decurrent on
the compressed and distally dilated petiole 1.5-4 mm. long; midvein convex above,
often pubescent in young leaves; lateral veins 5-8 pairs, in addition to some inter-
mediate ones, evident but rather inconspicuous on both surfaces, ascending; mar-
ginal vein about equaling the laterals, arched between them, 0.5-2 mm. from the
margins; leaves at flowering time thin, drying flat, usually darkening in drying,
paler beneath and with more prominent glands; blades at full maturity quite dif-
ferent in aspect, coriaceous, drying pale green and often nearly concolorous, lus-
trous and usually pale-mottled above, the margins pale and strongly revolute,
the lower surface thickly beset with pale and often convex glands; peduncles soli-
tary, axillary (near the base of the branchlet sometimes in the axils of much-

402 FIELDIANA: BOTANY, VOLUME 24

reduced foliaceous bracts), 7-12 mm. long and 1-flowered, or more often cymosely
3-flowered, the central (terminal) flower sessile, the lateral ones on branches up
to 7-8 mm. long; peduncles slender, somewhat compressed and up to 1 mm. wide
near apex, in 3-flowered dichasia often up to 2 cm. long; buds pyriform, 4-5.5 mm.
long, glabrous, contracted at base into a conic pseudostalk 0.5 mm. long, the distal
portion subglobose or ovoid, closed except a minute apical pore which bears a few
projecting hairs, often prolonged into a narrow apiculate tip up to 0.5 mm. long;
bracteoles linear, soon deciduous; calyx calyptrate, circumscissile at about the
level of the summit of the ovary, the conic or broadly dish-shaped calyptra some-
what persistent by one side; calyx appressed-pubescent within, splitting further
by irregular longitudinal breaks, into (usually 4) recurved lobes up to 1.5 mm.
long; style about 4-5 mm. long, pilose near the base, capitate; stamens about 100,
up to 5 mm. long, the anthers 0.4-0.5 mm. long; petals 1-4, white, concave, sub-
orbicular, 2-4 mm. long, sometimes one or more reduced in size; ovary (2-) 3-locu-
lar, the ovules about 20 in each locule, uniseriate, centrally directed from the
margins of a bilamellate axile placenta; fruit globose or pyriform, 1-1.5 cm. in
diameter, light yellow, resembling a guava in taste and color but with thin flesh
1-2 mm. thick; seeds 1-2 in each locule, with bony testa and C-shaped embryo,
up to 7-8 mm. long.

This tree seems to be very little known in Guatemala, although
it is a common and easily recognized forest tree throughout most of
southern and western Mexico, and is known from various localities
elsewhere in Central America.

In British Honduras and Yucatan and in Pete*n, this species is
represented by a scantily pubescent or essentially glabrous popula-
tion with rather long-acuminate leaves of a lanceolate type. This
is Psidium Sartorianum var. yucatanense McVaugh (P. yucatanense
Lundell). This variety is found also in Cuba, whence it was origi-
nally described as Calycorectes protractus Griseb. Specimens from
the Yucatan Peninsula can often be distinguished at once from other
Mexican and Central American specimens, most of which have short-
pointed or obtuse leaves that tend to be elliptic rather than lanceolate.

Another segregate of P. Sartorianum is P. Molinae Amshoff, of
Honduras. The leaves of this are said to be more obtuse than those
of P. Sartorianum, and the calyx in bud is shortly 4-lobed rather
than calyptrate. In general appearance the plants are like those of
P. Sartorianum.

Much taxonomic importance has been attached in the past to
the manner of opening of the calyx in Psidium. It is now known
that the calyx may open irregularly in some species with normally
calyptrate calyx, and the importance of this particular feature is not
fully understood. For notes on the taxonomy of P. Sartorianum and
its relatives see Fieldiana, Bot. 29: 525-529. 1963.

McVAUGH: FLORA OF GUATEMALA 403

UGNI Turcz.
Reference: Berg, Linnaea 27: 384-393. 1856.

Shrubs with numerous small coriaceous leaves mostly 2 cm. long or less.
Calyx-lobes distinct, 5, rarely 4. Flowers solitary, with persistent foliaceous
bracteoles. Stamens numerous (25 or more), the filaments somewhat flattened
and dilated, the outer longer and broader, the anthers sagittate, introrse, basifixed
and sometimes winged by the margins of the dilated connective. Ovary 3- or
4-locular (according to Berg), the numerous ovules horizontal, affixed to the cen-
tral bilamellate placentae. Seeds reniform or hippocrepiform, with bony testa,
the embryo arcuate.

A genus of about 10 species or fewer, in montane forests from
southern Mexico to Chile. It is distinguished technically from
Myrtus by the characteristic stamens, and from Myrteola by the
stamens, by the somewhat larger leaves and by the usually 5-merous
flowers.

Mexican and Central American plants of this genus seem only
doubtfully distinct from Ugni myricoides (HBK.) Berg, a species of
the higher Andes of northern South America. The South American
plant differs mostly in having white fruit and somewhat smaller
flowers. Berg distinguished four Central American species chiefly
on the basis of anther-shape and by the number of glands at the
tip of the anther. Neither of these characters seems to be consistent.
The glands sometimes vary from 1 to 3 in a single flower.

Probably all Mexican and Central American plants belong to the
following species. Costa Rican specimens, as noted by Berg in de-
scribing U. Oerstedii, often (but not always) have somewhat broader
leaves, and deltoid calyx-lobes 2-3 mm. long and wide. The type of
Myrtus montana presumably came from a plant with relatively small
bracts and leaves, but the specimen is not otherwise very different
from most Central American material. The type of M. Matudai
Lundell has bracteoles 4-7 mm. long and calyx-lobes 3-4 mm. long,
longer than those of the type of M . montana, as noted by Lundell,
but not unusually long when compared with most other specimens.

Ugni montana (Benth.) Berg, Linnaea 27: 392. 1856. Myrtus
montana Benth. PI. Hartw. 61. 1840 (type from Monte Pelado,
Oaxaca, Hartweg 459). Ugni Friedrichsthalii Berg, I.e. 388 (type
from Guatemala, Friedrichsthal, the exact locality not specified).
U. Friedrichsthalii a longipes Berg, I.e. 389. U. Friedrichsthalii fi
brevipes Berg, I.e. U. Oerstedii Berg, I.e. (type from Costa Rica,
Oersted). U. Warscewiczii Berg, I.e. 390 (type locality, "in Ve-
ragua," Warscewicz) . Myrtus Oerstedii (Berg) Hemsl. Biol. Centr.

404

FIELDIANA: BOTANY, VOLUME 24

Am. Bot. 1: 407. 1880. Eugenia Friedrichsthalii (Berg) Hemsl. I.e.
410. E. Warscewiczii (Berg) Hemsl. I.e. 411. Myrtus Friedrichs-
thalii (Berg) Bonn. Sm. Enum. PI. Guat. 3: 25. 1893. M. Matudai
Lundell, Phytologia 1: 246. 1937 (type from Chiapas). Sco (Volcan
de Zunil) ; mirto.

FIG. 55. Ugni montana (from Carlson 2390). A, Bud just before opening;
X 2.5. B, Slightly immature fruit; X 2.5. C, Dorsal, lateral, and ventral views
of anther; X 12.5.

Open, moist or dry, high mountain hillsides and meadows, or
often in dense Cupressus or Abies forest, 1500-3700 meters, most
abundant at higher elevations; El Progreso; Zacapa; Chimaltenango;
Solola; Quiche" ; Huehuetenango; Totonicapan; Quezaltenango; San
Marcos. Mountains of southern Mexico (Oaxaca, Chiapas) ; El Sal-
vador; Honduras; Costa Rica; Panama; perhaps the same species
in the Andes from Colombia and Venezuela to southern Ecuador.

A shrub or small tree 1-3 meters high, or often much lower, usually very
densely branched, the branches densely leafy; branchlets, inflorescence and young
foliage strigose-canescent with sordid hairs, the foliage and flowers soon glabrate;
leaves elliptic or occasionally ovate, rigidly coriaceous, mostly 5-10 (-15) mm.
wide, (7-) 10-18 (-28) mm. long, about twice as long as wide, acute at both ends
or in ovate leaves more rounded at base, the margins decurrent on the petiole
2 mm. long; midvein sulcate above, broad and prominent beneath, the other veins
scarcely apparent; blades dark green above, with revolute margins, sometimes
impressed-punctate; lower surface paler and dull, with a few small glandular dots;
flowers spreading or in age recurved, the pedicels 1-2 cm. long, 0.5 mm. thick;
bracteoles in texture like the leaves, linear or elliptic, spreading or at last recurved,
up to 1 mm. wide, 3-6 (-8) mm. long; hypanthium sessile in the bracteoles, cup-
shaped or hemispheric, 2-3 mm. high; calyx-lobes 5, foliaceous, acutely triangular,
loosely recurved or in fruit stellately spreading, 2-4 mm. long, 1.5-2 mm. wide
at base; disk glabrous, in fruit up to 3-3.5 mm. wide; style glabrous, 6-7 mm.
long, capitate; stamens about 50, probably pale pink to white, the outer ones up
to 4 mm. long; anthers 1-1.3 mm. long, sagittate, 0.5-0.6 mm. wide at base, with
1 (occasionally 2-3) apical glands; petals white, 5-7 mm. long; ovary imperfectly

McVAUGH: FLORA OF GUATEMALA 405

3-locular, the ovules about 10 in each locule, uniseriate on erect bilamellate axile
placentae; fruit globose, 6-8 mm. in diameter, at first red, turning black or pur-
plish black; seeds up to 20, broadly reniform, 1.5-2 mm. long, straw color, lustrous.

A very common and characteristic shrub at high elevations, espe-
cially in deep forest of Abies and Cupressus (Standley, mss.).

SPECIES OF UNCERTAIN GENERIC POSITION

Myrtus bullata Salisb. Prodr. 354. 1796. Calyptranthes? bullata
(Salisb.) DC. Prodr. 3: 258. 1828.

The description of this species, and the specific epithet, both
suggest Psidium Sartorianum. The type was from Honduras.

Publication 974

UNIVERSITY OF ILLINOIS-URBANA