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FLORA OF GUATEMALA

PAUL C. STANDLEY

AND

JULIAN A. STEYERMARK

FIELDIANA: BOTANY

VOLUME 24, PART III

Published by

CHICAGO NATURAL HISTORY MUSEUM
APRIL 25, 1952

FLORA OF GUATEMALA

PART III

FLORA OF GUATEMALA

PAUL C. STANDLEY

Curator Emeritus of the Herbarium

AND

JULIAN A. STEYERMARK

Curator of the Herbarium

FIELDIANA: BOTANY
VOLUME 24, PART III

Published by

CHICAGO NATURAL HISTORY MUSEUM
APRIL 25, 1952

MAY 5 - 1C52

UNIVERSITY OF ILLMOiS

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

CONTENTS

Families Included in Part III

PAGE

Commelinaceae ................. 1

Pontederiaceae .................. 42

Juncaceae ...................... 52

Liliaceae ....................... 59

Smilacaceae ..................... 92

Haemodoraceae ................. 100

Amaryllidaceae .................. 103

Dioscoreaceae ................... 145

Iridaceae ....................... 159

Musaceae ...................... 178

Zingiberaceae ................... 191

Cannaceae.. . 203

Marantaceae

Burmanniaceae

Casuarinaceae

Piperaceae

Chloranthaceae

Lacistemaceae

Salicaceae

Myricaceae

Juglandaceae

Betulaceae

Fagaceae

Urticaceae..

PAGE

207
221
227
228
337
340
342
348
352
359
369
396

COMMELINACEAE. Spiderwort Family

References: C. B. Clarke, Commelinaceae, in DC. Monogr.
Phan. 3: 113-324. 1881. Robert E. Woodson, Jr., Commentary on
the North American genera of Commelinaceae, Ann. Mo. Bot.
Card. 29: 141-154. 1942.

Annual or perennial, mostly succulent herbs, erect to procumbent or repent,
often rooting at the nodes; leaves alternate, entire, the petiole dilated into a basal
sheath; flowers perfect, mostly small, umbellate, cymose, racemose or capitate,
the bracts small or large, often spathaceous; perianth usually differentiated into
a distinct calyx and corolla; sepals 3, free or connate, imbricate, usually herbaceous,
the 2 inner ones generally subfalcate or asymmetric; petals usually 3, free or con-
nate, equal or unequal, marcescent, blue, purple, pink, or white; stamens normally
6 but sometimes fewer, all or partly fertile, usually free, the anthers 2-celled;
ovary superior, sessile, 3-2-celled, ovules orthotropous, few; stigma entire or
obscurely lobate; fruit capsular, or crustaceous and indehiscent.

An essentially tropical family, with about 25 genera, the plants
widely distributed in both hemispheres. Other genera represented
in southern Central America are Floscopa and Cochliostema. The
treatment used here is that proposed by Woodson in the paper
listed above. His treatment is radical in some respects, although
not startlingly so. Most of the groups are left undisturbed in their
traditional sense, while discordant elements have been removed
from some of the larger genera and united with minor groups to
form more easily recognized and consistent units.

Ultimate branches of the inflorescence composed of individual scorpioid cymes,
these appearing 1-sided superficially, solitary or variously clustered, very
rarely reduced to a single terminal flower; corolla regular or irregular.

Commelineae.

Cymes solitary, enclosed by a conspicuous compressed spathe-like bract.
Fruit dehiscent, capsular; sterile stamens, when present, with cordate anthers.

Commelina.
Fruit indehiscent, pergamentaceous; sterile stamens with hastate-triangular

anthers Phaeosphaerion.

Cymes variously clustered or compounded, rarely solitary but never enclosed

by a compressed spathaceous bract.
Anthers large, with an inconspicuous connective, dehiscent by apical pores;

seeds with a fleshy aril Dichorisandra.

Anthers small but with a conspicuous sterile connective, dehiscent longitudi-
nally; seeds dry, not arillate.

Flowers regular or essentially so Aneilema.

Flowers very strongly irregular Tinantia.

2 FIELDIANA: BOTANY, VOLUME 24

Ultimate branches of the inflorescence composed of paired sessile scorpioid cymes,
appearing as a 2-sided unit superficially, rarely reduced to a single flower;

corolla regular Tradescantieae.

Corolla gamopetalous, the petals united at the base.

Plants acaulescent or nearly so; flowers solitary and sessile in the axils of the

congested upper leaves; corolla tube 4-6.5 cm. long Weldenia.

Plants with elongate leafy stems; flowers in leafy-bracted cymes; corolla

tube 1-1.5 cm. long Zebrina.

Corolla polypetalous, the petals free to the base.

Paired cymes distinctly pedunculate, never sessile or subtended by leafy
bracts; stamens 6, usually in 2 very dissimilar series, the outer ones
occasionally sterile, rarely all the stamens fertile and essentially similar;

sepals foliaceous or petaloid Tripogandra.

Paired cymes sessile and subtended by conspicuous bracts similar to the
leaves (coriaceous spathes in Rhoeo), rarely appearing pedunculate and
the bracts greatly reduced but the stamens then usually 1-3, and the
sepals paleaceous.

Paired cymes sessile and subtended by more or less conspicuous, leafy
bracts, rarely appearing pedunculate and the bracts greatly reduced,
but the stamens usually 1-3, rarely 6, all fertile, and the sepals pale-
aceous Callisia.

Paired cymes sessile and subtended by conspicuous bracts essentially similar
to the leaves (coriaceous spathes in Rhoeo); stamens 6, all fertile and
essentially similar; sepals foliaceous or petaloid.

Cymes terminal on the main stem, occasionally also lateral in the upper
leaf axils; lateral cymes very rarely reduced to a single flower.

Tradescantia.

Cymes on slender peduncles lateral to the main stem.
Flowering peduncles elongate, usually branched; bracts foliaceous;

sepals becoming fleshy in fruit Campelia.

Flowering peduncles very short, simple; bracts appearing as coriaceous
spathes; sepals drying in fruit Rhoeo.

ANEILEMA R. Brown

Herbs with branched stems, the leaves narrow or rather broad; peduncles
axillary or terminal, many-flowered, paniculate, thyrsoid, or corymbose, or few-
flowered, rarely 1-flowered; lowest bracts resembling reduced leaves, never spa-
thaceous or complicate, the upper ones small, often amplexicaul, generally persist-
ent; sepals 3, free, the outer one cucullate, oblong-elliptic, the 2 inner ones obovate
or oblong, falcate, green, scarious, or petaloid, persistent; petals 3, free, all alike
or slightly unequal, blue or white, deciduous or marcescent; stamens 3-6, only
2 or 3 of them perfect, the filaments slender, barbate or glabrous; ovary sessile,
glabrous or pubescent, 2-3-celled, the dorsal cell smaller or wanting; ovules 1-20
in each cell; capsule dry, 2-3-celled, loculicidally 2-3-celled; seeds 1-20 in each
cell; seeds rugose or foveolate, often puberulent.

Species about sixty, in the tropics of both hemispheres, most of
them in the Old World. All the known Central American species
are listed below.

Plants erect; leaves glabrous or nearly so; bracts at the base of the ultimate
peduncles opposite, large and leaf-like A. aguensis.

Plants prostrate or procumbent; leaves densely long-pilose beneath; bracts at the
base of the ultimate peduncles small and inconspicuous A. geniculata.

FIG. 1. Aneilema geniculata. A. Habit; X / B. Stamen; X 11. C. Por-
tion of inflorescence with flower in anthesis; X 4.

4 FIELD IANA: BOTANY, VOLUME 24

Aneilema aguensis Standl. & Steyerm., comb. nov. Trades-
cantia aguensis Standl. & Steyerm. Field Mus. Bot. 23: 36. 1944.

Moist shaded banks and brushy slopes, 1,800-2,900 meters;
endemic; Huehuetenango; Sacatepequez (type collected on slopes
of Volcan de Agua above Santa Maria de Jesus, Standley 59358).

Plants slender, erect, about 50 cm. high, apparently with fibrous roots, simple
or branched, the stems densely or lightly villous; leaves thin, lanceolate or oblong-
lanceolate, 2-6.5 cm. long, 1-1.5 cm. wide, acuminate, subequally cuneate at the
base, sparsely pilose above, glabrous beneath or sparsely pilose along the costa,
ciliate; sheaths thin, 4-5 mm. long, 2-5 mm. broad, villous, the orifice and margins
ciliate; bracts at the base of the ultimate umbels 2 and opposite, lanceolate, 2-4 cm.
long; umbels on long slender glabrous peduncles, 3-4-flowered, the pedicels spread-
ing to deflexed, filiform, glabrous or glabrate, 4-11 mm. long; sepals thin, greenish,
scarious-marginate, 2-3 mm. long, lance-oblong, acute, glabrous; petals pale blue,
2.5 mm. long; filaments barbate at the middle; capsule 3-3.5 mm. long; seeds 5,
dark grayish brown, irregularly sulcate, about 1.2 mm. long and 1 mm. broad.

Aneilema geniculata (Jacq.) Woodson, Ann. Mo. Bot. Card.
29: 147. 1942. Tradescantia geniculata Jacq. Sel. Stirp. Amer. 94.
pi 64. 1763.

Moist or wet thickets or forest, 300 meters or lower; Pete"n;
Alta Verapaz; Izabal; Huehuetenango. Central and southern
Mexico; British Honduras to Panama; West Indies; South America.

Plants perennial, apparently with fibrous roots, the stems simple or branched,
prostrate or decumbent, rooting at the lower nodes, usually puberulent in a line
along one side, 20-60 cm. long; leaves thin when dried, oblong-lanceolate to ovate,
3-7 cm. long, 1-3 cm. wide, somewhat paler beneath, acute or acuminate, rounded
at the base or contracted at the somewhat unequal base, sparsely pilose or glabrate
above, usually long-pilose beneath, sometimes ciliate; sheaths thin, scarious, 5-7
mm. long, 2-4 mm. broad, long-pilose, pilose-ciliate; peduncles terminal or in the
upper leaf axils, filiform, 2.5-5 cm. long, glabrous or sparsely pilose; ultimate
umbels few-flowered, the pedicels filiform, 4-12 mm. long, glabrous to glandular-
pilose; bracts at the base of the peduncles greatly reduced, 1-2 mm. long; sepals
green, often tinged with purple, acute or acuminate, 2-3 mm. long, glabrous except
for the sparsely barbate tip; petals white, 3-5 mm. long; filaments barbate;
capsule 2-3 mm. long; seeds gray, finely and minutely reticulate, 1-1.5 mm. in
diameter. (Fig. 1.)

CALLISIA L.

Plants usually small, perennial, procumbent or prostrate, often with erect
branches; leaves generally pale green, ovate to elliptic-lanceolate, acute, the upper
ones subsessile, vaginate; flowers small, white, aggregate within the sheaths or
axillary, sometimes in axillary umbels and forming terminal panicles; bracts
uniform with the leaves, or the uppermost reduced almost to sheaths, the bractlets
small or very narrow; sepals 2-3, subequal or one of them smaller, free, elliptic
to oblong-linear, green or hyaline, erect and persistent in fruit; petals 2-3, sub-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 5

equal or one of them smaller, free, elliptic or lanceolate, marcescent; stamens
1-3, fertile, subequal, free, the filaments filiform, naked; anther cells rounded and
separated by the connective, or oblong and contiguous; ovary sessile, oblong,
2-3-celled, compressed or subtrigonous; style long, the stigma penicillate or short
and 3-lobate; ovules 2 in each cell; capsule small, thin, ellipsoid, 2-3-celled, loculi-
cidally 2-3-valvate; seeds 2 in each cell or rarely by abortion 1, superposed,
cylindric-trigonous or subtrapezoid, the testa fuscous or stramineous, smooth,
rugulose, or radiately striate.

Four species, in tropical America. The fourth one is native in
central Mexico.

Flowers scarcely exserted from the sheaths, in dense sessile clusters; style filiform

and elongate C. repens.

Flowers long-exserted from the sheaths, paniculate or umbellate, on obvious

pedicels; style short.
Leaves usually glabrous beneath, thin; leaf sheaths 4-5 mm. long; inflorescences

axillary and terminal C. monandra.

Leaves usually densely tomentulose beneath, firm-membranaceous; leaf sheaths
7-12 mm. long; inflorescences terminal, on a long peduncle. .C. multiflora.

Callisia monandra (Swartz) Schult. in Roem. & Schult. Syst.
Veg. 7: 1179. 1830. Tradescanlia monandra Swartz, Prodr. Veg.
Ind. Occ. 57. 1788. C. umbellulata Lam. 111. 1: 130. pi 35, /. 2.
1791. Matajalln (Jutiapa) ; Lochoch (fide Aguilar).

Moist or wet thickets or forest, sometimes in oak forest or in
roadside hedges, 1,700 meters or lower; Chiquimula; Jalapa; Jutiapa;
Retalhuleu. Southern Mexico; British Honduras to Costa Rica;
West Indies; South America.

Stems slender, pale green, simple or branched, 30-50 cm. long, rooting at the
nodes and often forming dense mats, glabrous below, glandular-pilose above; leaves
ovate or ovate-lanceolate, 2-5 cm. long, 1-1.5 cm. wide, acute or acuminate, round-
ed to subcordate or obliquely cuneate at the base, ciliolate, glabrous or sparsely
pilose on both surfaces, sessile or the lower leaves subpetiolate; sheaths small,
4-5 mm. long, villous-ciliate; peduncles axillary, bearing laterally 3-10-flowered
umbels, these sometimes forming lax terminal panicles, glandular-pilose; pedicels
filiform, 15 mm. long or shorter, glandular-pilose, the bractlets lanceolate, pilose;
flowers minute, 2-3-parted, white; sepals 1-2 mm. long, elliptic-oblong, sparsely
pilose, barbellate at the apex, green, scarious-marginate; petals smaller than the
sepals, lanceolate, hyaline; stamens 1-2, the filaments glabrous, the anther cells
rounded; capsule 1-1.3 mm. long, apiculate, 2-3-celled.

Callisia multiflora (Mart. & Gal.) Standl. Journ. Wash. Acad.
Sci. 15: 457. 1925. Commelina multiflora Mart. & Gal. Bull. Acad.
Brux. 9, pt. 2: 374. 1842. Tradescantia Martensiana Kunth, Enum.
PI. 4: 697. 1843. Callisia Martensiana C. B. Clarke in DC. Monogr.
Phan. 3: 312. 1881. Tzimaac (Coban, Quecchi).

FIG 2. Callisia repens. A. Habit; X \i. B. Characteristic crowded leaves
of sterile branch; X Y 2 . C. Flower; X 6. D. Capsule; X 22. E. Capsule
dehiscing; X 22.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 7

Moist or wet thickets or forest, often in wet places along streams,
sometimes a weed in cultivated ground, 350-2,500 meters; Alta
Verapaz; Baja Verapaz; Guatemala; Sacatepe"quez; Chimaltenango;
Huehuetenango. Southern Mexico; Honduras.

Stems rather stout, 3-4 mm. in diameter, pale green, procumbent, sometimes
with erect branches, 30-80 cm. long, somewhat branched above or simple, glabrous
or pilose; leaves lance-oblong or elliptic-lanceolate, 3-9 cm. long, 1-2.5 cm. wide,
acute or acuminate, sometimes abruptly so, rounded or subcordate at the sessile
base, pale green, usually densely tomentulose beneath, glabrate or tomentulose on
the upper surface; sheath usually very villous, sometimes glabrate, villous-ciliate,
7-12 mm. long, 3-6 mm. broad; peduncles elongate and branched, forming dichoto-
mous panicles, large and many-flowered, the branches glandular-pilose or glabrate,
the pedicels glandular-pilose, the bracts ovate-lanceolate; flowers 3-parted, in
clusters of 3-6; sepals 2-3 mm. long, green and scarious-marginate, glandular-
pubescent or glabrate; petals white, 3-4 mm. long, elliptic; stamens 3, the filaments
glabrous; capsule ellipsoid, 2 mm. long; seeds 6.

Called "carricillo" in Veracruz.

Callisia repens L. Sp. PL ed. 2. 62. 1762. Lochoch (fide Aguilar).

Moist or wet thickets or forest, or on shaded banks or rocks,
sometimes on wet stream banks, frequently a weed in cultivated
ground in the lowlands, 2,300 meters or lower; El Progreso; Izabal;
Zacapa; Jalapa; Jutiapa; Santa Rosa; Guatemala; Sacatepe"quez;
Chimaltenango; Huehuetenango; Quezaltenango; San Marcos.
Southern Mexico; British Honduras to Costa Rica; West Indies;
South America.

Stems slender, prostrate, rooting at the nodes, often forming large mats,
simple or branched, short or often much elongate, glabrous; leaves ovate, pale
green, somewhat succulent when fresh, membranaceous when dried, mostly 1-4 cm.
(rarely 7 cm.) long, 1-1.5 or rarely 2 cm. wide, acute or acuminate, rounded or
subcordate at the sessile base, or the lower leaves sometimes subcuneate and
petiolate, glabrous, ciliolate; sheaths usually glabrous, villous-ciliate; uppermost
leaves of sterile branches much crowded and imbricate, successively smaller;
flowering branches erect or ascending, the flowers 3-parted, in dense sessile clusters
of 3-9 on an elongate terminal axis, the flowers scarcely exserted beyond the
sheaths; sepals 2-3 mm. long, linear-oblong, green, pubescent, scarious-marginate;
petals 3, shorter than the sepals, white, oblong, hyaline; stamens 3, the filaments
glabrous, the anther cells rounded; capsule 1.5 mm. long, 6-seeded. (Fig. 2.)

This is an abundant weedy plant in wet places of the lowlands
and lower mountains in many parts of Central America, frequently
invading cultivated ground, especially cafetales. The plants often
form large mats, and cover large areas of ground in moist places.
As in other species, they usually are of a very pale shade of green,
thus being conspicuous in contrast to neighboring vegetation.

8 FIELDIANA: BOTANY, VOLUME 24

CAMPELIA L. Richard

Stout, usually erect, perennial herbs, glabrous or pubescent, the stems simple
or sparsely branched; leaves mostly lanceolate, succulent when fresh, sheathing
at the base; peduncles axillary, elongate, almost leafless, 1-2-dichotomous above
or simple, the branches bearing at the apex 2 subopposite, lanceolate, sometimes
complicate or cymbiform bracts; flowers fasciculate, short-pedicellate, subtended
by 2 bracts, usually little or not at all exceeding the bracts, the bractlets numerous,
small, ovate; flowers almost regular, the sepals 3, almost free, ovate-oblong,
herbaceous, somewhat accrescent in age, the outer one somewhat cucullate, all
the sepals succulent, persistent in fruit; petals 3, free, white; stamens 6, subequal,
the filaments barbate; anther cells oblong, separated by an oblong-triangular
connective; ovary sessile, 3-celled; capsule 3-celled or by abortion 2-celled, ellipsoid,
trigonous, loculicidally 3-valvate, the cells 1-2-seeded; seeds osseous, compressed-
ellipsoid, punctate.

About three species, in tropical America.

Sheaths of the leaves densely hirsute with spreading golden-brown hairs . C. hirsuta.
Sheaths of the leaves glabrous or sericeous with appressed, pale or dull hairs.
Stems, peduncles, and lower surface of the leaves glabrous; inflorescence sub-
tended by 2 reduced bracts, the larger one 1.5-6.5 cm. long and 1-2 cm.

wide; bractlets 1-3 mm. long C. Zanonia.

Stems, peduncles, and lower leaf surface more or less densely sericeous; inflores-
cence subtended by 2 reduced bracts and a large leaf 8-11 cm. long and
3-4.5 cm. wide; bractlets 7-10 mm. long C. Standleyi.

Campelia hirsuta Standl. Field Mus. Bot. 8: 135. 1930.

Dense wet mixed forest, 1,600 meters or lower; Alta Verapaz
(near Tactic); Izabal (Cerro San Gil). Atlantic lowlands of Hon-
duras, the type collected in the Lancetilla Valley, near Tela.

A stout erect herb a meter high or less, the stems simple, sparsely hirsute or
glabrate, sometimes purplish red; leaves oblong-oblanceolate or lanceolate, 14-20
cm. long, 4-5.5 cm. wide, abruptly long-acuminate, gradually attenuate to the
base into a petioliform portion; sheaths somewhat inflated, about 1 cm. long;
peduncles mostly longer than the leaves, sparsely or densely fulvous-hirsute;
flowers few, white; bracts unequal, the larger one oval-ovate, 4.5-6 cm. long, 2-3.5
cm. wide, abruptly acute, truncate or subcordate at the base, densely hirsute
on both surfaces, the smaller bract 1.5-3 cm. long, acute, complicate, densely
hirsute at the apex; sepals oblong, 4-5 mm. long, sparsely hirsute-ciliate, more
densely hirsute about the apex; petals white. (Fig. 3.)

Campelia Standleyi Steyermark in Standl. & Steyerm. Field
Mus. Bot. 23: 32. 1944.

Moist or wet forest, 200-1,500 meters; endemic; Alta Verapaz;
San Marcos (type from Volcan de Tajumulco, above Finca El
Porvenir, Steyermark 37187).

Plants stout, decumbent or erect, the stems purple or dull green mottled with
purple, 4-6 mm. thick, 60-100 cm. tall or more, sparsely or densely sericeous,

FIG. 3. Campelia hirsuta. Habit of upper portion of plant; X Y^

10 FIELD IANA: BOTANY, VOLUME 24

especially in the upper half; leaves deep green above and lustrous, pale or dull
green beneath, the main cauline ones broadly oblanceolate or oblong-elliptic, 9-25
cm. long, 2.5-6 cm. wide, acuminate or caudate-acuminate, gradually attenuate
below into a petioliform portion 1-2 cm. long, glabrous above, densely and shortly
sericeous beneath with dull hairs, the young leaves often with 2 silver stripes on
the upper surface; sheaths lax, membranaceous, 1.3-2.5 cm. long, 3-12 mm.
broad, sparsely or densely sericeous, the margins and orifice finely ciliate; peduncles
terminal and axillary, short or elongate, erect-ascending, 1-15 cm. long, densely
sericeous, naked below, leafy or subtended by well-developed leaves, especially
above; bracts at the base of the inflorescence 2, reduced and somewhat spathe-
like, ovate, caudate-acuminate, rounded or subcordate at the broad base, densely
short-sericeous beneath, 1-2 cm. long; pedicels short, the bractlets conspicuous,
brown-scarious, 7-10 mm. long; sepals 4.5-5 mm. long, firm-membranaceous;
petals rose-purple or white; seeds pale brownish, 2.3-3 mm. long, 1.5-2 mm.
broad, shallowly rugose.

Campelia Zanonia (L.) HBK. Nov. Gen. & Sp. 1: 264. 1816.
Commelina Zanonia L. Sp. PI. 41. 1753. Campelia glabrata Kunth,
Enum. PL 4: 109. 1843. Campelia mexicana Kunth, loc. cit. C.
Zanonia var. glabrata C. B. Clarke in DC. Monogr. Phan. 3: 315.
1881. Coyontura; Lochoc amargo; Oreja de burro.

Moist or wet, usually mixed forest, or in thickets, 2,400 meters
or lower; Pete"n; Alta Verapaz; Izabal; Zacapa; Chiquimula; Jalapa;
Santa Rosa; Guatemala; Sacatepe"quez; Quiche"; Huehuetenango;
Suchitepequez; Quezaltenango; San Marcos. Southern Mexico;
British Honduras to Panama; West Indies; South America.

A stout herb, the stems erect or decumbent, a meter high or less, usually
simple, glabrous, 8-10 mm. thick, dull green or mottled or streaked with dull
purple; leaves mostly crowded near the top of the stem, widely ascending, lanceo-
late to oblanceolate, 10-35 cm. long, 3.5-8 cm. wide, succulent, firm-membran-
aceous to subcoriaceous when dried, dark green above, pale dull green or silvery
purplish beneath, the young leaves sometimes striped with cream-color and green,
glabrous on both surfaces or appressed-pubescent beneath near the margins, long-
acuminate, gradually narrowed below into a petiolar portion 5-20 mm. long, or
subsessile; sheaths 1.5-2.5 cm. long, 1-2 cm. broad; pedicels 6 mm. long or shorter,
the bractlets 1-3 mm. long; sepals 3-5 mm. long, glabrous, firm-membranaceous;
petals white, lilac, or pinkish, 9-10 mm. long, obovate; capsule 3-5 mm. long,
black or dark purple.

COMMELINA L.

Perennial herbs, simple or branched, glabrous or pubescent; leaves mostly
broad, sheathing at the base, more or less succulent; peduncles subtended by
spathiform bracts, solitary or aggregate, scattered or crowded at the ends of the
branches, the peduncles bifid above, the 2 branches racemiform, the lower branches
1-3-flowered, with usually sterile flowers, the upper branches flowering later,
with 2-12 more or less secund flowers; sepals 3, the outer one cucullate, oblong-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 11

elliptic, the 2 inner ones obovate or oblong and falcate, green or petaloid, persistent,
sometimes accrescent in age; petals 3, free, the outer one cucullate, ovate, very
shortly unguiculate, usually small, sometimes absent, the 2 inner petals usually
long-unguiculate, ovate or cordate, generally blue, marcescent; perfect stamens 3,
2 or 3 others usually present but with sterile cruciate-quadrifid anthers, the fila-
ments long, slender, glabrous; anthers of the fertile stamens oblong; ovary sessile,
glabrous, 2-3-celled, the dorsal cell 1-ovulate, sometimes abortive, the 2 ventral
cells l-2-o vulate; capsule dry, chartaceous, loculicidally 2-3-valvate; seeds small,
pyramidal or ellipsoid, more or less compressed, the testa smooth or variously
roughened.

About ninety species, widely distributed in both hemispheres,
most numerous in tropical regions. Probably all the Central
American ones are included in this treatment.

Sheaths conspicuously ferruginous-ciliate at the orifice and along the margins.

C. robusta f. vestita.

Sheaths white-ciliate, or at least not as above.
Margins of the spathe united below.

Spathes and leaves scabrous-ciliate on the margins; sheaths not auriculate at
the summit, the lower and middle ones 3.5-4 cm. long; leaves scabrous

on both surfaces C. Standleyi.

Spathes and leaves not scabrous-ciliate on the margins; sheaths prominently
auriculate at the summit, the lower and middle ones 1-3 cm. long;
leaves not scabrous.
Larger leaves of the main branches lanceolate to lance-ovate, 1-4 cm. wide;

mature spathes 2-5 cm. long.
Petals deep or pale blue.

Spathes glabrous or minutely hirtellous C. erecia.

Spathes white-villous with long hairs at the base.C. erecia f. intercursa.

Petals white C. erecia f. Candida.

Larger leaves of the main branches narrowly linear to linear-lanceolate,

3-20 mm. wide; mature spathes 1-3 cm. long.
Sheaths usually glabrous or slightly pubescent; leaves glabrous to

sparsely pubescent.
Spathes densely short-villous.

Petals blue C. erecia var. angustifolia.

Petals white C. erecia var. angustifolia f. albina.

Spathe white-villous with long hairs.

Petals blue C. erecia var. angustifolia f. crispa.

Petals white C. erecia var. angustifolia f . cana.

Sheaths and both surfaces of the leaves densely pubescent.

C. erecia var. angustifolia f. villosa.
Margins of the spathe free.

Spathes not transversely striate, not with dark purple edges; plants glabrous

or glabrate; petals usually small, less than 5 mm. long C. diffusa.

Spathes conspicuously transverse-striate between the nerves, usually with
dark purple edges; leaves, stems, peduncles, and spathes scabrous or
finely pubescent; petals about 15 mm. long.

Plants acaulescent when young, in age branched only from the base, the
stem-like peduncles very stout and mostly 6-20 cm. long; spathes

2-3 cm. wide (when folded) and 3-4 cm. long C. alpestris.

Plants with well-developed and much elongate stems from the beginning
of anthesis, the stem bearing several or numerous spathes on slender,

12 FIELDIANA: BOTANY, VOLUME 24

relatively short peduncles; folded spathes mostly 1-1.5 cm. wide and
2-3 cm. long.
Leaves oblong-lanceolate to ovate-lanceolate, 2.5-4 cm. wide.

C. coelestis.
Leaves narrowly lanceolate to linear, 5-12 mm. wide.

C. coelestis var. Bourgeaui.

Commelina alpestris Standl. & Steyerm. Field Mus. Bot. 23:
213. 1947.

Moist meadows in the high mountains, usually in alpine regions,
often on limestone, 2,500-3,700 meters; endemic; Chimaltenango
(Cerro de Tecpam); Huehuetenango (Sierra de los Cuchumatanes;
type from vicinity of Che"mal, Steyermark 50265).

Plants perennial from a dense cluster of slender but fleshy roots, acaulescent
at the beginning of anthesis, in age branched from the base or shortly above the
base and becoming as much as 30 cm. high, or perhaps even taller, the height
consisting mostly of the length of the peduncles; leaves all basal or a few present
on the lower part of the stem, lanceolate, 11 cm. long and 2.5 cm. wide or smaller,
attenuate-acuminate, dilated and sheathing at the base, the sheaths membranous,
pale, about 2.5 cm. long and 1 cm. broad, eciliate, the blades glabrous, eciliate,
green above, slightly paler beneath, the margins slightly cartilaginous-thickened;
peduncles mostly appearing simple and scapiform, but actually arising from a short
stem bearing 1 or more large leaves near the base, the early and perhaps sometimes
all the peduncles very short and concealed by the leaf sheaths, the fruiting
peduncles much elongate, sometimes 22 cm. long but mostly shorter, very stout,
as much as 5 mm. thick, smooth and glabrous; spathe at anthesis green, about
3-3.5 cm. long and when folded 2-2.5 cm. wide, obtuse or acute, in age as much
as 3 cm. broad (when folded) and 4 cm. long, thinly hirsutulous, very strongly
trans verse- veined between the nerves; flowers numerous, borne on short stout
pedicels; sepals glabrous, pale green, 5 mm. long, subacute; petals large, about
15 mm. long, deep blue.

This species is closely related to C. coelestis, of which it might be
considered an extreme form, but apparently it is fully distinct and
better marked than many other members of the genus. In general
appearance it is quite distinct from C. coelestis in being acaulescent
or bearing greatly elongate, scapiform peduncles, which are much
stouter than in that species and at first glance appear to be stems
rather than peduncles. The spathes also are much larger than in
Guatemalan plants of C. coelestis, being both relatively and absolutely
broader. The plants grow and flower only when there is abundant
moisture, drying when the rains cease but remaining in place for a
long time afterward, when the dry stems are conspicuous in the
alpine meadows.

Commelina coelestis Willd. Enum. Hort. Berol. 1: 69. 1809.
C. pallida Willd. Hort. Berol. 2: 87. pi. 87. 1816. C. acuminata HBK.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 13

Nov. Gen. & Sp. 1: 258. 1815. China cucharita (Guatemala);
Lochoch (fide Aguilar).

Moist fields or meadows or open forest, often in pine-oak forest
or with Juniperus, 900-3,500 meters; Jalapa; Jutiapa; Guatemala;
Sacatepe"quez; Chimaltenango; Quiche"; Huehuetenango; Quezal-
tenango. Mexico.

Plants perennial from a dense cluster of elongate fleshy-thickened roots, erect,
stout or slender, usually 40-70 cm. high, generally branched above, the stems
scabrous, especially above, the peduncles also scabrous; leaves oblong-lanceolate
to ovate-lanceolate, 8-20 cm. long, 2.5-4 cm. wide, acute or acuminate, some-
what rounded or narrowed at the base, scaberulous above, villosulous or glabrate
beneath; sheaths thin, scarious, often purplish or striped with purple, rather lax,
1.5-2.5 cm. long, 5-7 mm. broad, white-ciliate, usually glabrous; peduncles slender,
often numerous, erect, 1-6.5 cm. long; spathes usually mottled or veined with
purple in the upper half, shortly and abruptly acute or acuminate, subcordate
or rounded at the base, short-villous throughout or villous at the base, mostly
2-3 cm. long and when folded 1-1.5 cm. wide, with conspicuous transverse veins
between the nerves; upper raceme of the inflorescence 4-10-flowered, the lower
raceme 1-2-flowered; petals deep blue, about 15 mm. long; seeds blackish brown,
foveolate.

This plant, like the preceding species, is found in fresh state only
when there is abundant moisture, but the dry stems and foliage
often persist long into the dry season.

Commelina coelestis var. Bourgeaui C. B. Clarke in DC.
Monogr. Phan. 3: 153. 1881. Coy ontur a; Lochoch (fide Aguilar).

Moist meadows or pine-oak forest, 1,000-2,100 meters; Zacapa;
Chimaltenango; Quiche"; Huehuetenango. Southern Mexico.

Similar to the species except for the narrowly lanceolate to linear leaves,
these only 5-12 mm. wide.

This form is not sharply distinguished from the typical one.

Commelina diffusa Burm. f. Fl. Ind. 18. pi. 7, f. 2. 1768.
C. nudiflora sensu Burm. f. op. cit. 17, and of many other later and
recent authors, not L. C. longicaulis Jacq. Coll. Bot. 3: 234. 1789.
Hierba de polio; Lochoch, Lochoch de sapo (fide Aguilar).

Moist or wet, open places, thickets, or forest, often a weed in
waste or cultivated ground, 1,600 meters or lower, most common in
the lowlands; Pete"n; Alta Verapaz; El Progreso; Izabal; Zacapa;
Chiquimula; Jalapa; Santa Rosa; Escuintla; Guatemala; Sacatepe"-
quez; Solola; Suchitepequez; Retalhuleu; Quezaltenango ; San
Marcos; Huehuetenango. Florida; Mexico; British Honduras to

14 FIELDIANA: BOTANY, VOLUME 24

Salvador and Panama; West Indies; South America; Old World
tropics.

Plants perennial, prostrate to ascending, green, the stems branched, rooting
at the nodes, usually glabrous or nearly so, the roots fibrous; leaves somewhat
paler beneath, lanceolate or ovate-lanceolate, 2.5-6 cm. long, 1-2 cm. wide, acute

FIG. 4. Commelina diffusa. A. Habit of plant; X J^. B. Portion of inflores-
cence; X 2.

to acuminate, rounded at the base, glabrous or nearly so; sheaths thin, scarious,
1-1.5 cm. long, 3-4 mm. broad, white-ciliate, glabrous; peduncles sometimes
almost obsolete, commonly 1-5 cm. long, spreading or ascending, glabrous or
sparsely puberulent above; spathes ovate-lanceolate, acute or acuminate, green,
8-20 mm. long, 5-10 mm. wide, glabrous, sometimes ciliate below; lower raceme
of the inflorescence 1-3-flowered; sepals delicate, green, scarious-marginate, 3-4
mm. long; petals blue, the upper 2 larger, 4-5 mm. long; capsule normally 5-seeded;
seeds black, reticulate. (Fig. 4.)

The Maya name of Yucatan is recorded as "bachaxiu." This
species has been reported from Guatemala as C. nudiflora L. and
C. longicaulis Jacq. For a discussion of the nomenclature of the
species see Merrill, Journ. Arnold Arb. 18: 64. 1937. This plant and
others of the genus are said to be much eaten by cattle. It is one
of the common dooryard weeds of Central America.

STANDLEY AND STEYERM ARK: FLORA OF GUATEMALA 15

Commelina erecta L. Sp. PI. 41. 1753. C. erecta var. typica
Fernald, Rhodora 42: 438. 1940. C. virginica of many authors, not
L. Hierba de polio ; Lochoch ; Ticuquito; Canutillo.

Moist or wet thickets or forest, often a weed in cultivated ground,
especially in cafetales, 1,300 meters or lower, most common at the
lower elevations; Pete*n; Izabal; Zacapa; Chiquimula; Jutiapa;
Santa Rosa; Escuintla; Suchitepequez; Retalhuleu; Quezaltenango;
Huehuetenango. Eastern and southern United States; Mexico;
British Honduras to Salvador and Panama; West Indies; South
America.

Stems slender, erect or decumbent, arising from a cluster of fleshy-fibrous
roots, simple or branched, 40-120 cm. long, glabrous; leaves lanceolate to ovate-
lanceolate, acute or acuminate, rounded at the base, abruptly contracted into the
sheath, usually glabrous; sheaths thin, membranous, glabrous or sparsely pubes-
cent, 22-35 mm. long, 4-6 mm. broad, the orifice sparsely ciliate; peduncles usually
less than 1 cm. long; mature spathes firm-membranaceous, ovate, green, shortly
and abruptly acuminate, 2-3.5 cm. long, glabrous or hirtellous; sepals 4-5 mm. long;
petals blue to very pale blue, 1-2.5 cm. long; seeds grayish brown, smooth, puberu-
lent, about 4 mm. long and 3 mm. broad.

Called "matalin" in Veracruz. This species, with its wide range
in both tropical and temperate regions, is variable in size, shape, and
pubescence of the leaves. Most of the Guatemalan material has
spathes 2-2.5 cm. long, and averaging smaller than the dimensions
given by Fernald (2.5-3.6 cm. long). The following varieties and
forms may be recognized in the Guatemalan material.

Commelina erecta f. intercursa Fernald, Rhodora 42: 439.
1940.

Moist or wet thickets, 200-500 meters; Zacapa. Of occasional
occurrence through much of the range of the species.

Like the typical form of the species, but more abundantly pubescent, the
spathes densely villous near the base with long white hairs; petals usually blue.

Commelina erecta f. Candida Standl. & Steyerm. Field Mus.
Bot. 23: 33. 1944.

Known in Guatemala only from the type, shaded slopes, Zacapa,
between Zacapa and Santa Maria, 200 meters, Steyermark 29267.
Chihuahua.

Spathes usually villous at the base with long white hairs, as in f. intercursa;
petals white.

Commelina erecta var. angustifolia (Michx.) Fernald,
Rhodora 42: 439. 1940. C. angustifolia Michx. Fl. Bor. Amer. 1:

16 FIELDIANA: BOTANY, VOLUME 24

24. 1803. C. virginica var. angustifolia C. B. Clarke in DC. Monogr.
Phan. 3: 183. 1881. Matalin de monte.

Usually on rather dry, brushy slopes or plains, in thickets or
thin forest, or in open places, 850 meters or lower; Izabal; Jutiapa;
Suchitepequez; Retalhuleu. Southern United States; Mexico;
British Honduras, and probably farther southward.

Stems glabrous or sparsely pubescent; leaves and sheaths glabrous to some-
what villous; similar to the typical variety and differing chiefly in the narrower and
often much smaller leaves; petals pale to deep blue.

Commelina erecta var. angustifolia f. albina Fernald,
Rhodora 42: 439. 1940.

Not yet known definitely in Guatemala, but to be expected there;
occurring in the Atlantic lowlands of Honduras, and scattered
through the general range of the variety.

Spathes glabrous to short-pubescent, as in C. erecta var. angustifolia, but
the petals white.

Commelina erecta var. angustifolia f. crispa (Wooton)
Fernald, Rhodora 42: 440. 1940. C. crispa Wooton, Bull. Torrey
Club 25: 451. 1898. C. erecta var. crispa Palmer & Steyerm. Ann.
Mo. Bot. Card. 22: 417. 1935. Hierba de polio.

At 1,100 meters or lower; Izabal (between Los Amates and
Quirigua); Guatemala (Lago de Amatitlan). Scattered throughout
the range of the variety.

Sheaths, stems, and leaves often pubescent; leaves narrowly lanceolate or
elliptic-lanceolate, 3-20 mm. wide; spathes sparsely or densely villous, with longer
white hairs at the base.

Commelina erecta var. angustifolia f. cana Standl. &
Steyerm. Field Mus. Bot. 23: 32. 1944.

Known definitely only from the type, Zacapa, along the railroad
between La Fragua and Estanzuela, 200 meters, Steyermark 29136.

Similar to f. crispa in the pubescence of the sheaths, and differing only in the
white petals.

Commelina erecta var. angustifolia f. villosa (C. B. Clarke)
Standl. & Steyerm. Field Mus. Bot. 23: 33. 1944. C. virginica var.
vittosa C. B. Clarke in DC. Monogr. Phan. 3: 183. 1881. C. elegans
var. hirsute, Standl. Field Mus. Bot. 8: 136. 1930.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 17

Rocky slopes or open forest or plains, sometimes in pine forest,
2,000 meters or lower; Alta Verapaz; Zacapa; Chiquimula; Huehue-
tenango. British Honduras; Honduras; South America.

Stems glabrous or villosulous; leaves lanceolate to ovate-lanceolate, 5-18 mm.
wide, usually villosulous on both surfaces, or the lower surface less pubescent;
sheaths villosulous; spathes usually rather densely pubescent; petals blue or white.

Commelina robusta Kunth, Enum. PL 4: 52. 1843.

In forest, at or little above sea level; British Honduras (Sittee
River, W. A. Schipp 746); Costa Rica; South America.

Plants rather stout, erect or decumbent, the stems 60-80 cm. long, glabrous;
leaves large, rather thin, broadly lanceolate, long-acuminate, rounded at the
narrow, somewhat unequal base, mostly 8-12 cm. long and 2.5-3.5 cm. wide,
transverse-veined between the nerves, glabrous or nearly so; sheaths 15-17 mm.
long, 3-4 mm. broad, ciliate, glabrate; peduncles less than 1 cm. long; spathes
several and crowded, usually terminal or nearly so, cordate-ovate, short-acute,
firm-membranaceous, transverse-veined between the nerves, glabrous, 20-27 mm.
long, about 20 mm. wide; petals blue or white; capsule normally with 5 seeds;
seeds of the ventral cell grayish, foveolate, 2-2.5 mm. long.

Commelina robusta f. vestita (C. B. Clarke) Standl. &
Steyerm. Field Mus. Bot. 23: 33. 1944. C. monticola Seub. in Mart.
Fl. Bras. 3, pt. 1: 264. 1855. C. monticola var. vestita C. B. Clarke
in DC. Monogr. Phan. 3: 162. 1882.

In forest, 400 meters; Quiche* (Finca Chaila, Zona Reina, A. F.
Skutch 1802). South America.

Like the typical form of the species but the leaves sparsely villosulous on
the upper surface, more densely villosulous on the lower surface; petals white in
the Guatemalan plant.

Commelina Standleyi Steyermark in Standl. & Steyerm.
Field Mus. Bot. 23: 33. 1944.

On rather dry slopes in pine forest, 1,000-2,000 meters; endemic;
Zacapa (Sierra de las Minas; type collected along trail between Rio
Hondo and summit of mountain at Finca Alejandrla, Steyermark
29644).

Plants erect, slender, sparsely leafy, the stems simple or sparsely branched,
50 cm. high or more, glabrate below, scaberulous above; leaves rather thick and
firm, linear, 7-13 cm. long, 2.5-4.5 mm. wide, attenuate, little narrowed at the
base, passing directly into the sheath, densely scaberulous on both surfaces,
scabrous-ciliate; lower and middle sheaths 22-40 mm. long, 2.5-3 mm. broad,
hirsutulous to glabrate, the orifice and margin white-ciliate; peduncles terminal,
1-2.5 cm. long, scabrous-hirtellous; spathe falcate-ovate, long-acuminate, less
than half as wide as long, 2-2.5 cm. long, 1-1.3 cm. wide, scabrous-hirtellous, not

FIELDIANA: BOTANY, VOLUME 24

evidently transverse-veined, the margins thickened and scabrous-hirtellous, the
closed basal portion about 5 mm. long.

DICHORISANDRA Mikan

Perennial herbs, usually large and coarse, the stems simple or branched,
erect or often scandent leaves mostly rather broad, vaginate at the base; inflores-

FIG. 5. Dichorisandra hexandra.
stem; X 1 A- B. Flower; X 2.

A. Habit of upper portion of flowering

cences usually terminal, racemose-paniculate, the racemes solitary, the branches
short, bracteate at the base; bracts linear or narrow, flat; pedicels short, often
subtended by ovate bractlets; flowers almost regular, the sepals 3, free, the outer
one cucullate, oblong-elliptic, the 2 inner ones oblong-falcate, green, sometimes
scarious-marginate, or often petaloid, persistent in fruit and erect; petals 3, free,
the 2 inner ones subfalcate, short-unguiculate, blue or purple, marcescent; stamens
usually 6, equal or those of the outer series slightly shorter, those of the inner
series subadnate to the petals, the filaments rather short, not barbate; anthers
elongate, 2-celled, the cells narrow, parallel, contiguous, dehiscent by apical pores;
ovary sessile, 3-celled, the cells generally 4-5-ovulate; capsule ovoid-trigonous,
3-celled, loculicidally 3-valvate; seeds several in each cell, covered with a pulpy

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 19

aril, subpeltate, rounded or angulate, the testa crustaceous, rugose or almost
smooth.

Species about thirty, in the American tropics, mostly in South
America. Only one is known from continental North America.
The South American ones are closely related, and their true number
probably is less than thirty.

Dichorisandra hexandra (Aubl.) Standl. in Standl. & Calderon,
Lista PI. Salvad. 48. 1925. Commelina hexandra Aubl. PL Guian.
35. pi. 12. 1775. D. Aubletiana Roem. & Schult. Syst. Veg. 7: 1181.
1830. Hierba de polio.

Moist or wet forests or thickets, 900 meters or lower; Alta
Verapaz; Izabal; Jutiapa; Santa Rosa; Suchitepequez; Huehue-
tenango. Southern Mexico; British Honduras to Salvador and
Panama; South America.

Plants usually scandent or subscandent, 1-5 meters long, the stems generally
branched, glabrous; leaves thick-membranaceous, oblong-elliptic to elliptic-lanceo-
late, 6-18 cm. long, 2-7 cm. wide, long-acuminate, often abruptly so, unequally
and broadly cuneate or somewhat rounded at the base, somewhat paler beneath,
finely many-nerved, scarcely or not at all transverse-striate, glabrous, short-
ciliate at the base, subsessile or narrowed into a petioliform portion 1-2 mm.
long; sheaths narrowly cylindric, close, glabrous to short-villosulous, especially
near the margins, the orifice sparsely short-ciliate, 1-2 cm. long, 3-5 mm. broad;
inflorescence racemose-paniculate, usually dense and many-flowered, sessile or
pedunculate, 4-8 cm. long; bracts at the base of the branches linear, often shorter
than the upper branches; axis of the panicle glabrous to densely hirsutulous;
sepals 6-8 mm. long, usually glabrous; petals oblong, obtuse, blue or purplish blue,
the margins and base edged with white, 10-13 mm. long; capsule 10-12 mm.
long, 9-10 mm. broad; seeds 4-5 in each cell, the aril bright orange. (Fig. 5.)

PHAEOSPHAERION Hasskarl

Plants branched, with elongate, ascending or scandent, herbaceous stems,
the leaves broad; peduncles solitary or aggregate toward the tips of the branches,
arising from conspicuous spathaceous bracts, bifid; sepals 3, almost free; petals 3,
free, the exterior one short-unguiculate, the 2 inner ones long-unguiculate, blue,
marcescent; perfect stamens 3, the filaments elongate, glabrous, the anthers
hastate-triangular; sterile stamens 2, opposite the inner petals; ovary 3-celled,
the dorsal cell 1-ovulate, the 2 ventral cells 2-ovulate; capsule globose or ellipsoid,
indehiscent, pergamentaceous, smooth and lustrous, blue, black, or white, contain-
ing normally 5 seeds.

About five species, in tropical America. No others are known
from Central America. The name Athyrocarpus has been used by
most authors of recent years for this group but, as pointed out by
Woodson, that name, appearing first in 1853, was not formally
published.

FIG. 6. Phaeosphaerion leiocarpum. A. Habit of upper portion of plant;
X 1 A- B. Stamen; X 5. C. Flower; X 5. D. Fruit; X 3.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 21

Spathes long-pedunculate, axillary; fruit dark blue P. leiocarpum.

Spathes short-pedunculate, chiefly terminal; fruit whitish.

Spathes glabrous or nearly so P. persicariaefolium.

Spathes densely rufous-hirsute P. rufipes.

Phaeosphaerion leiocarpum (Benth.) Hassk. Flora 49: 212.
1866. Commelina leiocarpa Benth. Bot. Voy. Sulph. 176. 1844
(type from Tigre Island, Golfo de Fonseca, Honduras). Athyrocarpus
leiocarpus Benth. & Hook, ex Hemsl. Biol. Centr. Amer. Bot. 3:
386. 1885. Lochoch, Lochoch amargo (fide Aguilar); Jicuquita.

Moist or wet thickets or mixed forest, often in rocky places,
1,900 meters or lower, most frequent at low elevations; Alta Verapaz;
Zacapa; Chiquimula; Jalapa; Jutiapa; Santa Rosa; Escuintla;
Guatemala; Sacatepe"quez; Chimaltenango; Retalhuleu; Huehue-
tenango. Southern Mexico; Salvador to Panama; northern South
America.

Plants large and diffuse, often much branched, frequently scandent over shrubs,
the stems scabrous or villosulous to glabrate, often purplish; leaves thin, ovate to
oblong-lanceolate, 2.5-10 cm. long, 2-3 cm. wide, long-acuminate, rounded or
subcordate at the base, sparsely pilose or glabrate, subsessile or abruptly contracted
into a petiole 3-7 mm. long, narrowed below into a purple-spotted, cylindric,
scarcely inflated, sparsely or densely pubescent, ciliate sheath 9-17 mm. long and
2-4 mm. broad; peduncles 1-3.5 cm. long, pubescent or glabrate; spathes 2-5.5
cm. long, ovate or lanceolate, long-acuminate, cordate at the base, finely pubescent
or glabrate; racemes 3-5-flowered; petals blue or pale blue; fruit globose, dark
blue or black, 6-7 mm. in diameter. (Fig. 6.)

Called "hierba de polio" in Salvador.

Phaeosphaerion persicariaefolium (DC.) C. B. Clarke in
DC. Monogr. Phan. 3: 137. 1881. Commelina persicariaefolia DC.
in Red. Liliac. 8: pi. 472. 1816. Athyrocarpus persicariaefolius
Hemsl. Biol. Centr. Amer. Bot. 3: 386. 1885. A. persicariaefoliiis f.
tetraspermus Donn. Smith, Enum. PI. Guat. 6: 54. 1903, nomen
nudum.

Moist or wet thickets and mixed forest, 200-900 meters; Alta
Verapaz; Jutiapa; Huehuetenango. Southern Mexico; Costa Rica;
Panama; West Indies; South America.

Stems elongate, sometimes a meter long or more, erect or reclining on other
plants, more or less branched, often rooting at the lower nodes, glabrous or nearly
so; leaves lanceolate, 8-16 cm. long, 2.5-4 cm. wide, long-acuminate, gradually
narrowed to the base, sparsely pubescent or glabrous, subsessile or narrowed into
a petiole 3-4 mm. long, the sheath sparsely pubescent or glabrate, rufous-ciliate;
peduncles less than 1 cm. long, terminal; spathes 2-3 cm. long, acute, rounded

22 FIELDIANA: BOTANY, VOLUME 24

at the base, densely and finely pubescent or glabrate; racemes densely flowered;
fruit about 5 mm. in diameter, white.

Phaeosphaerion rufipes (Seub.) Standl. & Steyerm., comb,
nov. Commelina rufipes Seub. in Mart. Fl. Bras. 3, pt. 1: 265. 1855.
P. persicariaefolium var. rufipes C. B. Clarke in DC. Monogr. Phan.
3: 137. 1881. Athyrocarpus rufipes Standl. in Standl. & Cald. Lista
PI. Salv. 47. 1925.

Moist or wet forest and thickets, 1,200 meters or lower; Pete*n;
Alta Verapaz; Santa Rosa; Escuintla; Chimaltenango. Salvador;
Costa Rica; Panama; South America.

Plants suberect to procumbent, the stems slender, rooting at the nodes,
glabrous or sparsely puberulent; leaves ovate-lanceolate or lanceolate, 7-10 cm.
long, 2-3 cm. wide, long-acuminate, obtuse or rounded at the base, villous to
glabrate beneath, usually glabrate above, the sheaths rufous-hirsute near and
along the margins or over the whole surface; peduncles about 5 mm. long; spathes
1.5-2 cm. long, acuminate, densely rufous-hirsute, especially near the margins, or
over the whole surface; racemes densely flowered; fruit 6 mm. in diameter, white.

Called "zapupa" in Salvador.

RHOEO Hance

Perennial herbs with short stems, or sometimes acaulescent, glabrous; leaves
large and rather broad, imbricate at the base; peduncles axillary, sometimes
divided, terminated at the apex by 2 large boat-shaped bracts, the flowers numer-
ous, included within the bracts, umbellately congested, the pedicels subtended at
the base by sheathing ovate bractlets; sepals 3, free, ovate-lanceolate, somewhat
petaloid, marcescent; petals 3, free, ovate, marcescent; stamens 6, hypogynous,
subequal, all fertile, the filaments barbate; anther cells oblong, separated by a
subquadrate connective; ovary sessile, ovoid, subtrigonous, 3-celled, the cells
1-ovulate; capsule 3-celled, or by abortion 2-celled, loculicidally 3-valvate; seeds
solitary, oblong-ellipsoid, rugose, the hilum ventral.

The genus consists of a single species.

Rhoeo discolor (L'Her.) Hance ex Walp. Ann. Bot. 3: 660.
1852-53. Tradescantia discolor L'He>. Sert. Angl. 8. pi. 12. 1788.

Moist or wet forest or thickets, often on rocky bluffs, said to be
common on old Maya ruins, at or little above sea level; Pete*n;
often grown for ornament in Guatemalan gardens, chiefly in the
tierra caliente. Yucatan Peninsula of Mexico; British Honduras;
West Indies.

Plants usually erect, somewhat fleshy, the stem glabrous, 10-20 cm. long or
almost none; leaves densely imbricate, few or numerous, linear-lanceolate, 20-35
cm. long, 3-5.5 cm. wide, acuminate, slightly narrowed to the sessile base, glabrous,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 23

green on both surfaces or often dark purple beneath; sheaths large, glabrous, or
sparsely pilose at the orifice, as much as 4 cm. broad; peduncles often branched
above, 2-4.5 cm. long, the bracts cymbiform, broadly ovate, opposite, glabrous,
acute, 3-4.5 cm. long; flowers numerous, the inflorescence shorter than the bracts
and included in them; sepals ovate-lanceolate, 3-4 mm. long; petals white, ovate,
longer than the sepals, 5-8 mm. long; seeds rugose, 3 mm. long, 1-1.5 mm. broad.

In Honduras, and probably elsewhere, this plant is known by
the picturesque and appropriate name "senoritas embarcadas."
From the form of the plant with bright purple leaves there may be
obtained a colored decoction that is used by the Yucatecan Mayas
as a cosmetic. It is probable that the plant is or has been used also
for coloring various articles.

TINANTIA Scheidweiler

Erect herbs, usually annual, simple or branched; leaves large, mostly elliptic,
usually pubescent, vaginate at the base; peduncles solitary and terminal on the
branches, the peduncles subumbellate, 1-4-fid, the pedicels densely racemose
or subumbellate, bracteolate or naked at the base; sepals 3, free, elliptic, green,
erect and persistent in fruit; petals 3, free, subequal, short-unguiculate, elliptic,
blue or purple, marcescent; stamens 6, free, all fertile, 3 of the filaments longer,
naked above, the anthers oblong, the 3 shorter filaments barbate at the middle,
the anthers smaller and rounded; anther cells parallel or curved, slightly separated;
ovary sessile, 3-celled, the cells mostly 3-5-ovulate, sometimes 2-ovulate; fruit
capsular, dry, 3-celled, loculicidally 3-valvate; seeds 2-4 in each cell, 1-seriate
and superposed, the testa rugose-roughened.

About seven species, in tropical America. Only the following
have been found in Central America but one other is known from
Mexico.

Sepals glabrous.

Stems usually conspicuously retrorse-pubescent; leaves densely and finely
puberulent, especially on the upper surface T. leiocalyx.

Stems glabrous or sparsely puberulent; upper leaf surface glabrous except for

scattered slender hairs near the apex T. leiocalyx f. glabra.

Sepals pubescent.

Petiolar sheath ciliate; middle and upper part of the stem with a vertical line
of puberulence; petals all blue, purple, or rose-colored; sepals 5-11 mm.
long; common peduncle 2.5-8 cm. long; leaves broadly ovate to elliptic,
broadest at or below the middle, 3-11 cm. long, 2-5 cm. wide. . .T. erecta.

Petiolar sheath not ciliate; middle and upper part of the stem without a vertical
line of puberulence; 2 or all the petals white; sepals 9-15 mm. long; common
peduncle 7-18 cm. long; leaves elliptic-obovate, broadest above the middle,
15-22 cm. long, 5-10 cm. wide T. Standleyi.

Tinantia erecta (Jacq.) Schlecht. Linnaea 25: 185. 1852.
Tradescantia erecta Jacq. Coll. Bot. 4: 113. 1790. Tinantia fugax
Scheidw. Allgem. Gartenzeit. 7: 365. 1839. Canutillo; Cana de
Cristo; Plateado.

FIG. 7. Tinantia erecta. A. Habit of upper portion of plant; X
B. Stamens, showing diverse forms within same flower; X 2. C. Pistil; X

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 25

Damp thickets or forest, river banks, or often a weed in cultivated
fields, 1,200-2,600 meters; Alta Verapaz; Chiquimula; Jalapa; Santa
Rosa; Escuintla; Guatemala; SacatepSquez; Chimaltenango; Quiche";
Huehuetenango; Quezaltenango; San Marcos. Mexico; Salvador
and Honduras to Panama; South America.

Plants annual, erect, a meter high or lower, simple or often much branched,
the stems succulent, often purplish, usually glabrous except for a line of puberulence
on one side; leaves thin, mostly 4-12 cm. long, acuminate, acute to rounded at the
base, with scattered hairs on both surfaces, deep green above, paler beneath,
ciliate, usually finely pubescent beneath along the costa, the petioliform base
5-15 mm. long; sheaths membranaceous, ciliate, glabrous or sparsely pubescent,
5-10 mm. long, 3-7 mm. broad; peduncles conspicuously glandular- villous, sub-
umbellate or 2-fid at the apex, 3-20-flowered, the inflorescence 1.5-5 cm. long and
3-7 cm. broad, the bracts conspicuous or absent; pedicels 7-22 mm. long, ascending
in anthesis, spreading or recurved in fruit; sepals subacute, sparsely or usually
densely glandular- villous; petals mostly blue to rose-purple, 1-1.5 cm. long;
capsule 7-11 mm. long, 4-5 mm. broad, rounded at the apex; seeds 2-3 in each cell,
pale gray-brown, coarsely rugose, 3-3.5 mm. long, 2-2.5 mm. broad. (Fig. 7.)

During the rainy season this plant often springs up abundantly
in gardens and grain fields, forming dense stands, but the leaves
and stems wither as soon as there is a scarcity of moisture.

Tinantia leiocalyx C. B. Clarke in Donn. Smith, Bot. Gaz.
18: 211. 1893. Pogomesia leiocalyx Standl. Journ. Wash. Acad. Sci.
17: 161. 1927.

Moist or wet thickets or forest, 500-1,700 meters; Chiquimula;
Retalhuleu (type from Rio Samala, W. C. Shannon 695); Quiche";
Huehuetenango. Southern and western Mexico; Honduras; Costa
Rica.

An erect annual 50-100 cm. high, the stems usually simple, sparsely or densely
villous in the upper half, puberulent below; leaves thin, bright green on both
surfaces, ovate or elliptic, 6-15 cm. long, 2.5-8 cm. wide, broadest at or below the
middle, abruptly acuminate, rounded at the base and abruptly contracted into
a petiole 1.5-5 cm. long, ciliate, finely pubescent on both surfaces; sheaths mem-
branous, ciliate, sparsely or densely villous, 6-8 mm. long, 5-7 mm. broad;
peduncles glabrous, about 3.5 cm. long, subumbellate or corymbiform, sparsely
5-7-flowered, the inflorescence 1.5-4 cm. long and about as broad, the bracts
oblong-lanceolate, acute, glabrous, 2-3 mm. long; pedicels 6-10 mm. long, ascend-
ing to reflexed; sepals subherbaceous, scarious-marginate, 9-13 mm. long, rounded
and broadly obtuse to narrowed at the apex, pale green, often petaloid and pur-
plish tinged; capsule 8-13 mm. long, 4-5 mm. broad; seeds coarsely rugose, 3 mm.
long, 2-2.5 mm. broad.

Tinantia leiocalyx f. glabra Standl. & Steyerm. Field Mus.
Bot. 23: 34. 1944.

26 FIELDIANA: BOTANY, VOLUME 24

At about 1,500 meters; Guatemala (without special locality,
Ignacio Aguilar 118). Mexico.

Stems glabrous or sparsely puberulent; leaves glabrous above except for
scattered hairs near the apex, usually glabrous beneath, finely puberulent above
close to the margins, ciliate, narrowed below into a petiole as much as 4 cm. long;
sheaths long-ciliate, glabrous or sparsely puberulent; peduncles glabrous or puberu-
lent; sepals glabrous, 8-11 mm. long, somewhat narrowed at the apex.

Tinantia longipedunculata Standl. & Steyerm. Field Mus.
Bot. 23: 35. 1944.

Moist thickets or forest, 300 meters or lower; Escuintla (type
from Santa Lucia, Heyde & Lux 6392); Suchitepequez. Central
and southern Mexico; Costa Rica.

An erect herb 30-50 cm. high or taller, the stems 3-4 mm. thick, glabrous to
sparsely antrorse-puberulent; leaves thin, 7-12 cm. long, 3-4.5 cm. wide, acuminate,
narrowed below into a petiole 1-3 cm. long, sparsely or densely appressed-pilose
above, glabrate or sparsely appressed-puberulent beneath, densely ciliate; sheaths
loosely ciliate, glabrate to sparsely appressed-pubescent, 4-8 mm. long, 3-8 mm.
broad; peduncles slender, 5-10 cm. long, finely and antrorsely appressed-pubescent,
simply racemose to 2-fid at the apex, 4-14-flowered, the inflorescence 2-5 cm. long,
3-4 cm. broad, the bracts very small, poorly developed, about 1 mm. long; pedicels
spreading to ascending, 7-10 mm. long, finely puberulent or glabrate; sepals
glabrous, 8-9 mm. long, narrowed and subobtuse at the apex; capsule about
9 mm. long and 5 mm. broad.

Tinantia Standleyi Steyermark in Standl. & Steyerm. Field
Mus. Bot. 23: 35. 1944. Cana de Cristo.

Moist or wet thickets and forest, 900-2,100 meters; Chiquimula;
Sacatepe"quez; Solola; Quezaltenango (type from lower slopes of
Volcan de Santa Maria, between Finca Pirineos and San Juan
Patzulin, Steyermark 33605) ; San Marcos. Costa Rica.

A stout erect herb about a meter high, the stems simple or sparsely branched,
usually much stouter than in other species, as much as 1 cm. thick, glabrous or
glabrate, often dark red below, purplish above; leaves thin, deep green above,
paler beneath, mostly 12-21 cm. long, abruptly acuminate, subsessile or gradually
narrowed at the base into a petiolar portion 1-2.5 cm. long, sparsely or densely and
finely pubescent above, glabrate to densely pubescent beneath; sheaths membran-
ous, glabrous or sparsely villosulous, 1.5-2.5 cm. long, 1-2 cm. broad; peduncles
conspicuously and densely glandular-villous, 2-6-fid at the apex, 16-100-flowered,
the inflorescence 6-13 cm. long, 5-14 cm. broad, the bracts conspicuously developed,
spreading, lanceolate to ovate, acute or acuminate, densely glandular-villous, 3-10
mm. long, 3-4 mm. wide; pedicels often purple, ascending in anthesis, reflexed in
fruit, 10-23 mm. long, densely glandular-villous; sepals dull green, subobtuse,
densely glandular-villous; petals 1 cm. long, all white or the lower one blue or
pale pink; anthers lilac; style yellow, barbate; capsule 10-13 mm. long, 4 mm.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 27

broad; seeds dull brown or gray-brown, coarsely rugose, 2.6-3 mm. long, 2.5 mm.
broad.

This showy plant sometimes forms a ground cover on forested
slopes of the volcanoes. From the common T. erecta it is distinguished
by its stouter and more luxuriant habit, non-ciliate sheaths, dif-
ferently colored flowers, long peduncles, and larger and differently
shaped leaves.

TRADESCANTIA L.

Perennial herbs, erect to prostrate, glabrous or pubescent; flowers umbellate,
the umbels several-many-flowered, sessile or nearly so, variously arranged, sub-
tended at the base by large leaf-like bracts; sepals 3, free, elliptic or oblong, green
or petaloid, persistent; petals 3, free, equal, broad, short-unguiculate, generally
blue or purple, sometimes white, marcescent; stamens 6, free, all fertile, subequal
or the 3 opposite the petals shorter, the filaments barbate or naked; anther cells
ellipsoid or oblong, dehiscent by a longitudinal slit, approximate and almost
parallel or separated by a rather broad connective; ovary sessile, 3-celled, the
cells 2-ovulate; capsule dry, 3-celled, loculicidally 3-valvate; seeds generally 2
in each cell, subpyramidal, usually rugose-reticulate.

Species perhaps forty, all American, in both tropical and tem-
perate regions. Probably all the Central American species appear
in the following list.

Umbels sessile along the branches, each umbel borne in the axis of a leaf-like
bract.

Leaves 12-16 cm. long, flat, not complicate, the margins not obviously thickened;
sepals 3.5-5 mm. long T. velutina.

Leaves 3-11 cm. long, usually complicate, the margins conspicuously thickened
and brown; sepals 6.5-8 mm. long T. crassifolia.

Umbels subtended at the base by 2 leaf-like bracts, pedunculate.
Stems covered throughout with long spreading hairs 1-3 mm. long.7 1 . Standleyi.
Stems glabrous or the hairs appressed or, if spreading, less than 1 mm. long.
Peduncles and upper portion of the stem densely sericeous with appressed
hairs; uppermost leaves densely soft-sericeous on both surfaces.

T. belizensis.

Peduncles and upper portion of the stem glabrous or villosulous with spread-
ing hairs; uppermost leaves glabrous or sparsely villous.

Bracts 2-5.5 cm. long, leaf-like; pedicels 10-20 mm. long; petals usually
blue, sometimes white; leaves not unequal at the base; sepals 5-6 mm.
long.

Petals blue T. guatemalensis.

Petals white T. guatemalensis f. alba.

Bracts 1-2 cm. long, spathe-like; pedicels 4-9 mm. long; petals usually

pink or lilac, sometimes white; leaves very unequal at the base; sepals

3-4 mm. long.

Leaves glabrous or glabrate on both surfaces; spathes glabrous or glabrate.

T. commelinoides var. glabrata.
Leaves pubescent on one or both surfaces; spathes more or less puberulent.

28 FIELDIANA: BOTANY, VOLUME 24

Petals usually pink or lilac; bracts ovate; upper leaves mostly oblong
or ovate-oblong, usually 5-9 cm. long and 2-3.5 cm. wide.

T. commelinoides.

Petals white; bracts almost orbicular or orbicular-ovate; upper leaves
mostly ovate, usually 3-5 cm. long and 1.5-2 cm. wide.

T. commelinoides var. rotundifolia.

Tradescantia belizensis Standl. Field Mus. Bot. 22: 5. 1940.

Moist forest, 1,000-2,000 meters; Chiquimula; Huehuetenango.
Veracruz; Chiapas; British Honduras, the type from Vaca, El
Cayo District, P. H. Gentle 2228.

A branched perennial herb, the stems procumbent or creeping, rooting at the
nodes, covered with very long, soft, spreading or appressed hairs, rarely glabrate;
leaves thin when dried, lance-oblong to ovate-lanceolate, 3-8 cm. long, 1.5-3 cm.
wide, acute or acuminate, unequally narrowed at the base, usually densely pilose
on both surfaces with long soft hairs, the older leaves sometimes glabrate or only
sparsely pilose; sheaths thin, scarious, 9-13 mm. long, 4-5 mm. broad, more or
less densely pilose with long soft hairs, the orifice and margins pilose-ciliate;
inflorescences terminal, on a long or rather short peduncle; bracts similar to the
leaves, unequal, dilated at the base, usually densely soft-pilose; flowers densely
crowded, on slender pubescent pedicels; petals pink, pinkish lilac, or white, 8 mm.
long; stamens unequal, 3 of them shorter, the filaments long-pilose.

Tradescantia commelinoides Roem. & Schult. Syst. Veg. 7:
1176. 1830. Lochoch (fide Aguilar).

Moist or wet thickets or forest, 900-3,000 meters; Alta Verapaz;
El Progreso; Santa Rosa; Escuintla; Sacatepe"quez; Chimaltenango;
Solola; Huehuetenango; Quezaltenango; San Marcos. Central and
southern Mexico; Honduras and Salvador to Panama.

Plants perennial, apparently with fibrous roots, branched, the stems prostrate
to procumbent and rooting at the nodes, sometimes ascending, generally pubescent
in a line along one side, or more densely pilose above, often purplish; leaves very
thin when dried, paler beneath and sometimes blotched with lilac, oblong-lanceolate
to ovate, 5-9 cm. long, 2-3.5 cm. wide, acute or acuminate, somewhat oblique at
the base, the petioliform portion 2-5 mm. long, the uppermost leaves sessile, sparsely
pilose on both surfaces or glabrate; sheaths scarious, long-pilose or glabrous, the
margins ciliate, 5-11 mm. long, 2-6 mm. broad; umbels terminal, 2-bracteate,
usually long-pedunculate, the peduncle slender, 1.5-5 cm. long, more or less short-
pilose; bracts spathe-like, ovate to rounded-ovate, acute or acuminate, rounded
to subcordate at the base, 1-2 cm. long, usually long-pilose at the base, sparsely
pilose or glabrous above, ciliate; umbels few-flowered, the pedicels villous; sepals
pale green, scarious-marginate, 3-4 mm. long, 2 of them usually glabrous, the
third asymmetric and pilose-ciliate on one margin; petals rose-purple to pink or
lilac, 5-7 mm. long; stamens subequal, the filaments barbate below. (Fig. 8.)

Tradescantia commelinoides var. glabrata Brueckner, Notiz-
bl. Bot. Gart. Berlin 10: 59. 1928. T. macropoda Greenm. Proc.
Amer. Acad. 32: 395. 1897.

FIG. 8. Tradescantia commelinoides. A. Habit of upper portion of plant;
X %. B. Two sizes of fertile stamens, and single pistil; X 4 } A. C. Flower; X 2.

30 FIELDIANA: BOTANY, VOLUME 24

Moist forest, 1,200-2,900 meters, or perhaps also at lower eleva-
tions; Escuintla; Suchitepequez; Huehuetenango. Southern Mexico.

A variety distinguished by its glabrous leaves, usually white flowers, and
sometimes larger leaves.

Tradescantia commelinoides var. rotundifolia C. B. Clarke
in DC. Monogr. Phan. 3: 296. 1881. T. gracillima Standl. Field
Mus. Bot. 8: 135. 1930 (type from Tela, Honduras).

Moist or wet thickets or forest, 1,200-1,700 meters; Chiquimula;
Guatemala. Honduras; Costa Rica; Panama.

Distinguished from the species by its proportionately broader and generally
smaller leaves, the upper leaves, especially, ovate, the bracts generally broader
and more rounded, the petals usually white.

Tradescantia crassifolia Cav. Icon. PI. 1: 54. pi. 75. 1791.

Zonji (Huehuetenango).

Open grassy slopes or fields, sometimes in pine forest, 1,600-2,500
meters; Chimaltenango; Solola; Huehuetenango. Widely distributed
in Mexico.

Plants erect or ascending, from tuberous-thickened roots, the stems simple or
sparsely branched, 12-60 cm. high, lanate or densely pilose; leaves thick and
fleshy when fresh, rather thick when dried, 2-ranked, linear-lanceolate to lance-
oblong, 3-11 cm. long, 1-3 cm. wide, acute, at the base slightly broader than
at the middle, the margins thickened, brownish, often undulate, sparsely pilose
or glabrate above, more or less lanate- villous beneath; sheaths passing directly
into the blade, sometimes almost obsolete, 4-8 mm. long, 4-10 mm. broad, lanate-
villous to glabrate; bracts 2, leaf-like, unequal, 1.5-6 cm. long, lanate-pilose or
pubescent beneath, complicate; umbels terminal or axillary, sessile, usually dense,
1-8 umbels present on a stem or branch; pedicels 6-14 mm. long, usually densely
sericeous-lanate or white-pilose; sepals oblong-elliptic, subacute or acute, 6.5-8
mm. long, membranaceous, scarious-marginate, white-lanate or white-pilose;
petals purplish rose or bluish, 1-1.5 cm. long; filaments equal, villosulous; ovary
hirsute.

T. crassifolia var. glabrata C. B. Clarke (in DC. Monogr. Phan.
3: 293. 1881) is a form with glabrous leaves, only the margins lanate.
It is known at present only in Mexico but may be expected in
Guatemala.

Tradescantia guatemalensis C. B. Clarke in Donn. Smith,
Bot. Gaz. 18: 210. 1893. T. anisophylla Standl. Field Mus. Bot.
22: 4. 1940 (type from Volcan de Tacana, Chiapas). T. tacanana
Standl. op. cit. 6. 1940 (type from Volcan de Tacana, Chiapas).
Coholdm (Coban, Quecchi); Hierba de potto; Lochoch (fide Aguilar).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 31

Moist banks or moist or wet forest and thickets, sometimes in
rocky places or along stream banks, 200-2,600 meters; Alta Verapaz;
El Progreso; Zacapa; Jalapa; Jutiapa; Santa Rosa (type from
Santa Rosa, Heyde &Lux 3515); Guatemala; Sacatepe"quez; Chimal-
tenango; Solola; Huehuetenango; Quezaltenango; San Marcos.
Chiapas; Salvador.

Plants perennial, apparently with fibrous roots, the stems procumbent or as-
cending, rooting at the lower nodes, 30-100 cm. long, simple or branched, appearing
glabrous or usually pubescent in a line along one side, the lower internodes rarely
sparsely pubescent; leaves thin when dried, lance-oblong to ovate-oblong, 3-9 cm.
long, 1-2.5 cm. wide, acuminate, rounded or narrowed at the subequal base, densely
short-pilose or glabrous on both surfaces; sheaths scarious, 8-15 mm. long, 5-6 mm.
broad, glabrous or sparsely villous, pilose-ciliate; bracts leaf -like, unequal, deltoid-
ovate to lanceolate, rounded at the base and contracted directly into the sheath,
2-5.5 cm. long, 1-2 cm. wide; peduncles terminal, usually solitary or sometimes 2,
commonly 1-8 cm. long; umbels several-many-flowered, the pedicels flexuous,
recurved after anthesis, 8-20 mm. long, more or less villous with spreading hairs;
sepals narrowly lanceolate or linear-oblong, 5-7 mm. long, 1-2 mm. wide, usually
glabrous except for the barbate apex, rarely pilose on the surface or margins;
petals blue or pale blue, venose, 7-9 mm. long; filaments pilose below; seeds 6,
dark gray, rugulose-tuberculate, about 1.5 mm. in diameter.

Tradescantia guatemalensis f. alba Standl. & Steyerm. Field
Mus. Bot. 23: 37. 1944.

Moist thickets or a weed in cafetales, 1,500-1,800 meters;
Sacatepe'quez (type collected at Antigua, Standley 58025; also col-
lected in a barranco above Duefias).

Differing from the typical form of the species only in having
white petals.

Tradescantia Standleyi Steyermark in Standl. & Steyerm.
Field Mus. Bot. 23: 37. 1944.

Moist or wet forest, often on exposed rocks, 1,000-2,100 meters;
endemic; Zacapa (Sierra de las Minas); Chiquimula (type from
Montana Nonoja, northeast of Camotdn, Steyermark 31696); Hue-
huetenango (region of La Libertad).

A large coarse perennial, the stems erect or ascending, 120 cm. high or less,
simple or usually branched above, stout, 4-6 mm. in diameter, densely hirsute
throughout with lax spreading brownish hairs as much as 3 mm. long; radical
leaves often numerous and forming a large rosette; principal cauline leaves few
or numerous, pale green, oblong-elliptic or ovate-elliptic, mostly 12-18 cm. long
and 5-6 cm. wide, acute, narrowed to the base and gradually passing into the
sheath, the uppermost leaves ovate and broadly rounded to subcordate at the base,
densely hirsutulous on both surfaces with long or short hairs; sheaths 10-15 mm.
long, 7-12 mm. broad, brownish-hirsute, the margins and orifice hirsute-ciliate;

32 FIELDIANA: BOTANY, VOLUME 24

peduncles terminal, 2.5-6 cm. long, forming a very large and open, dichotomous
inflorescence, densely hirsute with spreading brownish hairs; umbels few-flowered,
subtended by 2 spathe-like or leaf-like, ovate or rounded-ovate, subequal bracts
2-3 cm. long; pedicels about 9 mm. long, recurved after anthesis, densely villo-
sulous; sepals ovate, subobtuse, 3.5-5 mm. long, 2.5-3 mm. wide, 2 of them
glabrous or sparsely pilose near the base, the third densely brown-villosulous;
petals lilac; capsule 4 mm. high; seeds usually 6, gray-brown, 1.5-2 mm. long,
finely rugulose.

Tradescantia velutina Kunth & Bouche", Ind. Sem. Hort.
Berol. 12. 1848 (described from plants cultivated in Europe, "serre
de Rivage"; the type specimen formerly in the Berlin Herbarium
is labeled as having been collected in Guatemala by Warscewicz).

Moist or dry, brushy, rocky slopes, sometimes on steep rocky
banks along streams, 250-850 meters; Zacapa; Chiquimula; so far
as known, endemic.

A rather stout, usually simple herb 40-100 cm. high, erect or ascending, the
stems 5-7 mm. in diameter, softly and sparsely or densely villosulous; leaves
2-ranked, dull green and lustrous above (in the fresh state), somewhat paler
beneath, lanceolate or oblong-lanceolate, 12-16 cm. long, 1.5-3.5 cm. wide, acute
to long-acuminate, gradually narrowed or rounded or subcordate at the sessile
base, sparsely or densely short-pilose above, more densely villosulous beneath,
the margins not obviously thickened; sheath loose, 7-10 mm. long, 6-10 mm.
broad, sparsely or densely villosulous; umbels several to many along the upper
part of the stem, sessile, solitary, each subtended by a large leaf-like bract, the
bracts gradually reduced in size upward; pedicels 8-20 mm. long, densely villosu-
lous; sepals 3-5 mm. long, ovate-oblong, subacute, more or less densely villosulous;
petals lilac to bright rose-purple; stamens subequal, the filaments deep rose-lilac,
barbate below.

This plant is plentiful on the dry rocky hills about the divide
on the road between Zacapa and Chiquimula, where so many other
interesting and rare plants are found. It is a showy and handsome
plant, well worthy of cultivation. It grows only during the invierno,
for during the verano it would be impossible for any ordinary plant
to continue growth in this sun-baked locality, one of the most arid
in all Guatemala.

TRIPOGANDRA Rafinesque

Perennial herbs, succulent, the stems simple or branched, prostrate to erect,
the plants small or sometimes large; leaves linear to ovate; flowers umbellate,
or at least appearing so, the umbels long-pedunculate, never subtended by large
leaf-like bracts, the bracts usually very small and narrow, the inflorescences borne
in the axils of the upper leaves; sepals 3, free; petals 3, free, white or colored;
stamens 6, fertile, the 3 outer ones shorter, the anthers with a narrow connective
and parallel cells; 3 inner stamens longer, the anther connective 4-3-angulate, the

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 33

cells thus separated; filaments pilose or glabrous; ovary 3-celled, the cells 1-2-
ovulate; capsule 3-celled, the seeds 1 or usually 2 in each cell.

Perhaps thirty species, in tropical America. By most recent
authors the plants have been referred variously to Tradescantia,
Leptorrhoeo, and Neodonnellia. For a discussion of the genus and
its relationships see Woodson, Ann. Mo. Bot. Gard. 29: 150. 1942.
Probably all the Central American species are included in the follow-
ing account.

Stems naked, the leaves all in a basal rosette T. Warscewicziana.

Stems leafy, the plants without basal rosettes of leaves.

Leaves linear, about 2 mm. wide T. angmtifolia.

Leaves lanceolate or broader, 4-20 mm. wide or wider.

Leaves small, mostly 4-6 mm. wide and 3 cm. long or shorter. Flowers very

small, white T. floribunda.

Leaves large, mostly 1-2 cm. wide and much more than 3 cm. long.

Plants scandent. Petals white T. grandiflora.

Plants never scandent.

Petals normally white; peduncles shorter than the subtending leaves or
scarcely exceeding them.

Sepals and pedicels more or less glandular-pilose T. cumanensis.

Sepals and pedicels glabrous or nearly so .... T. cumanensis f. glabrior.
Petals lilac, pink, or purple; peduncles usually much exceeding the sub-
tending leaves.

Leaves cordate at the base and amplexicaul T. amplexicaulis.

Leaves not cordate at the base.

Peduncles generally in fascicles of 2-5, rarely solitary; principal
cauline leaves 5-13 cm. long, 1.5-3.5 cm. wide; roots at the
nodes generally brown- villous; shorter filaments glabrous or
with a few hairs; uppermost part of the peduncle usually densely
hirtellous.

Leaves glabrous on both surfaces T. elongate.

Leaves pilose on the upper surface, softly villous beneath.

T. elongata f. diuretica.

Peduncles generally solitary; principal cauline leaves 3.5-5 cm. long
and 1-2 cm. wide; rootlets at the nodes usually not brown-
villous; shorter filaments densely barbate at the apex; upper-
most part of the peduncles glabrous, glandular, or sometimes
villosulous.
Stems, sheaths, and lower leaf surfaces villous.

T. disgrega f. pubescens.
Stems, sheaths, and leaves glabrous.

Sepals more or less glandular-pilose, the hairs less than 1 mm.

long T. disgrega.

Sepals glandular-pilose with hairs 2-3 mm. long.

T. disgrega f. glandulosa.

Tripogandra amplexicaulis (Klotzsch) Woodson, Ann. Mo.
Bot. Gard. 29: 152. 1942. Tradescantia amplexicaulis Klotzsch ex
C. B. Clarke in DC. Monogr. Phan. 3: 304. 1881. Descantaria am-
plexicaulis Brueckner, Notizbl. Bot. Gart. Berlin 10: 56. 1927.

Lochoch (fide Aguilar).

34 FIELDIANA: BOTANY, VOLUME 24

Moist thickets or forest, 900-1,200 meters, or perhaps higher;
Jutiapa (near Jutiapa, Standley 75525); Guatemala; reported from
Volcan de Fuego, Chimaltenango. Central and southern Mexico;
reported from Costa Rica.

Plants perennial or perhaps annual, with fibrous roots, the stems erect, 20-50
cm. high, simple or branched, glabrous; leaves thin when dried, the middle and
upper ones ovate, acuminate or long-acuminate, cordate and amplexicaul at the
base, the lower cauline leaves ovate-elliptic to subovate, 2-8 cm. long, 1.5-3 cm.
wide, glabrous; sheaths scarious, ciliolate or glandular-ciliolate, 7-9 mm. long,
3-6 mm. wide; leaves at the summit of the stem reduced to sheaths; peduncles
terminal, slender, 3-7 cm. long, more or less glandular or glabrate, the pedicels
umbellate, 4-8 mm. long, glandular-pilosulous; bractlets ovate, glandular-pilosu-
lous; sepals 4-5 mm. long, ovate, acute or acuminate, glandular-pilosulous or gla-
brate; petals pale lavender to rose-purple, 6-7 mm. long; stamens unequal, the
3 longer filaments dilated upward, sparsely barbate below; ovary sparsely pubescent
at the apex; seeds 1.5 mm. in diameter, brown, rugose.

Tripogandra angustifolia (Robinson) Woodson, Ann. Mo. Bot.
Card. 29: 152. 1942. Tradescantia angustifolia Robinson, Proc.
Amer. Acad. 27: 185. 1892. Descantaria angustifolia Brueckner,
Notizbl. Bot. Gart. Berlin 10: 56. 1927.

Moist or wet thickets or forest, sometimes on rocks, 800-1,600
meters; Huehuetenango (between Santa Ana Huista and Nenton;
between San Ildefonso Ixtahuacan and Cuilco). Southern Mexico.

Plants perennial or perhaps also annual, with fibrous roots, very slender,
erect or ascending, simple or usually branched, glabrous; leaves linear, 2.5-5 cm.
long, about 2 mm. wide, acute, glabrous, the sheaths small, ciliate; flowers small,
about 6 mm. broad, mostly in few-flowered umbels, sometimes solitary, the
umbels on slender peduncles 5 cm. long or shorter, the bracts very small; sepals
ovate, acute; petals pale pink; outer filaments much longer than the inner ones,
glabrous, geniculate, the connective dilated, horseshoe-shaped, the cells small,
orange, transverse; inner stamens shorter, the anthers larger, pinkish, the con-
nective much less developed, the cells parallel or nearly so; seeds triangular,
brown, somewhat radiate-rugose.

Tripogandra cumanensis (Kunth) Woodson, Ann. Mo. Bot.
Gard. 29: 152. 1942. Tradescantia cumanensis Kunth, Enum. PI. 4:
96. 1843. Commelina floribunda HBK. Nov. Gen. 1: 260. 1816,
not Tripogandra floribunda Woodson, 1942. Descantaria cumanensis
Brueckner, Notizbl. Bot. Gart. Berlin 10: 56. 1927. Pie de potto;
Siempreviva; Canutillo; Rosana de llano (fide Aguilar).

Moist or wet thickets or forest, sometimes along streams or in
open fields, occasionally a weed in cultivated ground, especially in
cafetales, 1,500 meters or lower; Alta Verapaz; Izabal; Zacapa;
Santa Rosa; Escuintla; Guatemala; Suchitepequez; Retalhuleu;

B A

FIG. 9. Tripogandra cumanensis. A. Habit of upper portion of plant; X %
B. Flower; X 8. C. One of the inner stamens at extreme left, one of outer
stamens in center, and pistil at extreme right; X 8.

36 FIELDIANA: BOTANY, VOLUME 24

Quezaltenango; Huehuetenango. Southern Mexico; British Hon-
duras to Salvador and Panama; South America.

Plants perennial, with fibrous roots, the stems procumbent to suberect, often
rooting at the lower nodes, green or striped with purple or lavender, 15-50 cm.
tall, glabrous, very succulent; leaves fleshy when fresh, firm-membranaceous
when dried, somewhat paler beneath, oblong-lanceolate or lanceolate, 5-13 cm.
long, 1-2.5 cm. wide, acute to long-attenuate, rounded and abruptly contracted
at the sessile base, glabrous except for the scaberulous margins; sheaths scarious,
7-16 mm. long, 4-9 mm. broad, glabrous, the margins ciliate; peduncles 1-5,
terminal or from the uppermost leaf axils, 1.5-3 cm. long, glabrous or puberulent
in a line along one side; umbels several-many-flowered, the pedicels 2-5 mm. long,
more or less glandular-pilose; sepals pale green, often tinged with lavender,
scarious-marginate, cucullate-obtuse, 3-4 mm. long, sparsely or rather densely
and shortly glandular-pilose; petals usually white, sometimes pale lilac at the base,
4.5-5 mm. long; shorter filaments glabrous; ovary glabrous; seeds 6, gray, areolate-
reticulate, about 1 mm. in diameter. (Fig. 9.)

Tripogandra cumanensis f. glabrior (C. B. Clarke) Standl.
& Steyerm., comb. nov. Tradescantia cumanensis var. glabrior
C. B. Clarke in DC. Monogr. Phan. 3: 306. 1881.

Moist or wet fields or thickets, 850-1,300 meters; Jutiapa; San
Marcos. Honduras; Nicaragua; Panama; Ecuador.

Differing from the typical form of the species only in having glabrous sepals
and pedicels.

Tripogandra disgrega (Kunth) Woodson, Ann. Mo. Bot. Card.
29: 152. 1942. Tradescantia disgrega Kunth, Enum. PI. 4: 97. 1843.
Descantaria disgrega Brueckner, Notizbl. Bot. Gart. Berlin 10: 56.
1927. Tradescantia parvula Brandeg. Univ. Calif. Publ. Bot. 6: 51.
1914 (a depauperate form). Camotillo (Guatemala); Lochoch (fide
Aguilar); Coyontura.

Moist or wet thickets or forest, often in pine-oak forest, sometimes
on sandbars along streams, or a weed in cultivated ground, 600-2,400
meters; Chiquimula; Jalapa; Santa Rosa; Guatemala; Chimalte-
nango; Quiche"; Huehuetenango; San Marcos. Mexico; Costa Rica.

Plants perennial, succulent, with fibrous roots, the stems procumbent to
erect, 15-50 cm. long, simple or branched, often purplish at the nodes, glabrous;
leaves lance-oblong to ovate-lanceolate, membranaceous when dried, 3-5.5 cm.
long, 1-2 cm. wide, acute or acuminate, rounded and contracted at the base,
usually glabrous; sheaths scarious, 8-12 mm. long, 3-5 mm. broad, glabrous,
ciliate; peduncles terminal or from the upper leaf axils, usually solitary or binate,
1-7.5 cm. long, glabrous below, usually sparsely and shortly glandular-pilosulous
at the apex or throughout; umbels usually dense, the pedicels 3-8 mm. long,
glandular-pilose; sepals ovate, acute or subacute, somewhat scarious along the
margins, 3.5-5.5 mm. long, somewhat glandular-pilosulous; petals lilac to deep

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 37

rose, 5-7 mm. long; longer filaments naked, the 3 shorter ones densely barbate
at the apex; ovary glabrous; seeds 6, gray-brown, tuberculate, about 1.5 mm. in
diameter.

The specific name appeared originally as disgrega, but later
writers sometimes have written it Disgrega.

Tripogandra disgrega f. glandulosa Standl. & Steyerm., comb,
nov. Tradescantia disgrega L glandulosa Standl. & Steyerm. Field
Mus. Bot. 23: 36. 1944.

Moist or wet thickets or forest, 1,000-1,800 meters; known only
from Guatemala; Zacapa (type from Sierra de las Minas, along
trail between Rio Hondo and summit of mountain at Finca Ale-
jandria, Steyermark 29751); Chimaltenango (Finca Alameda).

Differing from the typical form in the much longer, gland-tipped hairs of the
sepals, and in the slightly larger sepals, 5-6 mm. long.

Tripogandra disgrega f. pubescens Standl. & Steyerm., comb,
nov. Tradescantia disgrega f. pubescens Standl. & Steyerm. Field
Mus. Bot. 23: 37. 1944. Borraja.

Known only from the type, Guatemala, near Guatemala, 1,485
meters, Jesus Morales R. 1106.

Differing from the typical form and from f. glandulosa in the villous stems,
sheaths, and lower leaf surfaces.

Tripogandra elongata (G. F. W. Mey.) Woodson, Ann. Mo.
Bot. Gard. 29: 152. 1942. Tradescantia elongata G. F. W. Mey.
Fl. Esseq. 146. 1818. Descantaria elongata Brueckner, Notizbl. Bot.
Gart. Berlin 10: 56. 1927. Tzimd (Coban, Quecchi).

Moist or wet thickets or mixed forest, sometimes in pine forest,
often on shaded banks or along streams, 250-2,300 meters; Alta
Verapaz; El Progreso; Zacapa; Chiquimula; Guatemala; Sacatepe"-
quez; Solola; Suchitepequez; Quezaltenango; San Marcos; Huehue-
tenango. Southern Mexico; Honduras; Costa Rica; South America.

Plants perennial, usually procumbent or decumbent and rooting at the lower
nodes, the roots fibrous, the stems ascending to erect, sometimes pendent from
banks, 3-6 mm. in diameter, simple or sparsely branched, 30-100 cm. long, glabrous
or glabrate; leaves lanceolate or lance-oblong, thick and fleshy when fresh, thick-
membranaceous when dried, paler beneath, the principal cauline leaves 5-13 cm.
long and 1.5-3.5 cm. wide, acute or acuminate, rounded and contracted at the base,
glabrous, minutely scaberulous on the margins; sheaths scarious, 10-17 mm. long,
5-9 mm. broad, glabrous, ciliate; peduncles 2-5, terminal, fasciculate, rarely
solitary, 2-8 cm. long, usually pubescent in lines, densely hirtellous near the
apex; umbels dense, the pedicels usually short, 3 mm. long or less, glabrous to

38 FIELDIANA: BOTANY, VOLUME 24

sparsely or densely glandular- villous; bractlets densely clustered at the apex of
the peduncle; sepals 3-4 mm. long, more or less glandular-pilose; petals rose-
purple, lilac, or pink, 7-8 mm. long; longer filaments barbate above, the 3 shorter
ones usually naked; ovary glabrous.

Tripogandra elongata f. diuretica (Mart.) Standl. & Steyerm.,
comb. nov. Tradescantia diuretica Mart, in Spix & Mart. Reise Bras.
1: 281. 1823. Tradescantia elongata var. diuretica C. B. Clarke in
DC. Monogr. Phan. 3: 303. 1881.

Reported by Clarke as collected in Guatemala by Skinner.
South America.

Distinguished from the typical forms of the species by having the leaves
pilose above and softly villous beneath, and by the villous sheaths.

Tripogandra floribunda (Hook. & Arn.) Woodson, Ann. Mo.
Bot. Gard. 29: 152. 1942. Aneilema floribunda Hook. & Arn. Bot.
Beechey Voy. 311. 1840. Tradescantia filiformis Mart. & Gal. Bull.
Acad. Brux. 9, pt. 2: 276. 1842. Leptorrhoeo filiformis C. B. Clarke
in Hemsl. Diag. PI. Mex. 55. 1880. L. floribunda Baill. Hist. PL
13: 218. 1894.

Moist or wet thickets or open fields, sometimes in moist rocky
places, often on sandbars along streams, or a weed in cultivated
ground, 200-1,600 meters; Zacapa; Chiquimula; Jalapa; Jutiapa;
Santa Rosa; Escuintla; Retalhuleu; Huehuetenango. Mexico;
Salvador; Honduras; Costa Rica; South America.

Plants low and very slender, perennial or probably in part annual, with
fibrous roots, the stems simple or branched, ascending or diffusely spreading,
rooting at the nodes, 10-20 cm. long, pale green, glabrous or with a vertical line
of pubescence along one side; leaves broadly linear to narrowly oblong-lanceolate,
mostly 1-4 cm. long and 4-6 mm. wide, pale green, acute, slightly narrowed to
the base, glabrous, ciliate; flowers white or pale bluish, small, the umbels 3-6-
flowered, pedunculate, terminal and axillary; peduncles very slender, 4 cm. long
or shorter, glabrous or sparsely puberulent, ascending or divaricate; pedicels obso-
lete or as much as 8 mm. long, glabrous or sparsely puberulent, the bractlets
minute, 1-2 mm. long, ovate, acute; sepals thin, green, scarious-marginate, 1.5-2
mm. long, villosulous; petals ovate-elliptic, about equaling the sepals; stamens
unequal, 3 of the filaments stouter and longer than the others, glabrous, the
anther cells oblong; ovary ovoid, glabrous; capsule 1.5 mm. long; seeds 3, rugose,
trapezoid.

Tripogandra grandiflora (Donn. Smith) Woodson, Ann. Mo.
Bot. Gard. 29: 153. 1942. Callisia grandiflora Donn. Smith, Bot.
Gaz. 31: 125. 1901. Donnellia grandiflora C. B. Clarke, in Donn.
Smith, Bot. Gaz. 33: 261. pi. 11. 1902. Neodonnellia grandiflora
Rose, Proc. Biol. Soc. Wash. 19: 96. 1906. Hoja de fluxidn (Pete"n).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 39

Moist or wet thickets or forest, 1,000 meters or lower; Pete"n;
Alta Verapaz (type from Cubilgiiitz, Turckheim 7684). Campeche;
British Honduras.

A large scandent herb, the stems glabrous, sometimes 3 meters long or more,
branched, often geniculate, 4-8 mm. in diameter; leaves divaricate, 2-ranked,
elliptic to oblong-lanceolate, firm-membranaceous when dried, 5.5-12 cm. long,
1.5-3 cm. wide, acute or acuminate, rounded and sessile at the base, glabrous,
ciliate at the base; sheath glabrous, long-ciliate; inflorescence often large and
open, terminal, composed of few-flowered pedunculate umbels alternately arranged
in an almost naked panicle 6-13 cm. long, the rachis flexuous or zigzag, glabrous;
lower bracts foliaceous, 1-5 cm. long, the upper ones greatly reduced, small,
subspathaceous; umbels 5-9-flowered, 7-13 mm. long, glabrous; sepals green,
white-punctate, oblong-elliptic, subacute, 5-7 mm. long, 2.5-3.5 mm. wide,
glabrous; petals equal, white, elliptic-oblong, obtuse, 9-10 mm. long, 4-6 mm. wide;
fertile stamens 3.5-7 mm. long, the filaments densely barbate above with long
yellow hairs, the anthers bright yellow, transverse-oval, 1.5 mm. broad, bifid at
the apex, the cells divergent, oblong; ovary oval, the stigma capitellate, papillose;
capsule oval, 5-6 mm. long; seeds usually 2, elongate-oblong, 4 mm. long, rugulose.

The plant is said to be common in climax forest in Pete"n, and it
is often planted in gardens there and elsewhere because of its hand-
some appearance, especially its panicles of rather large, pure white
flowers. The flowers are highly fragrant. The species is the type
of the genus Donnellia, later renamed Neodonnellia.

Tripogandra Warscewicziana (Kunth & Bouche*) Woodson,
Ann. Mo. Bot. Gard. 29: 154. 1942. Tradescantia Warscewicziana
Kunth & Bouche", Ind. Sem. Hort. Berol. Add. 11. 1847 (described
from plants grown in Berlin, said to have been collected somewhere
in Guatemala by Warscewicz). Dichorisandra Warscewicziana
Planch. Hort. Donat. 30. 1854-58. Tradescantia subscaposa C. B.
Clarke in Bonn. Smith, Bot. Gaz. 15: 29. 1890 (type from Santa
Rosa, Baja Verapaz, Turckheim 1213). Spironema Warscewiczianum
Brueckner, Nat. Pflanzenfam. ed. 2. 15a: 171. 1930.

On dry shaded rocks in oak or pine forest, 1,200-1,500 meters;
Baja Verapaz (region of Santa Rosa); Zacapa (Sierra de las Minas,
upper Rio Sitio Nuevo).

Plants subscapose, stout, erect, with a very thick, elongate caudex covered by
old leaf sheaths; stems erect, 10-40 cm. long, simple or furcate, glabrous, naked;
leaves radical, forming a dense rosette, fleshy-coriaceous, green above and mottled
with reddish purple, paler beneath and blotched or flushed with reddish or purplish,
narrowly to broadly oblong, 8-30 cm. long, 3-4 cm. wide, cuspidate-acuminate, ses-
sile and scarcely narrowed at the base, glabrous; inflorescence dense or rather lax,
forming a small or large panicle; lowest bracts sheathing the peduncles, lilac,
ovate-lanceolate, long-acuminate, 3 cm. long or shorter; pedicels lilac, 3-7 mm.
long, glabrous; bractlets ovate, scarious, 1-2 mm. long; sepals lilac, ovate, acute,

40 FIELDIANA: BOTANY, VOLUME 24

4-5 mm. long, glabrous; petals lilac or rose-purple, venose, 5 mm. long; filaments
naked; ovary glabrous.

The plant is in cultivation in Salvador, where it is called "pifia
japonesa." Because of its handsome appearance it has been grown
also in European greenhouses.

WELDENIA Schultes

Plants perennial, glabrous or nearly so, acaulescent or with very short stems,
arising from a cluster of fleshy roots; leaves narrow, sheathing at the base, the
lowest ones reduced to bladeless sheaths; flowers large, white, densely aggregate,
axillary, sessile; calyx tubular, subspathaceous, cleft above on one side, the limb
3-fid; corolla tube very long and slender, much exceeding the calyx, the 3 segments
of the limb ovate, horizontally spreading; stamens 6, equal, inserted in the upper
part of the corolla tube, the filaments linear, naked; anther cells oblong, parallel,
contiguous; ovary free, sessile, ovoid, 3-celled; style filiform, the stigma exserted,
penicillate; ovules about 6 in each cell; fruit capsular.

The genus consists of a single species.

Weldenia Candida Schult. f. Flora 12: 3. pi. 1, A. 1829. Lampra
volcanica Benth. PI. Hartweg. 95. 1842 (type from crater of Volcan
de Agua, Sacatepe"quez, Hartweg, without number). Flor blanca,
Loch (Huehuetenango).

Alpine meadows or hillsides, usually in moist places, often in
rocky situations, 3,300-4,500 meters; Sacatepe"quez (Volcan de
Agua); Totonicapan (Desconsuelo) ; Huehuetenango (Sierra de los
Cuchumatanes) ; San Marcos (Tajumulco, Tacana). High mountains
of central and southern Mexico.

Roots elongate, thick and fleshy; stems stout, erect, sometimes 30 cm. long
but usually short, and the plants often appearing acaulescent; leaves mostly flat
on the ground, linear-lanceolate or liguliform, 5-35 cm. long, 1-3 cm. wide, acute
to attenuate-acuminate, rather thin when dried, green and glabrous above, some-
what paler and sparsely villosulous beneath, especially along the costa, sparsely
ciliate near the base, the sheaths conspicuous, membranaceous, glabrous, 3-4.5
cm. long, 1.5 cm. broad; flowers 10-20, densely clustered at the apex of the stem,
sessile, not bracteate; calyx tube 3 cm. long, slightly ampliate above, sparsely
puberulent; corolla tube 4-6.5 cm. long, 1-1.5 mm. in diameter, the lobes white,
sometimes tinged with bluish, 1-2 cm. long, about 1 cm. wide; anther cells 2 mm.
long.

This is one of the common, characteristic, and conspicuous plants
of the alpine regions, abundant in some areas. The plants flower
only during the rainy months. During the dry season the rosettes
of leaves separate from the roots and become scattered over the
rocky open terrain, resembling cushions of leaves. The Indians of

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 41

the Sierra de los Cuchumatanes boil and eat the lower portions of
the tender leaves as "greens." The junior author sampled this dish
and found it fairly tasty.

ZEBRINA Schnizlein. Wandering Jew

Plants succulent, branched, more or less pubescent, the stems prostrate and
creeping or sometimes pendent; leaves ovate or oblong, scattered along the stems,
vaginate at the base; flowers small, aggregate between the 2 uppermost, somewhat
bract-like leaves, subsessile; sepals 3, connate below into a cylindric tube, petaloid,
white or scarious; petals 3, connate below into a slender tube longer than the calyx,
the blade spreading, ovate or lanceolate, rose-purple, bluish purple, or white;
stamens 6, equal or slightly unequal, inserted in the throat of the corolla, the
filaments barbate with moniliform hairs or glabrate; anther cells rotund, separated
by a narrow oblong connective; ovary ovoid, trigonous, 3-celled, the cells 2-ovulate;
capsule borne on a short recurved stipe, perforating the corolla tube, ovoid-oblong,
smooth; seeds 1-2 in each cell, ovoid, obtuse, somewhat rugulose.

The genus consists of two species, one endemic to Guatemala.

Leaves purple beneath, bluish green with usually 2 longitudinal silvery stripes

above; corolla rose-purple or bluish purple Z, pendula.

Leaves silvery green beneath, deep green above; corolla white Z. huehueteca.

Zebrina huehueteca Standl. & Steyerm. Field Mus. Bot. 23:
213. 1947.

Known only from the type, collected on barranco slopes, along
Rio Trapichillo, between Paso del Boqueron and below La Libertad
and Democracia, Huehuetenango, 1,000-2,100 meters, Steyermark
51016.

A fibrous-rooted perennial, the stems sprawling or ascending, glaucous, 85 cm.
long, simple or slightly branched, 4-8 mm. in diameter, glabrous; cauline leaves
subsessile, firmly membranous, deep green above, silvery green beneath, lanceolate
to elliptic-lanceolate, 9-15 cm. long, 3-6 cm. wide, long-acuminate, abruptly
narrowed at the base into the sheath, glabrous; sheaths 1.5-2.2 cm. long, 5-11 mm.
broad, glabrous to sparsely ciliate at the orifice; inflorescences terminal and axillary,
subtended by 2 bract-like leaves, the smaller included; bracts glabrous, unequal,
sessile, cordate, long-acuminate, 3-8 cm. long; flowers 10-12, subsessile, the sub-
tending bracts broadly ovate, asymmetric, 4 mm. long, glabrous; sepals connate,
the calyx tube 6 mm. long, the lobes 3 mm. long, oblong-lanceolate, acute, glabrous
throughout; corolla white, the tube 1.5 cm. long, 1-1.5 mm. broad, the lobes
elliptic-oblong, obtuse, 6-7 mm. long, 2.5 mm. wide; stamens 6, slightly unequal,
three of them slightly longer than the others inserted at the base of the corolla
lobes; filaments white, bearded in the lower half, 2-2.5 mm. long; anthers white,
the cells rotund, separated by a narrow linear connective; ovary glabrous; capsule
not seen.

Zebrina pendula Schnizl. Bot. Zeit. 7: 870. 1849. Z. pendula
var. villosa C. B. Clarke, Bot. Gaz. 37: 213. 1904 (type from Cubil-

42 FIELDIANA: BOTANY, VOLUME 24

giiitz, Alta Verapaz, Turckheim 8326). Z. Purpusii Brueckner,
Notizbl. Bot. Gart. Berlin 10: 57. 1928. Z. flocculosa Brueckner, op.
cit. 58. Hierba de polio; Adorno de Esquipulas (Huehuetenango) ;
Barbija (Jutiapa).

Moist or wet forest or thickets, frequently growing on rocks in
shaded or open places, or on banks, 2,000 meters or less, chiefly at
lower elevations; Pete"n; Alta Verapaz; Jalapa; Santa Rosa; Guate-
mala; Sacatepe"quez; Retalhuleu; Huehuetenango; doubtless in
several other departments; often planted in gardens for ornament.
Central and southern Mexico; British Honduras to Salvador and
Panama; West Indies.

Plants prostrate, often forming dense mats or colonies, the stems glabrous or
pilose; leaves rather thick and succulent, mostly ovate, 4-10 cm. long, 1.5-3 cm.
wide, acute, rounded at the sessile base, glabrous or sparsely pilose on the upper
surface, glabrous or pilose beneath, ciliate near the base, bluish green with usually
two longitudinal stripes of silver on the upper surface, purple beneath; sheaths
thin, membranaceous, 8-12 mm. long, 5-8 mm. broad, long-ciliate at the orifice,
otherwise glabrous or sparsely villous below; flowers clustered, subtended by two
large leaf -like bracts, the bractlets narrow, ciliate; corolla lobes ovate, obtuse; seeds
gray-brown. (Fig. 10.)

Called "matali" and "sangria" in Salvador. This is much planted
in gardens in Central America, and it is well known in the United
States, especially as a house plant, under the name "Wandering
Jew." Plants growing in exposed dry places, especially on rocks,
often assume a vivid and intense, red-purple coloring throughout.
They often become weeds in moist places about gardens and in
patios.

PONTEDERIACEAE. Pickerel-Weed Family

Reference: E. J. Alexander, Pontederiaceae, N. Amer. Flora 19:
51-60. 1937.

Perennial plants, aquatic or of wet soil, often floating; leaves vaginate, with
broad or narrow blades; inflorescence axillary from the rootstock or stem, spa-
thaceous, the 2 valves of the spathe similar or, when different, the lower leaf-like,
the upper rarely with a small blade; flowers spicate, umbellate, or paniculate, rarely
solitary, the bractlets minute or none; perianth marcescent, salverform or funnel-
form, 6-parted, or with a 6-parted limb, nearly regular or zygomorphic, the tube
well developed; stamens 3 or 6, commonly unequal and dissimilar, the anthers
introrse, basifixed or versatile; ovary free, superior, the stigmas terminal, 3-6-
lobate or 3-6-dentate, the style slender or clavate; ovary 1- or 3-celled, the ovules
numerous or by abortion 1; inflorescence usually decurved in fruit; fruit a 1- or
3-celled capsule, or 1-seeded and achene-like and enclosed in the fleshy, accrescent
base of the perianth.

FIG. 10. Zebrina pendula. A. Upper portion of plant; X 1 A- B- Flower; X

44 FIELDIANA: BOTANY, VOLUME 24

Plants of the family are widely distributed in temperate and
tropical regions of the earth. No other genera are known from
Central America.

Stamens 3; fruit capsular Heteranthera.

Stamens 6; fruit capsular or achene-like.

Ovary with 3 fertile cells, the ovules numerous; fruit capsular; petioles often
inflated Eichhornia.

Ovary with 1 fertile cell, the fruit achene-like; petioles not inflated . Pontederia.

EICHHORNIA Kunth. Water hyacinth

Plants aquatic, usually floating, the leaves petiolate, the petioles often inflated,
the blades broad or rarely wanting; inflorescence spike-like, rarely paniculate or
umbellate, pedunculate or very short-pediceled, the 2 spathe valves unlike, the
upper bractlike but frequently with a small dilated blade; perianth tubular, the
limb 6-parted, somewhat bilabiate, the 3 outer segments narrower than the 3
inner ones; stamens 6, the 3 anterior ones exserted, the 3 posterior ones included
in the throat of the perianth, the filaments of the exserted ones pubescent or
glabrous, those of the included ones glabrous, the anthers versatile; stigma sub-
capitate, shallowly 3- or 6-lobate, pilose; ovary 3-celled, many-ovulate; fruit
capsular, 3-celled, dehiscent; seeds numerous, ovoid, multicostate, obtuse at each
end; embryo cylindric, the endosperm farinaceous.

About a dozen species, in tropical America and Africa. One
other is known from Panama.

Corolla small, less than 3 cm. long; plants leafless; inflorescence surpassed by an

ensiform prolongation of the spathe E. paradoxa.

Corolla large, 3.5-4.5 cm. long; plants leafy; inflorescence not terminated by an

ensiform spathe.
Petioles strongly inflated, often subglobose; plants with a short naked stem

bearing new plants at the nodes; perianth lobes entire E. crassipes.

Petioles not or scarcely inflated, at most fusiform; plants with a continually
growing stem bearing leaves for its whole length; perianth lobes erose.

E. azurea.

Eichhornia azurea (Swartz) Kunth, Enum. PL 4: 129. 1843.
Pontederia azurea Swartz, Prodr. Veg. Ind. Occ. 57. 1788. Piaropus
azureus Raf. Fl. Tell. 2: 81. 1831. Ninfa.

In marshes or shallow quiet water, often in slow streams; in the
mountains at 1,500-1,900 meters, or more common at or near sea
level; Pete"n; doubtless in Izabal; Escuintla; Quezaltenango; Huehue-
tenango. Southern Mexico; Honduras to Panama; West Indies;
South America.

Plants usually rooting in mud and erect, often a meter high, generally forming
large dense colonies, the petioles as much as 30 cm. long, terete but often slightly
thickened and fusiform; leaf blades obovate to orbicular, 15 cm. long or less,
broadly rounded to obtuse at the apex, abruptly short-cuneate at the base; flower

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 45

ispikes 5-15 cm. long, many-flowered, glandular-puberulent, the flowers violet-
blue; perianth tube 2 cm. long, the lobes 1.5-3.5 cm. long; capsule about 1 cm.
Jong; seeds columnar, with 10 narrow wings.

A characteristic plant in open swamps or marshes of the coastal
regions, and doubtless abundant near the coasts of Guatemala,
! although we have few collections from the region. Often it occurs
in other parts of Central America in great abundance, forming exten-
sive and almost pure colonies. The plant grows typically in the tierra
caliente, but we have two collections from the highlands, probably
grown from seeds carried by birds.

Eichhornia crassipes (Mart.) Solms in DC. Monogr. Phan. 4:
527. 1883. Pontederia crassipes Mart. Nov. Gen. 1: 9. 1823. Pia-
ropus crassipes Raf. Fl. Tell. 2: 81. 1837. Balsa; Ninfa; Lechuguilla.

Frequent in marshes and lagoons of the tierra caliente, found
locally in lakes and ponds at much higher elevations, frequently
planted in fountains of patios and gardens, 2,300 meters or lower,
most plentiful at low elevations; Pete"n; Alta Verapaz; Baja Verapaz;
Jutiapa; Santa Rosa; Escuintla; Sacatepe"quez; Chimaltenango;
Solold; Huehuetenango. Florida; Mexico; British Honduras to
Salvador and Panama; West Indies; South America.

Plants floating, the stems very short, the roots numerous, long, pendent,
plumosely branched; petioles 2-30 cm. long, the shorter ones much inflated and
globose, the longer ones less inflated; leaf blades rounded-reniform, often lacerate,
variable in size; flower spikes 4-15 cm. long, puberulent; flowers lilac or rarely
white, the perianth tube 1.5-2 cm. long, the lobes about 3 cm. long; capsule 1.5
cm. long; seeds narrowly 10-winged. (Fig. 11.)

Sometimes known in Salvador as "lechuga" and "lechuga de
concha." The common water hyacinth is frequent in many localities
in Central America, but is far less common than the preceding species;
there is, in fact, some possibility that it may be an introduced rather
than a native plant. In some parts of its range, where it grows
abundantly, it often obstructs navigation, as in the St. Johns River
in Florida (where introduced) and in the Panama Canal, where
constant dredging is necessary to keep it under control. All species
of the genus are handsome and decorative plants because of their
large spikes of beautifully colored flowers.

This species is widespread in small lakes and ponds of the moun-
tains of Guatemala, sometimes almost filling them. For instance,
in the Laguna de Ocubila near Huehuetenango the plants cover large
areas to the exclusion of other vegetation. Near the shore, in shallow
water, all the plants have spindle-shaped narrow petioles; farther

FIG. 11. Eichhornia crassipes. Habit of plant; X

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 47

out, in deep water, they grade gradually into plants with globose
petioles. The alteration is very gradual, as if all the plants were of
the same species. In January, 1941, many of the leaves were
brown, apparently having been frosted severely. There is consider-
able doubt that E. azurea and E. crassipes really are distinct species,
at least as represented in Central America. Spruce states that along
the Amazon when plants lodge and grow on the mur 1 they no longer
produce inflated petioles, having no need for them. Both the stems
and the petioles, especially when inflated, contain large air spaces
and are thus enabled to float on the water.

Eichhornia paradoxa (Kunth) Seub. in DC. Monogr. Phan. 4:
531. 1883. Pontederia paradoxa Kunth in Roem. & Schult. Syst.
Veg. 7: 1144. 1839. E. Schultesiana Seub. in Mart. Fl. Bras. 3, pt.
1:94. 1871.

Known in North America and Guatemala from a single locality,
Chiquimula, near Finca San Jose", southeast of Conception de las
Minas, 1,500 meters, Steyermark 31167, in bogs in pine forest.
Venezuela and Brazil.

Plants aquatic, rooting in mud, about 50 cm. high, leafless; scapes all radical,
10-12 from each plant, erect, very spongy, pale green, 7-10 mm. thick; leaf sheath
ventricose, ovate-oblong, 2.5-3 cm. long, opening longitudinally along the lateral
margin, closed above the middle, prolonged above into a petioliform portion 18-23
cm. long, the upper part of the petiole constricted into a green blade-like organ,
this membranaceous and oblong-lanceolate, 6-7 cm. long, 1-1.2 cm. wide; flowers
5-7, subumbellate at the base of the sheath, 1.5-4 mm. long, 1.5-2 mm. thick,
protruding from the open portion of the sheath; perianth lobes purple with yellow
stripes on the upper lobe; perianth tube 1.5-2 cm. long, the lobes about 1 cm. long;
filaments glabrous; capsule elliptic-lanceolate, trigonous, 14-17 mm. long; seeds
barrel-shaped or columnar, rufous-brown, about 10-costate, 1 mm. long, 0.5 mm.
broad, finely horizontally rugulose-striate.

This species, unknown elsewhere in Central America, reappears
in parts of South America. The South American plant appears to
have a scape half as thick as the Guatemalan specimen, and the spathe
is terminated by a shorter prolongation with a narrower, linear-
instead of oblong-lanceolate foliar portion at the tip. Whether
these differences are real ones can be decided only when more collec-
tions are available for study.

HETERANTHERA Ruiz & Pavon

Plants perennial, usually growing on mud, not in water; leaves with broad
or narrow blades; inflorescence 1-flowered or spicate, pedunculate or sessile;

48 FIELDIANA: BOTANY, VOLUME 24

spathe valves 2, the lower leaf-like, the upper bract-like; perianth almost regular,
salverform, the 3 outer lobes narrower than the inner ones; stamens 3, the middle
one with a larger anther and longer filament than the other 2, the anthers basifixed,
erect; ovary incompletely 3-celled by intrusion of the placentae, many-ovulate;
fruit a dehiscent capsule; seeds very numerous.

The genus is a small one, of tropical America and Africa. One
other Central American species is reported from Panama.

Spathe 1-flowered; leaf blades ovate, rounded at the base H. limosa.

Spathe 2-6-flowered; leaf blades reniform, cordate at the base H. reniformis.

Heteranthera limosa (Swartz) Willd. Ges. Nat. Freunde Berlin
Neue Schrift. 3: 439. 1801. Pontederia limosa Swartz, Prodr. Veg.
Ind. Occ. 57. 1788.

Usually in mud about pools or along small streams or in ditches,
200-3,000 meters; Zacapa; Jalapa; Jutiapa; Quiche"; Huehuetenango;
Totonicapan. United States; Mexico; El Salvador; Honduras;
West Indies; South America.

Plants small, with succulent and spongy, usually creeping stems; petioles
erect, 10 cm. long or shorter, the leaf blades lanceolate to almost orbicular, 1-5
cm. long, rounded or obtuse at the apex, cuneate to subcordate at the base; stipules
5 cm. long or shorter, rounded or emarginate at the apex; lower bract of the spathe
similar to the leaves, the upper bract oblong, 1.5-4 cm. long, caudate; spathes
1-flowered; perianth 2-6 cm. long, the tube 1-4 cm. long, whitish, the lobes violet-
blue or white, linear-lanceolate; capsule oblong or narrowly ellipsoid, 1-2.5 cm.
long; seeds 0.6-0.8 mm. long, gray-brown, 10-12-costate, coarsely striate horizon-
tally.

The Maya name of Yucatan is recorded as "hacolel."

Heteranthera reniformis Ruiz & Pavon, Fl. Peruv. 1: 43. pi. 2.
1798. Ninfa; Lechuguilla; Chispi (Retalhuleu).

Usually in mud, about ponds or lakes or along streams or ditches,
2,000 meters or lower; Alta Verapaz; Baja Verapaz; Izabal; Zacapa;
Chiquimula; Jalapa; Jutiapa; Santa Rosa; Escuintla; Guatemala;
Chimaltenango; Solola; Suchitepequez; Retalhuleu; Huehuetenango.
United States; Mexico; Honduras and Salvador to Panama; West
Indies; South America.

Plants usually creeping on mud, rarely floating; leaves erect, the petioles
1-5 cm. long, the leaf blades 5 cm. wide or smaller, reniform, rounded at the apex,
cordate at the base; flower spikes 5 cm. long or shorter; perianth tube slender,
6-9 mm. long, the limb spreading, 8-12 mm. broad, white or pale blue, the lobes
lanceolate; capsule narrowly conic, 8-12 mm. long; seeds pale yellow-brown,
columnar, 0.5-0.8 mm. long, narrowly 10-winged, the wings evanescent and leaving
ridges at maturity. (Fig. 12.)

FIG. 12. Heteranthera reniformis. A. Habit of plant; X %. B. Flower;
X 3. C. Capsule; X 4%. D. Portion of flower, showing stamens and pistil; X 6.

50 FIELDIANA: BOTANY, VOLUME 24

PONTEDERIA L. Pickerel-Weed

Coarse and often tall herbs of marshes; leaves mostly erect, the blades narrow
or broad, many- veined; inflorescence spike-like, pedunculate; spathe valves very
unlike, the lower leaf-like, the upper bractlike; perianth pilose outside, funnel-
form, the limb bilabiate; stamens 6, the 3 anterior ones exserted, the 3 posterior
ones included in the throat, the anthers versatile; ovary 3-celled, 2 of the cells
abortive and empty, the fertile one with a single ovule; fruit achene-like, enclosed
in the accrescent base of the perianth tube, beaked by the persistent style base.

The few species are all American. One other species is reported
from southern Central America (Panama).

Inflorescence as broad as long or nearly so; plants with elongate floating stems;

perianth tube with elongate glands in the hairs P. rotundifolia.

Inflorescence much longer than broad; rootstock creeping under ground, the
plants acaulescent; perianth tube with globose or ellipsoid glands (or none)
in the hairs.
Basal auricles of the leaves spreading; perianth lobes lanceolate or lance-elliptic,

7-8 mm. long, the resin streaks not prominent P. lanceolata.

Basal auricles of the leaves directed downward; perianth lobes obovate or
oblanceolate, 8-12 mm. long, the resin streaks very prominent. P. sagittata.

Pontederia lanceolata Nutt. Gen. PL 1: 216. 1818.

In marshes, at or little above sea level; Pete*n; Izabal. Eastern
and southern United States; Mexico; British Honduras; Honduras;
Cuba; South America.

Plants stout, usually a meter high or less, with thick creeping rootstocks;
leaves erect, the blades variable in shape and size, lanceolate to cordate or hastate,
18 cm. long or usually smaller; stipules truncate, the costa produced into a mucro;
lower spathe valve leaf-like, the upper one loosely vaginate, more or less recurved,
3-6 cm. long, caudate; inflorescence glabrate or hirtellous, the flowering portion
5-15 cm. long; perianth glabrate to densely glandular-pilose, violet-blue or white,
the tube 5-7 mm. long; fruit 5-6 mm. long, broadly ovoid, the ridges of the crests
with angular obtuse divisions.

Pontederia rotundifolia L. f. Suppl. PL 192. 1781. Lechuga
de agua.

In marshes and borders of slow streams, 1,300 meters or lower;
Alta Verapaz; Izabal. Honduras to Panama; South America.

Plants large and coarse, sometimes floating, usually creeping in mud, the
stems branched, erect, a meter high or lower; leaf blades ovate to rounded-sagittate
or reniform-cordate, as much as 12 cm. long and 18 cm. wide, obtuse or rounded
at the apex, the basal lobes rounded; stipules truncate; lower spathe valve similar
to the leaves, the upper one obovate, acute, erect-spreading; inflorescence short-
spicate, the rachis pilose with long crispate hairs; perianth lilac, pilose outside,
the tube 6-8 mm. long, the lobes 9-12 mm. long; fruit ovoid, rostrate, 7 mm. long,
with spinose-cristate ridges. (Fig. 13.)

52 FIELDIANA: BOTANY, VOLUME 24

Called "balsa" in Salvador. Material of this species, as well as
of the others known from Central America, has often been referred
to P. cordata L., a plant of the United States and Canada. All
the species are large and showy plants with handsome flowers, often
forming large colonies in marshes or along the borders of slow streams.
P. rotundifolia is especially plentiful in ditches and marshes about
Coban and in the North Coast.

Pontederia sagittata Presl, Rel. Haenk. 1: 116. 1827.

Swamps or marshes, usually in open places, sometimes about
the borders of lakes, 500 meters or lower; Alta Verapaz; Izabal;
Zacapa; San Marcos. Southern Mexico; Honduras.

Plants erect, frequently a meter high or more; leaf blades sagittate, often
broadly so, as much as 24 cm. long and 15 cm. wide, mostly smaller, obtuse, the
basal auricles usually long and narrow, directed downward; upper spathe valve
laxly vaginate, the upper portion spreading, 5-9 cm. long, abruptly caudate;
inflorescence glabrous to pilose, the flowering portion 5-15 cm. long; perianth
lilac, sparsely glandular-pilose or glabrate, the tube 6-7 mm. long, the lobes
spreading, 8-10 mm. long, broadly elliptic to ovate, with resinous streaks outside,
the upper middle one with a large yellow blotch; filaments glabrous, the anthers
ovate-sagittate; stigma 3-6-fid.

JUNCACEAE. Rush Family
Reference: Buchenau, Pflanzenreich IV. 36: 1-284. 1906.

Grass-like plants, herbaceous, annual or usually perennial; stems slender,
simple, terete; leaves narrow and grass-like, flat or terete, with open or closed
sheaths; inflorescence simple or compound, paniculate or corymbose; flowers
small, perfect, with or without bractlets; perianth regular, the 6 segments gluma-
ceous; stamens 6 or sometimes 3, the filaments triangular to filiform, the anthers
basifixed, 2-celled; ovary superior, 1- or 3-celled; ovules 3 to many, ascending,
anatropous; stigmas 3; fruit a 1- or 3-celled, loculicidally dehiscent capsule.

This family is one of world-wide distribution, but is more abun-
dant in the temperate and arctic regions than in the tropics. In
Guatemala the species are most commonly found in wet meadows,
along streams and margins of lakes, usually in regions of higher
altitudes. The genus Luzula is most frequently found in moist
coniferous mountain forests or on exposed alpine summits of vol-
canoes and mountain ranges. Of the three genera known from
Central America, two of them are represented in Guatemala.

Plants glabrous; leaf sheaths open; capsule many-seeded Juncus.

Plants pubescent; leaf sheaths closed; capsule 3-seeded Luzula.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 53

JUNCUS L.

Mostly perennial plants, glabrous; stems pithy or hollow; leaves glabrous, the
blades terete or flattened, the sheaths open and usually with 2 auricles at the
summit; inflorescence cymose, paniculate, or glomerate, often unilateral; perianth
segments glumaceous, usually greenish or brownish, the margins membranaceous;
stamens 6 or rarely 3; ovary 1- or 3-celled; capsule 3-celled with a central placenta,
or 1 -celled with parietal placentae, the seeds numerous, sometimes append-
aged.

About 250 species, most abundant and widely dispersed in
temperate and cold regions of both hemispheres. Besides the
Central American species listed here, another is known in Costa
Rica.

Lowest bract of the inflorescence terete, appearing like a continuation of the
stem, the inflorescence apparently lateral; leaves all reduced to bladeless
sheaths.

Stamens 6; perianth segments 5-6 mm. long J. andicola.

Stamens 3; perianth segments 2.5-4 mm. long.

Perianth dark brown to castaneous J. effusus var. aemulans.

Perianth stramineous, greenish, or pale brown J. effusus var. solutus.

Lowest bract of the inflorescence flat or channeled along the upper side, not
appearing like a continuation of the stem, the inflorescence obviously terminal;
leaves with well-developed blades.
Leaves terete, septate.

Style very short, inconspicuous J. microcephalus.

Style elongate, usually very conspicuous J. trinervis.

Leaves flat, not septate.

Leaves 2-4 mm. wide; flowers in dense cymose-paniculate heads; stamens 3.

J. marginatus var. setosus.

Leaves 0.8-1.5 mm. wide; flowers inserted singly or clustered; stamens 6.
Flowers mostly clustered at the tips of the branches of the inflorescence;

perianth segments about equal, 3-4.5 mm. long J. tennis.

Flowers scattered or somewhat secund along the branches of the inflores-
cence; outer perianth segments attenuate-subulate, conspicuously
longer than the inner ones, 5-6 mm. long. .../. tennis f. discretiflorus.

Juncus andicola Hook. Icon. PI. 8. pi. 714- 1848.

Huehuetenango, wet ground along streams, 2,450-3,500 meters
(Soloma; Tunima). South America.

Plants coarse and stout, with thick creeping rhizomes, densely cespitose;
stems 55-145 cm. tall, 5-8 mm. thick at the base and middle, 3.5-4 mm. thick
below the inflorescence, fleshy-subcoriaceous, terete, finely striate; leaves reduced
to bladeless sheaths, the leaf scales (cataphylls) several at the base of the stem,
large, black to yellow-brown, shining, obtuse; involucral bract erect, similar to
and a continuation of the naked scape, 15-21 cm. long, gradually acuminate;
inflorescence compound, dense, laterally spreading; flowers numerous, 5-6 mm.
long, rigid and firm in texture; perianth segments narrowly lanceolate, acuminate,
stiffly erect-ascending, the midrib buff to stramineous, conspicuous, the sides
castaneous to dark brown; stamens 6, the filaments about equaling or slightly

54 FIELDIANA: BOTANY, VOLUME 24

shorter than the anthers; capsule slightly shorter than the perianth, 4-4.5 mm.
long, trigonous-ovate, mucronate, dark brown, almost 3-celled; seeds without
tail-like appendages, 0.8-0.9 mm. long, pale or rich brown.

Juncus effusus L. var. aemulans (Liebmann) Buchenau,
Pflanzenreich IV. 36: 136. 1906. J. aemulans Liebmann, Vid.
Medd. Nat. For. Kjoebenh. 38. 1850. Tule.

Swamps and bogs in open meadows, 2,100-3,400 meters; San
Marcos; Totonicapan; Huehuetenango. Southern Mexico.

Plants forming large clumps, densely cespitose, the rootstocks short-creeping,
with inconspicuous internodes; stems 60-80 cm. tall, 2-4.5 mm. thick at the base
and middle, 1.5-3 mm. thick below the inflorescence, firmly subcoriaceous, terete,
inconspicuously many-striate; leaves reduced to bladeless sheaths, the leaf scales
several at the base of the stem, firmly membranaceous, reddish brown at the base,
obtuse, mucronate; involucral bract erect, 15-30 cm. long, aciculate; inflorescence
compound, loosely many-flowered, laterally spreading; flowers 3-3.5 mm. long,
membranous or becoming firmer in age; perianth segments linear-lanceolate, acumi-
nate, erect-ascending, castaneous or reddish brown; capsule slightly shorter than
the perianth, 2-2.5 mm. long, dull brown, retuse, almost 3-celled; seeds with short
points, without tail-like appendages, 0.6-0.7 mm. long, pale brown.

Juncus effusus var. solutus Fern. & Wieg. Rhodora 12: 90.
1910. Tule.

Borders of ponds, swamps, and wet meadows, or around dried
pools on forested slopes, and infrequently along banks of large
streams, 1,200-1,900 meters, or along the Motagua River descending
to 250 meters; Alta Verapaz; Zacapa; Baja Verapaz; Jalapa; Quiche*.
Eastern, central, and southern United States; Mexico; Costa Rica.

Similar in most respects to the preceding variety; stems 60-130 cm. tall, not
sulcate; leaf scales dark brown to reddish brown at the base; involucral bract
10-40 cm. long; inflorescence mostly loosely flowered; flowers 2.5-3.5 mm. long,
firmly membranaceous; perianth segments erect-ascending, subequal, stramineous
or greenish brown; capsule equaling or slightly shorter than the perianth, 2-2.5
mm. long, dull brown; seeds without tail-like appendages, 0.5 mm. long, pale
brown.

Typical J. effusus with much smaller flowers, non-sulcate stems,
and pale basal sheaths is a European species not known from the
New World. The relative diffuseness of the inflorescence appears
rather variable, and some of the Guatemalan collections examined
seem to intergrade with forms of J. effusus var. compactus Lejeune
and J. effusus var. prolifer Sender. In the region about Coban,
the stems are used for weaving mats. The inside of the stems contains
a white spongy pith.

FIG. 14. Juncus marginatus. A. Habit of plant; X 1 A- B- Flower, opened,
with sepals shown on outside; X 10. C. Mature capsule in position with perianth
segments; X 10.

56 FIELDIANA: BOTANY, VOLUME 24

Juncus marginatus Rostk. var. setosus Coville, Proc. Biol.
Soc. Wash. 8: 124. 1893. J. setosus Small, Fl. Southeast. U. S.
258. 1903.

Borders of swamps, ponds, wet meadows, sometimes in wet open
fields and on banks along small streams, 1,300-1,900 meters; Alta
Verapaz; Zacapa; Quiche"; Huehuetenango. United States and
Mexico; Honduras.

Plants perennial from a bulbous thickened base; culms densely cespitose,
erect, 20-65 cm. tall, 2-3 mm. thick below middle; leaves flat, shorter than the
culms, 2-4 mm. broad; bracts reduced and inconspicuous; inflorescence compound,
with 20-35 heads, rather densely flowered, the heads 2-7-flowered; flowers 2.5-
3.5 mm. long; outer perianth segments broadly lanceolate, abruptly attenuate-
aristate, deep or rufous brown, the inner perianth segments slightly longer and
broader than the outer ones, elliptic-ovate, subobtuse, abruptly aristate, similar
in color to the outer ones but more scarious on margins; stamens 3, not exserted
in fruit; capsule glossy, about equaling the perianth, subglobose-turbinate, deep
brown to castaneous, imperfectly 3-celled; seeds apiculate, 0.5-0.7 mm. long,
strongly ribbed, rufous brown. (Fig. 14.)

Typical J. marginatus Rostk., with the inner perianth segments
obtuse and the capsule dull and lusterless, has not been found in '
Guatemala. Coville (Proc. Biol. Soc. Wash. 8: 123. 1893) referred
Turckheim 431 from Coban to J. marginatus var. aristulatus (Michx.)
Coville. Fernald (Rhodora 37: 156. 1935) and Hermann have shown
that Michaux's J. aristulatus (transferred by Coville as a variety
under J. marginatus) is synonymous with J. biflorus Ell., a species
with elongate nodulose rhizomes and stamens exserted in fruit. Since
the specimens from Coban and elsewhere in Guatemala have bulbous
thickened bases and stamens included in fruit, they must be placed
with J. marginatus.

Juncus microcephalus HBK. Nov. Gen. & Sp. 1: 327. 1815.

Borders of swamps, ponds, and small streams, 1,300-3,100 meters;
Alta Verapaz; Zacapa; Jalapa; Chimaltenango; Huehuetenango.
Mexico; Honduras; Costa Rica; South America.

Plants perennial from a slightly thickened base; culms densely cespitose,
erect, 20-90 cm. tall, 2-3 mm. thick below middle, compressed; leaves subterete,
septate, rather soft, about two-thirds the height of the culms, 2-4 mm. broad;
bracts much reduced or inconspicuous; inflorescence compound, with 12-40 heads,
the heads 2-8-flowered; flowers 3-3.5 mm. long; perianth segments subequal, ap-
pressed-ascending, dull brown along the central portion, gray-buff and scarious
on margins, lanceolate, acuminate; stamens 6; capsule slightly shorter than the
perianth, obovate, obtuse, dull brown, 1-celled; seeds apiculate, without tail-like
appendages, 0.4-0.5 mm. long, pale brown.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 57

Juncus tenuis Willd. Sp. PI. 2: 214. 1800. J. macer S. F. Gray,
Nat. Arr. Brit. PL 2: 164. 1821.

Wet banks, fields, and meadows, along roadside ditches or on
sandbars along streams, sometimes along brooks in forest, 1,400-
3,500 meters; Zacapa; Baja Verapaz; Jalapa; Chimaltenango; Que-
zaltenango; Huehuetenango; San Marcos. Widely distributed in
North America; Mexico; Costa Rica; also in the Old World.

A perennial with wiry roots, densely tufted; stems 5-45 cm. tall, stiff and
wiry, slender, 0.5-1 mm. thick; leaves flat, 0.8-1.5 mm. broad, shorter than the
culms; auricles at the summit of the leaf sheath entire, white, scarious, con-
spicuously produced beyond the point of insertion, 0.5-1.5 mm. long; bracts (at
least the lowermost) exceeding the inflorescence; inflorescence erect, few- to many-
flowered, open; flowers more or less clustered at the tips of the branches, 3-4.5
mm. long; perianth segments lanceolate, acuminate, stramineous to greenish
brown in the middle, scarious on the margins, slightly spreading in age; capsule
slightly shorter than or almost equaling the perianth, ovoid to obovoid, retuse,
dull greenish brown to stramineous (dull brown in age), falsely 1-celled; seeds
short-pointed, without tail-like appendages, 0.5-0.6 mm. long, pale brown.

In the United States this species of rush is exceedingly common,
frequently being found along paths and in grassy places. The more
common form found in Guatemala is the following:

Juncus tenuis f. discretiflorus (Hermann) Fernald, Rhodora
47: 123. 1945. J. macer f. discretiflorus Hermann, Rhodora 40: 82.
1938.

Distribution in Guatemala similar to that of the typical form,
but more common. Eastern United States.

In most respects like typical J. tenuis, but the flowers scattered or secund
along the branches, larger, 5-6 mm. long, the outer perianth segments conspicuously
longer than the inner ones.

For a discussion of the nomenclature of this species see Fernald,
Rhodora 47: 117. 1945.

Juncus trinervis Liebm. Naturh. For. Kjoebenhavn Vid. Medd.
41. 1850.

Moist or wet meadows, often along brooks, 2,500-2,800 meters;
Huehuetenango (Sierra de los Cuchumatanes: Cerro Canana; near
San Mateo Ixtatan). Central and southern Mexico.

Plants erect, with horizontal rhizomes, often producing stolons, the stems
slender, 20-40 cm. high; leaf blades terete or somewhat compressed, evidently
septate; inflorescence compound or decompound, often umbelliform, the heads
usually few, hemispheric or subglobose, chestnut-colored; flowers 4-5 mm. long;
sepals all about equal in length, lanceolate, broadly marginate; stamens 6, some-

58 FIELDIANA: BOTANY, VOLUME 24

what shorter than the sepals, the anthers linear, longer than the filaments; style
long and slender, often projecting beyond the sepals, the stigmas elongate, con-
spicuously exserted; fruit usually conspicuously longer than the perianth, trigonous,
gradually attenuate into the style; seeds obovate, apiculate, lineolate.

LUZULA De Candolle

Perennial plants, the stems more or less leafy; leaves sparsely or densely ciliate,
the blades usually flat, the sheaths without auricles; inflorescence compound,
dense or open, the flowers in heads, spikes, or loose cymes; perianth dark brown;
stamens usually 6, rarely 3, shorter than the perianth; ovary 1-celled; capsule
1-celled, 3-valvate; seeds 3.

About sixty species, chiefly in temperate and arctic regions. All
the known Central American species are listed here.

Inflorescence loose and open, corymbose, the flowers mostly solitary . . L. gigantea.
Inflorescence very dense and crowded, spike-like.

Lowest branch of the inflorescence remote from the other branches; inflorescence

erect L. caricina.

Lowest branch of the inflorescence close to the other branches, appearing to
be part of the main spike-like inflorescence; inflorescence nodding.

J. racemosa.

Luzula caricina E. Mey. Linnaea 22: 418. 1849.

Alpine meadows, 2,500-3,700 meters; Huehuetenango (Sierra de
los Cuchumatanes). Central and southern Mexico.

Plants cespitose, the stems strictly erect, 8-30 cm. tall, 1-1.5 mm. thick,
usually leafy to the inflorescence; leaves grass-like, conspicuously ciliate, 5-20
cm. long, 2-5 mm. broad, callose-obtuse at the apex; inflorescence spike-like, erect,
1.5-4 cm. long, 5-8 mm. thick, the few-flowered glomerules becoming looser in
age, the lowest branch of the inflorescence more or less distant from the other
branches and appearing to be removed from the main inflorescence, elongate-
stipitate; flowers about 3 mm. long; perianth segments subequal, lanceolate,
long-acuminate, castaneous with the upper half of the margins buff; capsule
about equaling the perianth, trigonous-ovate, obtuse; seeds large, 1.5 mm. long,
castaneous, the basilar caruncle large, white-stramineous.

Luzula gigantea Desv. Journ. Bot. 1: 145. 1808.

Cool high coniferous mountain forests, sometimes in sphagnum
bogs, 2,500-3,800 meters; El Progreso; Sacatepe"quez; Chimalte-
nango; Quezaltenango; San Marcos. Mexico; Costa Rica; Panama;
South America.

Plants densely cespitose and stoloniferous; stems copiously leafy at the base,
ascending, 20-90 cm. tall, 1.5-3.5 mm. thick below the middle; leaves bright
green, firmly membranaceous, ascending, mostly 5-25 cm. long, 5-14 mm. wide,
densely ciliate or in age becoming glabrate; cauline leaves 2-4, the basal ones
numerous; lowest bract of the inflorescence leaf-like, shorter than the inflores-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 59

cence; inflorescence loosely corymbose, usually nodding, the branches filiform;
flowers 2-2.5 mm. long; perianth segments subequal, lanceolate, long-acuminate,
dull or dark brown; capsule equaling or slightly exceeding the perianth, trigonous-
ovate, obtuse, deep brown to castaneous; seeds 1-1.3 mm. long, rufous-brown to
castaneous, obliquely obovate, shortly apiculate at the grayish apex.

Luzula racemosa Desv. Jour. Bot. 1: 162. pi. 6, f. 3. 1808.
Sacatillo de pena.

Alpine meadows, exposed rocky summits of volcanoes, and open
pine woods at high elevations, 3,500-4,500 meters; Quezaltenango;
Huehuetenango; San Marcos; reported by Hemsley from Volcan
de Fuego. Mexico. South America.

Plants densely cespitose; stems very leafy at the base, strictly erect, mostly
10-45 cm. tall, 0.5-1.5 mm. thick; leaves deep green, firmly membranaceous to
subcoriaceous, much shorter than the stems, ascending to erect, densely ciliate
or in age glabrate, 2-12 cm. long, 2-5 mm. wide, flat or becoming convolute,
curved, or more or less canaliculate; cauline leaves usually 2, mostly smaller than
the numerous basal leaves; lowest bract prolonged, more or less leaf-like; inflores-
cence conic to oblong-cylindric, nodding, 2-4 cm. long, 4-12 mm. thick, interrupted
near the base, densely flowered, simple or somewhat lobate; flowers 2.5-3 mm.
long; bracts conspicuously exceeding the flowers, long-ciliate; perianth segments
subequal or the outer slightly longer, lanceolate, caudate, the tips conspicuously
hyaline as are the margins, dark brown to castaneous in the central portion and
at the base; stamens mostly 3; capsule shorter than the perianth, broadly ovoid
to trigonous-globose, very shortly mucronate, rufous- brown to castaneous; seeds
1-1.3 mm. long, castaneous to ferruginous, obliquely ovate, shortly apiculate at
the gray apex.

Buchenau lists several varieties, but there does not appear to
be any substantial basis for their segregation.

LILIACEAE. Lily Family

Perennial herbs or shrubs, rarely arborescent, often with rhizomes or bulbs,
the root fibers often thick and fleshy or even tuber-like; stems herbaceous or woody,
sometimes scandent, leafy or naked; inflorescence very variable in form, the flowers
small or often large and showy, usually perfect and regular; perianth inferior and
free from the ovary, or rarely adnate at its base, often marcescent and persistent,
corolla-like, the segments free or more or less united, normally 6, more or less
definitely biseriate; stamens usually 6, hypogynous or affixed to the perianth,
opposite the segments, the filaments free or connate; anthers long or short, basifixed
or dorsifixed, 2-celled, the cells dehiscent usually by longitudinal slits; ovary
superior to partly inferior, usually 3-celled, rarely 1-celled, with axial placentae,
the style filiform or columnar, with a small, terminal, capitate or 3-lobate stigma,
or the styles sometimes 3 and apical; ovules 2-many in each cell; fruit baccate
and indehiscent, more often dry and capsular and loculicidally or septicidally
dehiscent; seeds many or few, sometimes by abortion only 1; endosperm
copious.

60 FIELDIANA: BOTANY, VOLUME 24

A large family, with many species in North America, chiefly in
temperate regions, and well represented in all the continents. All
the genera native in Central America are listed here.

Plants more or less woody, treelike or shrublike, not scandent, the stems bearing

many long leaves.
Leaves mostly 6-12 cm. wide, often reddish or purplish; stems slender, simple;

cultivated plants Taetsia.

Leaves narrower; stems stout, often 10 cm. or more in diameter, often branched.

Flowers about 4 cm. long; leaves stiff and hard Yucca.

Flowers less than 1 cm. long; leaves flexible.

Fruit dry, winged Beaucarnea.

Fruit fleshy, not winged Dracaena.

Plants herbaceous, low, sometimes woody but then with very slender, scandent

stems.
Leaves cauline, no evident basal ones present.

Stems much branched, often scandent Asparagus.

Stems simple.

Flowers very large, 15 cm. or more in length; cultivated plants. . . .Lilium.
Flowers relatively small, 2 cm. long or shorter; native plants.

Leaves linear; stems with usually 2 flowers Calochortus.

Leaves mostly elliptic; flowers very numerous Smilacina.

Leaves all or chiefly basal, or the stems rarely bearing a few leaves.
Flowers in umbels.

Plants without bulbs or corms, cultivated. Flowers 3.5-5 cm. long, blue

or white Agapanthus.

Plants with bulbs or bulb-like corms.

Perianth segments 2-3 cm. long or larger; flowers usually 2-4 Milla.

Perianth segments less than 1 cm. long; flowers usually numerous.

Perianth segments connate near the base; plants without an onion odor.

Nothoscordum.

Perianth segments distinct; plants with an onion-like odor. . . .Allium.
Flowers not in umbels.

Plants with bulbs; perianth segments often with glands within.

Flowers solitary; perianth segments not glandular within Milla.

Flowers numerous; perianth segments with glands within.

Flowers spicate; ovary superior Schoenocaulon.

Flowers racemose; ovary partly inferior Zygadenus.

Plants never with bulbs; perianth segments without glands.

Leaves thick and fleshy, with spinelike teeth; perianth cylindric. . .Aloe.
Leaves not fleshy, unarmed.

Perianth tubular, the lobes much shorter than the tube .... Kniphofia.

Perianth not tubular, the lobes distinct or much longer than the tube.

Leaves stiff and hard, erect, dark green mottled with white or gray;

flowers white or greenish white Sansevieria.

Leaves thin and soft, usually recurved, green.

Flowers 10-12 cm. long Hemerocallis.

Flowers less than 2 cm. long, usually 1-1.5 cm.

Anthers coherent into a tube about the style Echeandia.

Anthers free . . Anthericum.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 61

Several genera besides those listed here may be seen occasionally
in cultivation in Guatemala, especially the Aspidistra (Aspidistra
elatior Blume, native of Japan), grown as a pot plant. Tulips (Tulipa
spp.; "Tulipan") and hyacinths (Hyacinthus; "Jacinto") probably
have been grown in pots, but the spring-flowering bulbous plants
as a rule do not succeed out of doors even in the mountains of
Central America.

AGAPANTHUS L'HeYitier

Large succulent plants with short rhizomes and thick fleshy root fibers;
leaves radical, broadly linear or lorate; scape simple, naked, the flowers numerous
in a terminal umbel, large and showy, usually blue, sometimes white; involucral
bracts 2, broad, caducous, the interior bracts numerous, linear-setaceous; perianth
funnelform, the tube cylindric, usually short, the 6 segments subequal, longer
than the tube, erect-spreading; stamens 6, inserted in the throat, shorter than the
perianth, the filaments filiform, the anthers oblong or oblong-linear, dorsifixed,
bilobate at the base, the cells introrsely dehiscent; ovary sessile, 3-celled, the
style filiform, the stigma small; ovules numerous; capsule oblong, obtuse, somewhat
trilobate; seeds numerous, ascending, compressed, black, the testa produced as
an oblong wing.

About three species in South Africa, one of them grown for
ornament in most tropical regions.

Agapanthus africanus (L.) Hoffmannsegg, Verz. Pflanzenkult.
35. 1824. Crinum africanum L. Sp. PL 419. 1753. A. umbellatus
L'He>. Sert. Angl. 18. 1788. Agapanto.

Cultivated for ornament in the central highlands but not very
common; also at lower elevations, and especially frequent about
Coban, where it perhaps is sometimes naturalized in hedges. Native
of South Africa, but now cultivated in many tropical regions.

A large and rather robust plant, often forming dense clumps, the leaves about
60 cm. long, rather broad, succulent; scape a meter high or less, bearing at the
apex an umbel of 10-30 large, blue or sometimes white flowers; perianth 3.5-5
cm. long.

The plant is a handsome and showy one, with a rather short
blooming season, apparently. It is able to endure neglect and a
cool climate, and was noted as high as the gardens of San Marcos
(above 2,500 meters).

ALLIUM L. Onion, Garlic, Leek

Plants with bulbs, strong-scented, the bulbs tunicate, solitary or sometimes
several and arising from a short rhizome; leaves basal, usually linear, sometimes

62 FIELDIANA: BOTANY, VOLUME 24

terete or fistulose; scape simple, leafless, the flowers umbellate, few or numerous,
on short or elongate pedicels, usually small, sometimes replaced by bulblets; in-
volucral bracts 2 or very rarely 3, often united below or throughout; perianth
segments 6, free, equal or the inner ones slightly larger, spreading or campanulate-
connivent; stamens 6, inserted at the base of the perianth segments, longer or
shorter than the perianth segments, the filaments free or sometimes connate at
the base, the anthers oblong, dorsifixed at the middle, the cells introrsely dehiscent;
ovary sessile or short-stipitate, 3-celled, the style filiform, short or elongate, the
stigma small, sometimes shortly trifid; ovules usually 2 in each cell; capsule
membranaceous, surrounded by the persistent perianth, loculicidally dehiscent;
seeds angulate-compressed or often disklike, the testa membranaceous, black.

A large genus of about 270 species, widely distributed in temperate
and cold regions of both hemispheres, in America none of them strictly
tropical. A single native species has been found in Central America.
Several of the species are of great importance as food plants.

Native plants, the leaves about 2 mm. wide A. glandulosum.

Cultivated plants, the leaves much broader.

Leaves terete, hollow A. Cepa.

Leaves flat, not hollow.

Leaves broad, strongly conduplicate or carinate; flowers white or pinkish;

bulb simple, scarcely more than an enlargement of the stalk. A. Porrum.

Leaves narrow, carinate; flowers purple, often replaced by bulblets; bulbs

small, breaking up into numerous small ones that are covered with dry

scales A. sativum.

Allium Cepa L. Sp. PI. 300. 1753. Ceboll (Quecchi); Sipoix
(Quiche*); Cebolla; Cucut (Maya). Onion.

Native of Persia and adjacent regions, now cultivated in almost
all parts of the earth.

The onion is one of the commonest and most important vegetables
of Guatemala and all Central America. There is, in fact, a common
saying that sin cebollas no hay cocina. It is used either dry or fresh.
No sight is more common on the roads and trails of Guatemala than
the many Indians who carry heavy loads of onions from the produc-
ing regions to the various remote markets of the country. Onions are
grown almost anywhere in Guatemala, and there are large fields,
or fields of beds, of them about Almolonga and Zunil, to supply
the densely populated region of Quezaltenango. The two principal
centers of onion (and garlic) production, however, are Panajachel,
on the shores of Lake Atitlan, and Aguacatan, in Huehuetenango.
About the latter place the extensive irrigated valley is given over
almost wholly to cultivation of onions and garlic, which are carried
either fresh or dry, but especially fresh, to all parts of the country,
and even to Chiapas. During the dry season when the surrounding

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 63

hills are parched and brown, these great onion fields, vivid green,
are a beautiful sight. The plants are watered by men, women, and
children, who toss the water upon the beds from the ditches of clear
water that are conducted through the fields, the same process used
in watering most other vegetables where running water is available.
Most of the onions grown in Guatemala are white, but red ones are
seen occasionally.

Allium glandulosum Link & Otto, Icon. Rar. 1: 33. pi. 17.
1841.

Meadows or open forest, 1,200-3,800 meters; El Progreso; Jalapa;
Chimaltenango; Totonicapan; El Quiche"; Huehuetenango; San
Marcos. Southern Mexico.

Plants with small bulbs usually about 1.5 cm. in diameter, producing long
stolons; leaves often longer than the scapes, sometimes shorter, about 2 mm. wide,
flat; scapes slender, subterete, 20-40 cm. tall, the flowers usually few, on slender
pedicels 2.5 cm. long or shorter; perianth segments deep purple-red, usually with
white margins, sometimes white with lavender or purple stripes, 7-8 mm. long,
linear-oblong; stamens slightly shorter than the perianth; capsule subglobose,
3.5 mm. high.

This is the southernmost representative of the genus in North
America. Several additional species are native in Mexico, and in
the western United States there are many other species.

Allium Porrum L. Sp. PI. 295. 1753. Puerro. Leek.

Native of Europe, now grown in many temperate regions for
its roots and leaves.

Leeks are seldom seen in markets of the United States, but
they are plentiful in those of Guatemala, especially at Quezaltenango
and Guatemala. They are planted to some extent about Coban,
and doubtless also in other regions. They probably are not used
by the Indians, unless it be those who grow them for market. The
center of production seems to be at Almolonga and Zunil, and leeks
are plentiful in the Quezaltenango market, large and fine ones,
fully equal in quality to any grown in the United States. They are
used principally if not wholly for preparing a very good soup that
is common on tables of the upper classes of society.

Allium sativum L. Sp. PI. 296. 1753. Ajo; Cucut (Maya);
Acuc ( Jacaltenango) ; Axu (Quiche"); Hanx, Anx (Quecchi). Garlic.

64 FIELDI AN A: BOTANY, VOLUME 24

Native of Europe, now grown in most temperate and tropical
regions.

Garlic is planted extensively in many or most parts of Guatemala
and consumed all too extensively, at least too generally for Nortl
American palates that are unaccustomed to it. It is found in almost
all dishes, it would seem, except desserts, and often in too great
amounts, even according to Central American standards. Excessive
use of garlic is long established in the Iberian Peninsula, and thos
familiar with the dicta of Don Quixote will recall his admonitions
on the subject to Sancho Panza. Garlic, like onions, is gro\
extensively about Aguacatan and Lake Atitlan. It is used only ii
the dry state, and for transportation usually the tops of the plants
are braided together to form bunches, just as in the United States.
In the markets the cloves often are separated, so that one may
purchase a very small quantity if desired. The name "ajo" appear
geographically as in the case of a caserio of Huehuetenango, calk
Ajal, i.e., a place where ajo is grown. The Indians of Alta Verapz
often carry an onion in their pockets or elsewhere about their clothing
to avert misfortunes and disease.

Allium Schoenoprasum L., Chives, native of Europe, is gro\
rarely in Guatemala but commonly in the United States. The ver
slender and abundant, grass-like leaves are used in the latter country

for flavoring soups and salads.
/

ALOE L.

Succulent plants, without bulbs, the caudex very short or tall and shrubby
or arborescent, sometimes branched; leaves thick, rosulate or often distichous or
scattered, sometimes spinose-dentate; scapes or peduncles axillary or pseudo-
terminal, usually elongate, simple or branched, the flowers mostly small, racemose,
often nutant, solitary within the bracts; perianth cylindric or contracted above
the ovary, the segments connivent or coherent or connate, erect or with spreading
tips; stamens 6, hypogynous, equaling or longer than the perianth, the anthers
oblong or oblong-linear, dorsifixed near the base, the filaments subulate, the cells
introrsely dehiscent; ovary sessile, oblong or ovoid, 3-celled, the style filiform,
the stigma small; ovules numerous; capsule coriaceous, ovoid or oblong, loculicidally
dehiscent; seeds numerous, triquetrous or compressed, sometimes winged, the
testa black.

About one hundred species, in Africa and western Asia, none
native in America. Because of their often bizarre forms, some
species are grown as pot plants, and some of these have found their
way to Guatemala. The following species has become widely dis-
persed in tropical regions because of its medicinal properties.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 65

Aloe vera L. Sp. PI. 320. 1753. S&bito.

Planted commonly in the Pacific bocacosta, and in some places
thoroughly naturalized in hedges and thickets; seen also in the
Oriente, and even as high as Antigua (1,500 meters); probably to
be found in gardens in almost all parts of the country. Native of
the Mediterranean region.

Plants acaulescent or nearly so, producing long stolons; leaves narrowly
lanceolate, 30-60 cm. long, turgid, pale green, the margins with distant spinelike
teeth; scape stout, a meter high or less, bearing a few distant scales; racemes 10-30
cm. long, dense, the bracts lanceolate or ovate, longer than the short pedicels;
flowers yellow, about 2.5 cm. long.

The aloe plant probably was introduced into Central America
soon after the conquest. It has long been used in medicine as a
bitter purgative. The watery juice upon exposure to air soon
hardens, and in this form is the aloes (acibar) of pharmacy. In
recent years it has been found that the fresh sap is an effective
agent in healing persistent sores or wounds, and the leaves are
shipped for this purpose from Florida to the North. In Guatemala
poultices of the crushed leaves are applied to heal sores, and the
sap is applied to boils, to bring them to a head. Women sometimes
wean nursing children by rubbing the bitter sap upon their nipples.
The Maya name is reported from Yucatan as "hunpetskinci."

| ANTHERICUM L.

Plants with short or no rhizomes, bearing a cluster of numerous thick fleshy
fascicled roots, the base of the plant often covered with persistent sheaths of
branched, withered leaves; leaves basal, linear, sometimes broadly so; scape simple
or sometimes with a few leaflike bracts, the flowers slender-pedicellate, racemose,
solitary or fasciculate in the axils of the bracts, these mostly scarious; perianth
marcescent, usually persistent upon the fruit, the segments distinct, spreading,
subequal, 3-5-nerved; stamens 6, hypogynous, shorter than the perianth, the
filaments filiform, the anthers linear, erect, the cells introrsely dehiscent; ovary
3-celled, sessile, the style filiform, the stigma small; ovules 4-8 in each cell; capsule
oblong to globose or shallowly 3-lobate, loculicidally dehiscent; seeds often only
1-2 in the cell, angulate or angulate-compressed, the testa black.

A group of 50 or more species in Europe, Africa, and America.
An additional Central American species occurs in Panama. The
genus is in need of revision, and the species are not well known.

Anthericum Liliago L., sometimes known by the name St.
Bernard's lily, native of southern Europe and northern Africa, is
seen occasionally as a house plant in the patios of Guatemala, and
is rather frequent in cultivation in the United States. It has slender

FIG. 15. Anthericum aurantiacum. A. Habit of plant; X 1 A- B. Portion
of inflorescence; X 1. C. Stamen; X 3. D. Capsule; X 1.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 67

racemes of small white flowers and long narrow leaves. It is note-
worthy for its habit of producing long stolons, usually above ground,
which bear at the apex a tuft of small leaves. By using these for
propagation, new plants are easily obtained. A variety with white-
striped leaves cultivated at Quezaltenango is known there by the
name ' 'listen."

Plants densely hirsute on the stems and leaves A. vestitum.

Plants glabrous.

Anthers only 1 mm. long; perianth 16-18 mm. long A. Skinneri.

Anthers 3-4 mm. long; perianth 10-14 mm. long.

Leaves mostly 5-12 mm. wide A. aurantiacum.

Leaves mostly 15-25 mm. wide A. eleutherandrum.

Anthericum aurantiacum J. G. Baker ex Hemsl. Biol. Centr.
Amer. Bot. 4: 91. 1887; Bot. Jahrb. 8: 209. 1887. A. apodastanthum
Donn. Smith, Bot. Gaz. 19: 265. 1894.

Moist thickets, open pine forest, rocky slopes, or open hillsides,
1,200-3,300 meters; Chiquimula; Jutiapa (type from Rio Paz,
Lehmann 1721); Sacatepe*quez (type of A. apodastanthum from San
Bartolo, 1,500 meters, Heyde & Lux 4644); Chimaltenango; Huehue-
tenango. Honduras; Costa Rica; Panama.

Plants usually cespitose, the root fibers very numerous, thick and fleshy or
sometimes rather slender; leaves thin, bright green, mostly 5-12 mm. wide; scapes
usually simple, sometimes with a single branch, the flowers slender-pedicellate,
orange-yellow; perianth segments 3-nerved, 10-14 mm. long; anthers 3 mm. long;
capsule oblong-oval, about 13 mm. long, the seeds 10 or fewer in each cell. (Fig. 15.)

We have seen no authentic material of A. aurantiacum, but the
description indicates no characters by which it and A. apodastanthum
may be separated, in spite of the fact that Captain Smith, in publish-
ing the latter, states that the type had been seen by Baker, who
reported it to be an undescribed species.

Anthericum eleutherandrum (Koch) H. E. Moore, Gent.
Herb. 8, fasc. Ill: 246. 1951. Anthericum platyphyllum Greenm.
Proc. Amer. Acad. 33: 472. 1898. Chuichuisa.

Brushy banks or hillsides, sometimes in damp pine forest, 1,300-
2,200 meters; Alta Verapaz; Jalapa; Huehuetenango. Mexico.

Plants with dense clusters of leaf fibers at the base; leaves basal, bright green
and rather soft, 15-40 cm. long, mostly 1.5-3 cm. wide, long-attenuate to each
end, glabrous, ciliolate; scapes 40-70 cm. tall or more, usually sparsely branched
above, the flowers numerous, orange-yellow, 2-4 in the axils of each bract, the
pedicels 3-12 mm. long, articulate below the middle; perianth segments 10-12 mm.
long, 3-nerved, oblong, obtuse; filaments muricate, the anthers 3-4 mm. long;
capsule oblong, 10-15 mm. long, emarginate; seeds angulate, black.

68 FIELDIANA: BOTANY, VOLUME 24

Anthericum Skinneri J. G. Baker, Journ. Linn. Soc. Bot. 15:
318. 1876.

Type from Guatemala, collected by Skinner, the locality not
known.

Leaves 15-20 cm. long, 4-6 mm. wide, ciliolate; scape glabrous, as much as
75 cm. tall, bearing a single small bract; raceme lax, 10-12 cm. long, the pedicels
12-16 mm. long, articulate at the middle; perianth segments 3-5-nerved.

Known to the authors only from description. The very small
anthers and unusually elongate pedicels should be distinctive,
especially if the anther size is characteristic and not abnormal.

Anthericum vestitum Baker, in Journ. Linn. Soc. 15: 307.
1876. Anthericum crinitum Standl. Field Mus. Bot. 17: 228. 1937.
Grassy slopes or plains, 1,600-2,000 meters; endemic; Chimal-
tenango (type from Finca Alameda, near Chimaltenango, J. R.
Johnston 714); Huehuetenango.

Roots numerous, fleshy, much thickened; leaves all basal, linear, 5.5-18 cm.
long, 6-7 mm. wide, long-attenuate to each end, hirsute with rather long, thick,
spreading hairs; scape simple, stout, 18-35 cm. tall, hirsute, the flowers few or
numerous, forming an interrupted raceme as much as 17 cm. long, the pedicels
stout and stiff, scarcely 3 mm. long; perianth glabrous, orange-yellow, the segments
12 mm. long, 3-nerved; capsule sessile, glabrous, oblong, 1 cm. long, 5 mm. broad,
emarginate at the apex.

ASPARAGUS L. Asparagus

Herbs or often more or less woody vines, or erect shrubs, the stems, at least
in age, becoming much branched, the ultimate branchlets (cladodes) filiform and
terete or angulate, or sometimes leaflike, solitary or fasciculate in the axils of small
scales; flowers small, solitary or clustered at the bases of the cladodes, sometimes
short-racemose; perianth marcescent and persistent, the segments subequal,
distinct; stamens 6, hypogynous or affixed to the bases of the perianth segments,
the filaments filiform, the anthers ovate or oblong, dorsifixed, the cells introrsely
dehiscent; ovary sessile, 3-celled, the style stout or slender, short or elongate, the
3 stigmas short, recurved; ovules 2-many in each cell; fruit baccate, globose,
the seeds solitary or few, subglobose, usually black.

About 100 species, in temperate and warm regions of the Old
World; none native in America.

Cladodes leaf-like, ovate A. asparagoides.

Cladodes linear or terete.

Cladodes linear, flat A. Sprengeri.

Cladodes terete.

Plants erect, herbaceous A. officinalis.

Plants scandent, woody A. plumosus.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 69

Asparagus asparagoides (L.) W. F. Wight, Cent. Diet. 11: 845.
1913. Medeola asparagoides L. Sp. PI. 339. 1753. Myrsiphyllum
asparagoides Willd. Ges. Naturf. Freund. Berl. Mag. 2: 25. 1808.

Planted occasionally for ornament, chiefly in patios. Native of
South Africa.

A very slender and usually interlaced, small vine with lustrous, pale green
foliage; cladodes solitary (clustered in the other species), ovate, acute or acuminate,
1-2 cm. long; flowers greenish white; berries dark purple.

Known by the name "smilax" among florists of the United
States, where formerly, at least, the plant was much used for making
floral designs. The color and form of the foliage are well preserved
for a long time after the branches are cut.

Asparagus officinalis L. Sp. PI. 313. 1753. Espdrrago. As-
paragus.

Grown as a vegetable, and often planted in gardens for ornament.
Native of Europe.

Plants herbaceous, erect, with much branched rootstocks, the young stems
stout, simple, covered with large scales, later branching and often a meter tall
or more; cladodes filiform, 6-15 mm. long, clustered in the axils of minute scales;
flowers mostly solitary at the nodes of the branches, green, pendent on filiform
articulate pedicels; perianth campanulate, about 5 mm. long, with linear obtuse
segments; stamens shorter than the perianth; berries red, about 8 mm. in diameter.

Asparagus is hot a common vegetable of Central America, but
is seen more frequently in Guatemala than elsewhere, principally
at rather high elevations. Most of that in the Guatemala City
market is said to come from the vicinity of Amatitlan. It is moder-
ately large and of good appearance, and is sometimes blanched before
being cut. It is grown also about Coban, but the stalks eaten there
are all or mostly rather slender. Tinned asparagus from the United
States is sold in small quantities in the more expensive food shops.

Asparagus plumosus J. G. Baker, Journ. Linn. Soc. 14: 613.
1875. Velo de novia.

Grown commonly in gardens for ornament in most regions of
Guatemala. Native of South Africa.

Plants very slender and intricately branched and tangled, often climbing to
a height of 3 meters or more, deep green, woody; stems green; cladodes numerous,
8-20 in a cluster, mostly 5-7 mm. long, terete; leaf scales on the main branches
developing into short woody spines; flowers borne at the ends of the branchlets,
white, the perianth lobes spreading, obovate; berry 1-3-seeded, purple-black.

70 FIELDIANA: BOTANY, VOLUME 24

A rather common ornamental plant in gardens, in both hot and
cool regions.

Asparagus Sprengeri Regel, Act. Hort. Petrop. 11: 302. 1890.
Planted commonly in gardens for ornament. Native of Natal.

A woody vine with tuberous roots; stems often 3 meters long or more, rather
stout, the branchlets angled, pale; cladodes in clusters of 3-8, linear, flat, 12-35
mm. long, pointed; leaf scales on the main stems with a slender brown brittle
spine; flowers in open racemes, pale pink, fragrant; berries 1-3-seeded, bright
red, 1 cm. or less in diameter.

The plant is especially handsome when covered with the showy
bright-colored berries.

BEAUCARNEA Lemaire
Reference: Trelease, Proc. Amer. Phil. Soc. 50: 437-441. 1911.

Trees with clean trunks and few branches, the trunk usually enlarged at the
base, the branches densely leafy at the ends; leaves elongate-linear, coriaceous,
entire or minutely serrulate, dilated at the base; flowers small, dioecious, in large,
much branched panicles, short-pedicellate and fasciculate in the axils of bracts,
the pedicels articulate below the flowers; perianth campanulate, the segments
distinct, acute, entire, subeqUal, spreading in anthesis; stamens 6, attached to
the base of the segments, the filaments filiform, the anthers ovate or oblong,
dorsifixed; ovary sessile, 1-celled; fruit dry, trigonous, 3- winged, neither lobate nor
inflated, irregularly rupturing at maturity; seeds 3-sulcate or 3-lobate.

About nine species are known, in Guatemala and Mexico. By
some authors the genus has been united with Nolina, in which the
ovary is 3-celled and the fruit is not winged.

Nerves of the leaves minutely papillose, the leaves rough to the touch.

B. guatemalensis.
Nerves of the leaves not papillose, the leaves smooth to the touch.

Leaves up to 3 cm. wide in the upper portion, about 1-1.5 cm. wide above the

basal constriction B. Ameliae.

Leaves up to 2 cm. wide in the upper portion, 7 mm. wide above the basal
constriction B. petenensis.

Beaucarnea Ameliae Lundell, Bull. Torrey Club 66: 585. /. 1.
1939. Corcho; Chicu (Huehuetenango).

Dry or wet, rocky, mountain slopes, 600-1,200 meters; Pete*n;
Alta Verapaz (Lanquin); Huehuetenango. Yucatan; central British
Honduras.

A shrub or tree 3-8 meters high, the trunk 20-45 cm. in diameter, thickened
at the base; branches few, erect; leaves clustered at the ends of the branches,
usually spreading or pendent, flat, the enlarged base 2.5-5 cm. wide, the blades

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 71

mostly 85-105 cm. long, tapering to a long setiform apex, the margins minutely
serrulate with appressed teeth; staminate inflorescence as much as 75 cm. long;
flowers in fascicles of 2-3, pale yellow or almost white; pedicels 5-7 mm. long;
seeds shallowly lobate.

The Maya names are reported from Yucatan as "tsipil" and
"chit." It is doubtful whether this species is distinct from B.
petenensis.

Beaucarnea guatemalensis Rose, Contr. U. S. Nat. Herb. 10:
88. /. 1. 1906. Izote real; Izote de montana (fide Aguilar).

Generally on dry rocky hillsides, 700-1,600 meters; Baja Verapaz
(type from Sierra de las Minas opposite El Rancho, probably on
the road to Coban, W. A. Kellerman 4320); Jalapa (Volcan de
Jumay); cultivated in the parks and gardens of Guatemala City;
endemic.

A low tree with a tall trunk and few erect branches; leaves rather thin, a meter
long or less, 2-3 cm. wide, the edges smooth or roughened, the faces rough to the
touch; inflorescence a large ovoid panicle, much branched, the branches 30 cm.
long or less; perianth segments about 3 mm. long; fruit elliptic-obovate, 15-18
mm. long and 13-15 mm. broad, emarginate at the base and apex; seeds 5 mm.
in diameter, irregularly trilobate, smooth.

The species is cultivated in Guatemala in the Jardin Botanico,
Finca La Aurora, and elsewhere.

Beaucarnea petenensis Lundell, Bull. Torrey Club 66: 586.
1939. Dracaena petenensis Lundell, Journ. Wash. Acad. Sci. 25:
230. 1935. Izote (Chiquimula).

Pete*n (type from Monte Hiltun, a grove in limestone valley forest,
Lundell 3271); Chiquimula (Montana Nube, southeast of Conception
de las Minas, 1,500-1,700 meters, on open wind-swept slopes covered
with small shrubs); endemic.

A tree 6-12 meters tall, the trunk 20-30 cm. in diameter, its base bulbous-
thickened, the branches few; leaves densely crowded at the ends of the branches,
grass-green, soft and flexible, pendent, flat, the enlarged base 3.5 cm. wide or less,
the upper portion of the blade as much as 2 cm. wide, tapering to a long setiform
tip, as much as 140 cm. long, the margins minutely serrulate with appressed teeth;
inflorescence as much as 30 cm. long and probably much larger, rather dense, the
branches suberect or strongly ascending, the lowest subtended by long thin bracts.

Plants that probably belong to this species are occasional in
cultivation in the city of Quezaltenango. The source of these
cultivated plants is unknown, but they may well have been brought
from northern Quiche* or Alta Verapaz, by some of the Indians who
bring live birds and mammals as well as cacti and orchids for sale.

72 FIELDIANA: BOTANY, VOLUME 24

GALOCHORTUS Pursh

Reference: Marion Ownbey, A monograph of the genus Calo-
chortus, Ann. Mo. Bot. Gard. 27: 371-561. 1940.

Plants glabrous, arising from tunicated bulbs with membranaceous or fibrous-
reticulate coats; stems scapiform or leafy, often branched; leaves mostly linear,
the solitary basal ones often conspicuous, the cauline ones reduced upward; inflores-
cences monochasial or subumbellate, the bracts usually equaling the pedicels in
number and opposite them; flowers small or rather large, erect or nutant, globose
to broadly campanulate; outer perianth segments finally valvate in bud, more or
less sepaloid, obtuse to attenuate, usually naked; inner perianth segments con-
volute in bud, usually more or less barbate on the inner face and with a glandular
spot near the base; stamens 6, in 2 series, the anthers oblong to linear, obtuse or
apiculate, the base prolonged below the attachment of the filament as a tubular
sheath; filaments subulate, dilated at the base; ovary triangular or 3- winged,
abruptly contracted at the apex or attenuate to a persistent 3-fid stigma; ovules
in 2 rows in each cell; capsule orbicular to linear, 3-angulate or 3-winged, erect
or nutant, septicidally dehiscent; seeds irregular or compressed, the testa usually
hexagonal-reticulate.

Species about 55, all in western North America. The genus
reaches its southern limit of distribution in Guatemala. In Cali-
fornia the plants are well known by the name "mariposa" (Spanish,
"butterfly") or "mariposa lily," and some of them have been intro-
duced into cultivation because of their handsome flowers.

Flowers erect; petals deep purple; sepals with a glandular spot near the base.

C. Ghiesbreghtii.

Flowers nutant; petals white or whitish; sepals with a median glandular spot
within C. fuscus.

Calochortus fuscus Schult. f. Bijdr. Nat. Wet. 4: 131. 1829.
Jekelen.

At 1,400-1,600 meters; Huehuetenango (Sierra de los Cuchu-
matanes, between Las Palmas and Chacula, Steyermark 51760).
Widely distributed in Mexico.

Bulb ovoid, with fibrous-reticulate coats; stems slender, erect, 60 cm. high or
less, sparsely branched, bulbiferous in the axils of the upper leaves and bracts;
basal leaf linear-attenuate, not exceeding the stem; cauline leaves successively
shorter upward, the lower ones linear, 5 mm. wide, the upper ones linear-lanceolate
and amplexicaul; flowers usually 2, campanulate, nutant; sepals shorter than the
petals, obtuse or acute, with an oblong median spot bordered above by a laciniate
membrane, dull lavender outside, creamy white within; petals narrowly elliptic
to cuneate-obovate, about 15 mm. long, acute or obtuse, ciliate distally, sparsely
barbate above the gland, white outside and inside, the gland depressed, oblong,
bordered above with a laciniate membrane; anthers yellow, about 5 mm. long,
shorter than the filaments; capsule linear-oblong, acute at each end, erect.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 73

Calochortus Ghiesbreghtii Wats. Proc. Amer. Acad. 14: 268.
1879.

Mountain slopes, usually on limestone, about 1,400-2,000 meters;
Huehuetenango (Sierra de los Cuchumatanes: Trinidad; Cumbre
Papal, between Cuilco and Ixmoqui). Southern Mexico.

Bulb small, ovoid, with thick fibrous-reticulate coats; stems slender, erect,
often branched, rarely bulbiferous, about 60 cm. high or lower; basal leaf linear,
attenuate, almost equaling the stem, the cauline leaves linear-attenuate; flowers
2 or more, erect, broadly campanulate, the pedicels often greatly elongate; sepals
about equaling the petals, elliptic, acute or obtuse, with a circular glandular spot
near the base, dull purple outside, purple within; petals obovate, cuneate at the
base, acute or obtuse, sparsely or densely barbate with short hairs about the gland,
deep dull purple; gland not depressed, naked, bordered above by a deeply laciniate
membrane; anthers apiculate, shorter than the filaments; capsule linear-oblong,
3-angulate, erect.

DRACAENA L.

Trees or shrubs with a thick woody caudex; leaves coriaceous, closely parallel-
veined, usually crowded at the ends of the branches; flowers usually densely
fasciculate along the branches of a small or large panicle, the bracts scarious,
large or small; perianth funnelform or narrowly campanulate, the tube short or
somewhat elongate, the lobes narrow, subequal, longer or shorter than the tube;
stamens 6, affixed to the tube at the base of the lobes, the filaments filiform or
somewhat complanate, the anthers oblong, dorsifixed at the middle, versatile,
the cells introrsely dehiscent; ovary sessile or subsessile, 3-celled, the style fili-
form, the stigma capitate; ovules solitary in the cell, erect; fruit baccate, globose,
smooth or shallowly sulcate; seeds usually solitary in the fruit, globose.

About 35 species, all except the following native of the warmer
parts of the Old World.

Dracaena americana Bonn. Smith, Trees & Shrubs 1: 207.
pi. 98. 1905. Izote de montana; Cana de arco; Cukil (Kekchi);
Cerbatana (Pete"n); Halal, Ikaax (Pete*n, Maya).

In dense wet forest or in thickets of the Atlantic lowlands,
ascending from sea level to about 1,350 meters; Pete"n (Cerro Ceibal) ;
Alta Verapaz; Izabal. Tabasco to British Honduras and Costa Rica.

A tree, sometimes 12 meters high but usually lower and often only a shrub,
the trunk often 30 cm. in diameter, the bark grayish and somewhat exfoliating,
the branches few, the branchlets marked by the oblique scars of fallen leaves;
leaves linear-ensiform, 20-35 cm. long, 1-2.5 cm. wide, somewhat dilated and
clasping at the base, attenuate to the apex, bright green, rather soft; panicles
ovoid, dense and many-flowered, about 20-30 cm. long, the branches leafy-bracted ;
pedicels 3-6 mm. long, articulate above the middle; perianth white or creamy
white, about 7 mm. long; berries oval or globose, yellowish green, 2 cm. or less in
diameter, containing 1-3 seeds; seeds subglobose, 10-12 mm. in diameter. (Fig. 16.)

FIG. 16. Dracaena americana. A. Habit of uppermost portion of plant i
anthesis; X 2 A- B. Stamen; X 4. C. Portion of inflorescence; X 2. D. Pist
with ovary cut transversely; X 4. E. Fruit; X 1.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 75

Called "candlewood" in British Honduras, and "izote" in Hon-
duras. We have not found the tree abundant in Guatemala, but it
may well be plentiful in regions we have not visited. It is a hand-
some and ornamental plant, and has been introduced into cultiva-
tion, as at Quirigua Hospital, and in Finca La Aurora, Guatemala.

ECHEANDIA Ortega

Reference: C. A. Weatherby, Synopsis of the genus Echeandia,
Proc. Amer. Acad. 45: 387-394. 1910.

Plants with short rhizomes, the root fibers fasciculate, thick and fleshy, some-
times fusiform; leaves all or mostly radical, long and narrow, linear or linear-
lanceolate, membranaceous, distichous, narrowed at the base into long sheaths;
scapes slender, simple or sparsely branched; flowers fasciculate along the scape
or its branches, slender-pedicellate, white or yellow, the bracts small, scarious;
perianth marcescent, the segments distinct, rotate-spreading, equal or the inner
somewhat wider, 3-5-nerved; stamens 6, hypogynous, shorter than the perianth,
ithe filaments short, filiform, the anthers linear, basifixed, equaling or slightly
longer than the filaments, coherent about the style to form a cylinder, the cells
introrsely dehiscent; ovary sessile, 3-celled, the style filiform, slightly exserted
beyond the anthers, the stigma small, capitate; ovules numerous in the cell; capsule
ovoid or oblong, trigonous, loculicidally dehiscent; seeds numerous, angulate-
compressed, black.

About ten species, ranging from Mexico to Venezuela, only two
known from Central America. In general appearance as well as in
most details the plants are exactly like species of Anthericum, and
the genus can be determined only by examination of the anthers.
Since many Guatemalan specimens are in fruit only, the deter-
mination in some instances is questionable.

Capsules 6-9 mm. long; filaments smooth E. parviflora.

I Capsules mostly 12-18 mm. long; filaments roughened E. macrocarpa.

Echeandia macrocarpa Greenm. Proc. Amer. Acad. 39: 73.
1903.

Damp fields or oak forest in the mountains, sometimes in Juni-
perus forest, 1,400-1,800 meters; Jalapa; Guatemala; Sacatepe"quez;
Huehuetenango. Southern Mexico.

Leaves all or chiefly basal, much elongate, 6-15 mm. wide, glabrous; scape
glabrous, simple or with few branches, 40-60 cm. tall, bearing 1-2 bracts; pedicels
articulate below the middle, stout, in fruit becoming 1-1.5 cm. long; flowers yellow,
the perianth segments 15-17 mm. long, the inner ones ovate-lanceolate; filaments
equaling or slightly longer than the anthers; capsule oblong, 4-6 mm. broad.

76 FIELDI ANA: BOTANY, VOLUME 24

It is perhaps this species that was reported from Volcan de Fuego
by Hemsley as E. terniflora Cav.

Echeandia parviflora J. G. Baker, Bot. Jahrb. 8: 209. 1887.

Open ground, 900-1,600 meters; Guatemala (type from Amatitlan,
F. C. Lehmann 1647); Santa Rosa; material from Pete"n probably
belongs here, and possibly a poor collection from Zacapa; Huehue-
tenango. Southern Mexico.

Leaves linear, thin, 4-8 mm. wide, 20 cm. long or often longer, erect or some-
what spreading; scape scabrous or short-hirtellous below, simple or with a few
slender branches, as much as 80 cm. tall; pedicels rather short and stout, in fruit
6-8 mm. long or longer, articulate below the middle or near the base; perianth
creamy white, 5-6 mm. long or somewhat longer; anthers about 5 mm. long;
capsule oval or broadly oblong, usually 6-8 mm. long and 4-5 mm. broad.

HEMEROCALLIS L.

Tall glabrous herbs with fibrous roots, the leaves basal, linear; flowers large
and lily-like, orange or yellow, grouped at the ends of leafless scapes; perianth
funnelform, the lobes oblong or spatulate, much longer than the cylindric tube;
stamens 6, inserted at the summit of the perianth tube, shorter than the lobes,
declined, the filaments filiform, the anthers linear-oblong, the cells introrsely
dehiscent; ovary oblong, 3-celled, the ovules numerous in each cell; style slender,
declined, the stigma small, capitate; capsule oblong or ovoid, thick-walled, 3-
angulate, loculicidally 3-valvate.

About ten species, natives of Eurasia. Two of them are widely
cultivated in temperate regions.

Hemerocallis fulva L. Sp. PI. ed. 2. 462. 1762.

Occasionally cultivated for ornament at middle elevations; noted
at Huehuetenango, Finca El Zapote in Escuintla, Coban, Antigua,
and elsewhere; tending to escape along hedges and roadsides in the
mountains of Alta Verapaz. Native of Europe and Asia.

Leaves 8-12 mm. wide, tapering to an acute apex; scapes often a meter high
or even taller, bearing several short bracts above, the flowers 6-15, short-pedicel-
late, reddish orange, paniculate, 10-12 cm. long, opening for one day only; perianth
tube 2.5-3 cm. long, the lobes oblong, somewhat spreading.

Usually called "orange day-lily" in the United States, where
the plant has long been under cultivation in country gardens. Some
of the plants grown in Guatemala have double flowers. In Salvador
the plant is called "clavel de Italia."

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 77

KNIPHOFIA Moench

Large coarse glabrous plants with short rhizomes, the root fibers numerous,
thick and fleshy; leaves radical, long and linear, green; scape leafless, tall, simple,
the flowers showy, very densely racemose or spicate, yellow or red, subsessile or
short-pedicellate, deflexed; bracts small but rather conspicuous because of their
great number, scarious; perianth tubular, the segments 1-nerved, connate for most
of their length, the short free portion ovate or deltoid; stamens 6, hypogynous,
slightly shorter than the perianth or often exserted, the filaments filiform, the
anthers oblong or linear, dorsifixed, the cells introrsely dehiscent; ovary sessile,
3-celled, the style filiform, often exserted, the stigma small; ovules numerous in
each cell; capsule globose or globose-ovoid, coriaceous, loculicidally dehiscent;
seeds angulate, black.

About 25 species, in South Africa and Madagascar.

Kniphofia Uvaria (L.) Hook. Bot. Mag. pi. 4816. 1854. Aloe
Uvaria L. Sp. PI. 323. 1753. Espech (Quezaltenango) ; Candela
(San Marcos); Pompdn.

Abundantly naturalized in Quezaltenango and San Marcos, in
fence-rows and along roadsides, very plentiful and conspicuous in
many places, especially above Palestina and on the plains west and
north of Quezaltenango; noted also in Totonicapan and at Huehue-
tenango; planted for ornament in many gardens of the highlands.
Native of South Africa.

Leaves very numerous and usually forming dense clumps, recurved and often
lying upon the ground, pale green, linear, usually 1-1.5 cm. wide, 60-90 cm. long,
carinate, rough-margined, soft; racemes 15 cm. long or less, spikelike, very dense
and many-flowered; upper flowers bright red, the lower ones yellow, reflexed, the
cylindric perianth 2.5-3.5 cm. long; bracts scarious and pale, a third as long as the
flowers or shorter.

Known in the United States, where sometimes cultivated, as
"poker plant." When in flower, in the highlands of Guatemala in
January, and probably also in most of the other months, the plant
is a showy one because of its brilliant stalks of flowers. It was
noted in abundance only in Quezaltenango and San Marcos, where
it is conspicuous during the winter months because of its dense masses
of green leaves, there being but little green at that season of the
year at these high elevations. Evidently the foliage is not eaten
by stock.

LILIUM L. Lily

Large or small plants with bulbs, the bulbs usually composed of thick fleshy
scales; stems erect, leafy, simple or branched above; leaves various in shape,
sessile or petiolate, verticillate or scattered; flowers large and showy, solitary

78 FIELDIANA: BOTANY, VOLUME 24

at the apex of the stem or several or many and racemose or umbellate; perianth
deciduous, funnelform or subcampanulate, the segments distinct, more or less
unguiculate, the upper portion often recurved or revolute; stamens 6, hypogynous,
shorter than the perianth segments, the filaments filiform or somewhat complanate,
the anthers linear or oblong-linear, dorsifixed, versatile, the cells introrsely dehis-
cent; ovary sessile, narrow, 3-celled, the style elongate, usually clavate at the
apex; ovules numerous in the cell; capsule oblong, chartaceous or coriaceous,
loculicidally dehiscent; seeds numerous, compressed, brown or pale, often marginate.

About 100 species, in temperate regions of the northern hemi-
sphere. Numerous species are native in the United States, but no
native ones are found as far south as Central America. Only one
species is cultivated at all commonly in Central America, but a
few others may be found in private gardens of Guatemala.

Lilium longiflorum Thumb. Trans. Linn. Soc. 2: 333. 1794.
Azucena; Saquicotzij (Totonicapan, fide Tejada).

Grown generally for ornament, chiefly at middle and higher
elevations, but sometimes even in the lowlands. Native of China
and Japan.

Bulbs 5-10 cm. in diameter, subglobose or oblate, white or yellowish; stems
stout, a meter high or less, smooth, green; leaves numerous, spreading or the
upper suberect, linear-lanceolate, mostly 7-12 cm. long and 9-15 mm. wide;
flowers 1-10, pure white, fragrant, often tinged with green near the base, about
17 cm. long; anthers yellow.

One of the favorite garden flowers of Guatemala and of the moun-
tain regions generally of Central America, to be seen in gardens of
rich and poor. Large quantities of the flowers are grown for market
at Almolonga and Zunil (Quezaltenango), and presumably also not
far from Guatemala, since they usually are plentiful in the Guate-
mala markets. The flowers are used everywhere for making funeral
coronas or wreaths, and as decorations upon altars and in religious
processions, where thousands of them sometimes may be seen. The
lily does not thrive in the lowlands, but the flowers grown in the
highlands are carried down commonly by cargadores to the tierra
caliente for sale there.

Lilium tigrinum L. Lirio rojo. A native of Europe, rarely
grown in Guatemalan gardens. The large flowers are orange-red;
the leaves usually bear small dark-colored bulblets in their axils.
Known in the United States as "tiger lily."

MILL A Cavanilles

Plants arising from a small corm, the root fibers fasciculate, sometimes fleshy-
thickened; leaves few, radical, narrowly linear; scapes simple, naked, the flowers

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 79

i2-4 or rarely 1, forming a terminal umbel, long-pedicellate, large, white; involucral
: bracts 2-3, small and inconspicuous; perianth salverform, the tube cylindric,
'equal, little constricted at the mouth, the lobes rotate-spreading, slightly unequal;
stamens 6, inserted in the mouth of the tube, exserted, the filaments very short;
anthers oblong, erect, connivent about the style, the cells introrsely dehiscent;
: ovary sessile, elongate, 3-celled; style slightly longer than the anthers, the stigma
: little dilated, 3-lobate; ovules numerous in each cell; capsule oblong, obtuse,
membranaceous, loculicidally dehiscent, the valves septicidally parted; seeds
compressed, angulate, black.

The genus consists of a single species.

Milla biflora Cav. Icon. PI. 2: 76. pi 196. 1793. Jacinto de
monte; Chumimi.

At 800-1,700 meters; Huehuetenango; Quiche'. Mexico.

Corms about 1.5 cm. in diameter, brown, with a loose outer coat; leaves few
or numerous, often withering, 1 mm. wide or sometimes wider, usually shorter
than the scapes; scapes slender, generally 30 cm. high or less, the pedicels rather
stout, shorter than the scapes or frequently equaling them; perianth lobes 2-3 cm.
long, elliptic-oblong, obtuse-apiculate, white within, pale greenish outside, with a
median several-nerved stripe; anthers narrowly oblong, about 6 mm. long; capsule
about 1.5 cm. long.

A handsome and rather conspicuous plant, common at some places
in the Sierra de los Cuchumatanes of Huehuetenango.

NOTHOSCORDUM Kunth

Low plants with tunicated bulbs, in general appearance exactly like some
species of Attium, but without alliaceous odor; leaves radical, linear, flat; scape
slender, naked, the flowers in a terminal umbel, not articulate with the pedicels;
involucral bracts 2, connate at the base; perianth marcescent, persistent, the 6
segments connate at the base or to the middle, spreading or campanulate-con-
nivent, subequal, 1-nerved; stamens 6, affixed to the bases of the perianth segments,
shorter than the perianth, the filaments more or less dilated but subulate and entire
at the apex; anthers oblong, dorsifixed, the cells introrsely dehiscent; ovary sessile,
3-celled, the style filiform, the stigma small; ovules usually 6-12 in each cell;
capsule membranaceous, 3-lobate, loculicidally dehiscent; seeds angulate-com-
pressed or almost flat, black.

About 10 species, one Chinese, the others American. Only one
is found in Central America.

Nothoscordum fragrans (Vent.) Kunth, Enum. PI. 4: 461.
1843. Allium fragrans Vent. Descr. PI. Nouv. Jard. Gels 26. 1800.
Cebollin (fide Aguilar).

A weed in waste or cultivated ground, 1,250-1,500 meters; Alta
Verapaz (Coban and vicinity); Guatemala (Guatemala). Mexico
and southeastern United States; Costa Rica; West Indies.

80 FIELDIANA: BOTANY, VOLUME 24

Plants onion-like in appearance, with small white bulbs; leaves few or numer-
ous, erect or weak and recurved, usually as long as the scape, mostly 2-8 mm.
wide, succulent and soft; scape slender, 30-60 cm. tall, glabrous, bearing at the
apex an umbel of few or numerous greenish white flowers, the slender pedicels
unequal, 2-3 cm. long; perianth 8-10 mm. long, the segments obtuse; capsule
5-6 mm. high.

Apparently this plant is introduced in Guatemala, for it is found
in few localities, and not far from dwellings, just as is the case in
Costa Rica. Except for its lack of an onion odor, there is little to
distinguish the genus from Allium.

Phormium tenax Forest., New Zealand flax (known locally as
"lirio de espada"), native of New Zealand, is planted rarely for
ornament in Guatemala. It is a coarse acaulescent plant with sword-
like leaves 5-12 cm. wide. The form in cultivation here is P. tenax
var. variegatum Hort., in which the leaves are bordered or striped
with creamy white. In its native country the plant is of great
economic importance because of the very strong fiber obtainable
from its leaves.

Ruscus aculeatus L., native of the Mediterranean region, is seen
very rarely in Guatemala as an ornamental plant, usually in pots.
It is much grown in southern France and elsewhere for ornamental
purposes, the dried branches and leaves being dyed and used to
make funeral wreaths and decorative articles.

SANSEVIERIA Thunb.

Perennial herbs with creeping rootstocks; leaves flat or terete, coriaceous,
thick and tough, cespitose, often mottled with light and dark green; scapes simple,
much shorter than the leaves, the flowers greenish white, racemose or paniculate;
perianth segments united below to form a tube, the lobes narrow, spreading;
stamens 6, inserted at the bases of the perianth segments, the filaments filiform,
the anthers oblong or linear; ovary 3-celled, the ovules solitary in each cell; seeds
1-3, subglobose, fleshy.

About 50 species, in Africa and Asia. The tough fiber of the
leaves of some species is utilized commercially.

Sansevieria guineensis (L.) Willd. Sp. PI. 2: 159. 1799.
Aletris hyacinthoides var. guineensis L. Sp. PI. ed. 2. 456. 1762.
Curarina; Oreja de burro; Quina (Escuintla).

Planted commonly for ornament at middle and low elevations,
often grown for hedges; thoroughly naturalized in many parts of

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 81

the lowlands, at least in hedges and thickets near dwellings, or on
the borders of fields. Native of tropical Africa.

Leaves linear-lanceolate, concave, 50-140 cm. long, 5-9 cm. wide, erect,
narrowed at each end, dark green, mottled with white or gray; scapes usually
shorter than the leaves and often almost concealed by them, the white flowers
forming a narrow panicle, the pedicels 5-8 mm. long; perianth segments linear
or linear-spatulate, the whole flower 1.5 cm. long or less, the recurved lobes about
equaling the tube.

The fiber of this species has been utilized in some regions where
the plant is native or cultivated. The plant is well known in cultiva-
tion in the United States, particularly as a pot plant, since it stands
the dry heat of apartments in winter time. During the past few
years the name "mother-in-law plant" a name of no apparent
application has come into rather general use for it among United
States florists. It would be interesting to know how the name
"curarina" came to be applied to this plant, for there is no relation-
ship between Sansevieria and the genus Strychnos, which supplies,
in part, the drug curare. The local name is well established, how-
ever, and everywhere in Guatemala even the children know the
plant as "curarina." It is much used in domestic medicine, being,
apparently, one of the most esteemed of local plants. It is even
reported to have the properties of quinine, for which there is probably
little or no basis. In Salvador Sansevieria is known by the names
"espada de Judas" and "espada del diablo."

SCHOENOCAULON Gray

Reference: Robert R. Brinker, Monograph of Schoenocaulon, Ann.
Mo. Bot. Gard. 29: 287-314. 1942.

Bulbs usually narrow, fibrose-tunicate; leaves radical, elongate-linear; scapes
tall, simple, leafless, the flowers small, subsessile, forming a long, usually dense
spike, the filaments and perianth long-persistent, the bracts small; perianth seg-
ments distinct, subequal, narrowly lanceolate or linear, erect-spreading, bearing
within at the base a pit or nectary; stamens 6, hypogynous, longer than the
perianth, the filaments thick-filiform, erect; anthers small, subglobose, the cells
confluent, explanate and orbicular-peltate after emission of the pollen; ovary
ovoid, slightly contracted at the base, free from the perianth, shallowly 3-lobate
at the apex, 3-celled, the lobes bearing short, introrsely stigmatose styles; ovules
4-8 in each cell ; capsule ovoid or oblong, acuminate, separating septicidally into
3 carpels; seeds oblong, often curved, angulate, sometimes acuminate, black or
fuscous, the testa thick, lax; endosperm carnose.

About five species, in Texas, Mexico, and Central America, one
extending to Venezuela.

82 FIELDI AN A: BOTANY, VOLUME 24

Schoenocaulon officinale (Schlecht. & Cham.) Gray ex Benth.
PL Hartweg. 29. 1939. Veratrum officinale Schlecht. & Cham.
Linnaea 6: 45. 1831. Sabadilla officinalis Standl. in Standl. & Cald.
Lista PI. Salv. 49. 1925. Cebadilla; Cebollajo; Chaguitera (fide
Aguilar) ; Boom (Huehuetenango).

Brushy or grassy slopes, usually in pine or oak forest, mostly in
rather dry and exposed places, sometimes on moist or wet slopes,
chiefly at 500-2,000 meters; Zacapa; Chiquimula; Jutiapa; Santa
Rosa; Guatemala; Chimaltenango; Huehuetenango; reported from
Baja Verapaz. Southern Mexico (Mexico to Veracruz and Chiapas);
Salvador; Honduras; reported from Costa Rica, but perhaps only
in cultivation; Venezuela.

Bulbs long and narrow, densely covered with very numerous, long, loosely
interlaced, black fibers; leaves rather hard and dry, a meter long or less, 1 cm. wide
or narrower, closely and conspicuously nerved, glabrous; scape usually about a
meter tall, stout, terete, the spikes often 30 cm. long or more, very dense, in
flower 1.5 cm. thick, the flowers cream-colored or greenish white, opening from
below upward; capsules pale, 11-12 mm. long, the carpels rostrate. (Fig. 17.)

The plant has been referred also to the genera Asagraea and
Helonias, and there are numerous synonyms. The seeds are
4-6 mm. long, wrinkled, black or dark brown outside, whitish
within, hard, inodorous, and have an exceedingly acrid, burning
taste. They contain veratrin and other alkaloids, and are poisonous.
They formerly constituted an official drug of the British Pharma-
copoeia. They have emetic-cathartic properties, operating some-
times with great violence, and overdoses produce death. They were
known in Europe as early as 1752, and were formerly used to some
extent to expel tapeworms. The seeds are the chief ingredients of
pulvis Capucinorum, sometimes used in Europe for destruction of
vermin in the hair. During the first World War there was a great
demand in central Europe for the seeds, for use in the armies,
especially the German ones. Large amounts of seed are exported
from Venezuela, where the plant is cultivated. They are said to
be exported also from Mexico, and even from Guatemala, but
shipment from the latter country seems doubtful, since persons in
the country who were consulted knew nothing of their export, and
the people seem to have little or no knowledge of the plant or
its properties. Although widely scattered in Guatemala, the plant
is not at all plentiful, and we have not found it in quantity
anywhere. It flowers and matures seeds during the invierno, and
the leaves wither soon after the rains cease.

FIG. 17. Schoenocaulon offlcinale. A. Habit of plant with detached in-
florescence at right; X M- B. Flower viewed from above, with pistil detached;
X 4; C. Flower viewed from the side, with pistil in natural position; X 4. D.
Capsule, dehiscing; X 2. E. Longitudinal section through portion of pistil; X 2.

84 FIELDIANA: BOTANY, VOLUME 24

SMILACINA Desf.

Reference: Ralph W. Emons, A revision of the Central American
species of Smilacina, Ann. Mo. Bot. Card. 32: 395-411. 1945.

Low herbs, sometimes epiphytic, with short or elongate, thick or slender
rhizomes, the stems erect, simple, leafy above; leaves alternate, almost sessile,
chiefly ovate or lanceolate; inflorescence terminal, racemose or paniculate, the
flowers small, pedicellate, solitary or in fascicles of 2-3 within the bracts; perianth
finally deciduous, the segments distinct or nearly so, subequal, spreading; stamens
6, hypogynous or affixed to the very base of the perianth, the filaments short and
broad or elongate and filiform, the anthers ovate, dorsifixed, the cells introrsely
dehiscent; ovary sessile, subglobose, 3-celled; style columnar, the stigma small or
sometimes 3-lobate; ovules 2 in each cell; fruit baccate, globose; seeds few, often
solitary, subglobose.

About 25 species in North America and Asia. In America the
genus extends south to Panama, where one additional species, S. gigas
Woodson, a giant terrestrial plant, by Emons considered synony-
mous with S. paniculata, is known.

Stems strongly bulbous-thickened at the base; plants epiphytic; inflorescence
paniculate or racemose, very dense and many-flowered in anthesis, the pedicels

straight, erect or ascending; perianth bright rose S. amoena.

Stems not bulbous-thickened at the base, or only slightly so; plants commonly
terrestrial; inflorescence paniculate or racemose, not very dense even at first,
sometimes few-flowered, the pedicels often curved; perianth usually white
or creamy white, sometimes pale pink.
Inflorescence paniculate.

Perianth 3-4 mm. long; branches of the panicle usually many-flowered, the

pedicels mostly very slender S. paniculata.

Perianth 6-7 mm. long; branches of the panicle few-flowered, the pedicels

very stout S. crassipes.

Inflorescence racemose.

Inflorescence few-flowered, the rachis straight or nearly so S. scilloidea.

Inflorescence many-flowered, the rachis zigzag or strongly flexuous . S. flexuosa.

Smilacina amoena Wendl. in Otto & Dietr. Allg. Gart. Zeit.
18: 137. 1850. Trovaria Salvini Baker, Journ. Linn. Soc. Bot. 14:
567. 1875 (type from Volcan de Atitlan, Solola, Salviri). S. Salvini
Hemsl. Biol. Centr. Amer. Bot. 3: 368. 1884. Convallaria Salvini
Druce, Bot. Exch. Club Brit. Isl. 3: 408. 1914. Vagnera Salvini
Standl. Journ. Wash. Acad. Sci. 15: 457. 1925. Ramillete de Mag-
dalena (fide Aguilar).

On mossy tree trunks in dense forest, 2,200-3,800 meters; El
Progreso; Chimaltenango; Solola; Quiche"; Quezaltenango; San
Marcos. Southern Mexico; Costa Rica.

Stems solitary or cespitose on tree trunks, 30-50 cm. tall or even larger, stout,
usually much thickened and bulblike at the base, arising from thick and often

FIG. 18. Smilacina flexuosa. A. Habit of plant; X
C. Flower; X 4.

- B. Fruit; X 3.

86 FIELDIANA: BOTANY, VOLUME 24

elongate rhizomes, bearing few or numerous leaves; leaves sessile or nearly so,
broadly ovate to elliptic-lanceolate, 5-12 cm. long, commonly 3-4 cm. wide, acute
to long-acuminate, glabrous; inflorescence at first very dense and almost head-
like, in age more open, the pedicels and short branches straight, erect or strongly
ascending, the pedicels mostly 1-1.5 cm. long; perianth deep rose or rose-red, about
5 mm. long, the segments broadly rounded or somewhat emarginate at the apex.

This plant is an exceptionally beautiful one on account of its
large clusters of brilliantly colored flowers. Because of its epiphytic
habit, frequently it is mistaken for an orchid by the local people or
by strangers, and sometimes is offered for sale as one. Emons refers
most of the Guatemalan material to var. Salvini (Baker) Emons,
but the varietal characters are not obvious.

Smilacina crassipes Standl. & Steyerm. Field Mus. Bot. 23:
214. 1947.

On rocks or sometimes epiphytic on tree trunks, 1,300-1,900
meters; endemic; Huehuetenango (type from rocky slopes above
La Libertad, on Cerro Pueblo Viejo, Steyermark 51003); San Marcos
(Volcan de Tajumulco near Finca El Porvenir).

Plants erect or pendent, sometimes a meter high, the stems stout, almost
1 cm. thick below, somewhat thickened and bulb-like at the base; leaves numerous,
membranaceous, lanceolate to oblong-lanceolate or oblong-elliptic, mostly 17-20
cm. long and 4.5-5.5 cm. wide, sometimes as much as 8.5 cm. wide, attenuate-
acuminate, obtuse or rounded at the base and contracted into a stout petiole 7
mm. long or shorter; inflorescence narrowly paniculate, 10-15 cm. long, 5-6 cm.
wide, the axis very stout and angulate, the lateral branches few, divaricate at
almost a right angle, very stout, angulate, mostly 2-4-flowered; pedicels 5 mm.
long or usually shorter, thick, stout, acutely angulate when dry; perianth pale
yellow, campanulate, the segments thick, oval or oblong-oval, 6-7 mm. long,
rounded at the apex, suberect in anthesis, more or less persistent; anthers about
1.3 mm. long, the stamens somewhat shorter than the perianth; berries blood-red,
about 8 mm. in diameter.

Smilacina flexuosa Bertol. Fl. Guat. 411. pi. 39. 1840. S.
Bertolonii Kunth, Enum. PI. 5: 151. 1850. Convallaria flexuosa
Druce, Bot. Exch. Club Brit. Isl. 3: 408. 1914. Vagnera flexuosa
Standl. Journ. Wash. Acad. Sci. 15: 457. 1925. Ramillete de la
Virgen (fide Aguilar); Diente de perro.

Dense, damp or wet forest, terrestrial or rarely on tree trunks,
1,300-3,000 meters; type collected in the mountains of Guatemala
by Velasquez, the locality not specified; Alta Verapaz; Baja Verapaz;
Zacapa; Guatemala; Sacatepe"quez; Chimaltenango; Solola; Suchite-
pequez; Quiche"; Huehuetenango; San Marcos; reported from Santa
Rosa. Southern Mexico; El Salvador; Honduras; Costa Rica.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 87

Stems arising from thick elongate rhizomes, not or scarcely thickened at the
base, bearing few or numerous leaves, commonly 40-100 cm. tall; leaves narrowly
oblong-lanceolate to ovate-elliptic, mostly 11-16 cm. long, bright green, glabrous;
inflorescence simply racemose, short and rather few-flowered or sometimes much
elongate and many-flowered, often recurved, the slender rachis often zigzag,
always strongly flexuous, the slender, more or less curved, rather stiff pedicels
1-2 cm. long, the whole inflorescence in fruit often recurved or pendent by the
weight of the fruits; perianth pink, white, or dull purplish, 6-7 mm. long; fruit
bright or dull red, subglobose or shallowly trilobate, about 1 cm. broad, usually
3-seeded. (Fig. 18.)

A variety erubescens Emons with pink flowers is known from
several departments in Guatemala, typical S. flexuosa being main-
tained by Emons for the more common white-flowered form.

Smilacina paniculata Mart. & Gal. Bull. Acad. Brux. 9, pt. 2:
388. 1842.

Dense or wet forest, 1,300-3,300 meters, usually terrestrial,
rarely epiphytic; Alta Verapaz; Baja Verapaz; El Progreso; Zacapa;
Suchitepequez; Quiche"; Huehuetenango; San Marcos. Southern
Mexico; El Salvador; Costa Rica; Panama.

Plants glabrous, the stems a meter high or less, rarely almost 2 meters tall,
not thickened at the base, bearing numerous leaves; leaves short-petiolate, narrowly
lanceolate to ovate-elliptic, 8-20 cm. long, gradually or abruptly acuminate;
panicles usually pyramidal, large and many-flowered, lax and much branched, the
branches dull red or lilac, straight or nearly so, the pedicels 1 cm. long or less,
straight, often conspicuously thickened; perianth white or pinkish, 3-4 mm. long;
fruit about 8 mm. in diameter, at first green spotted with dark purple, turning
i blood-red.

Smilacina scilloidea Mart. & Gal. Bull. Acad. Brux. 9, pt. 2:
1388. 1842.

Dense or open pine, Juniperus, or Abies forest of the central
.and western mountains, 2,700-3,800 meters; Chimaltenango (Volcan
de Acatenango); Huehuetenango; Quezaltenango; San Marcos.
Southern Mexico; Honduras.

Plants small, with long slender repent rhizomes, the stems mostly 15-30 cm.
tall, flexuous, more slender than in other species, with usually few, distant leaves;
leaves bright green, thin, glabrous, mostly 5-10 cm. long, oblong-lanceolate to
'narrowly lanceolate, acute to long-attenuate; racemes small and with very few
flowers, the rachis and pedicels straight or nearly so, the pedicels 5 mm. long or less;
perianth white to deep purplish, 3-4 mm. long; fruit mostly 8 mm. or less in diam-
eter, bright red.

Part of the Guatemalan material is referred by Emons to var.
rosea Emons, which probably is a pink- or purple-flowered form
of the species.

88 FIELDI ANA: BOTANY, VOLUME 24

TAETSIA Medic.

Tall, somewhat shrubby plants, the caudex ligneous, often stoloniferous;
leaves crowded at or near the apex of the stem, petiolate or subsessile, lanceolate
or narrower; flowers small, paniculate, short-pedicellate, solitary or few within
each bract, the bracts small and scarious; perianth cylindric or narrowly cam-
panulate, with a short tube, the lobes oblong, erect or somewhat spreading, equal
or the inner slightly longer; stamens 6, inserted on the tube at the base of the
lobes, about equaling the perianth, the filaments filiform or complanate; anthers
narrowly oblong, dorsifixed, the cells introrsely dehiscent; ovary sessile, 3-celled,
the style filiform, the stigma capitate or shallowly 3-lobate; ovules 4-16 in each
cell; fruit globose or trilobate, fleshy, 3-celled, indehiscent or sometimes loculi-
cidally dehiscent; seeds in each cell numerous or by abortion 1, obovoid or some-
what compressed, black and lustrous.

About ten species, chiefly in the East Indies, Pacific islands, and
Australia.

Taetsia fruticosa (L.) Merrill, Interpret. Herb. Amboin. 137.
1917. Convallaria fruticosa L. in Stickm. Herb. Amboin. 16. 1754.
Asparagus terminalis L. Sp. PI. ed. 2. 450. 1762. Cordyline terminalis
Kunth, Abh. Acad. Berlin 30. 1820. Liston, Croton listado (fide
Aguilar).

Planted everywhere in the lowlands, in gardens and in roadside
hedges, escaping to thickets and naturalized in many localities;
also in gardens of the central and other mountain regions, but not
so common or altogether absent above 1,500 meters. Probably a
native of Malaya.

Plants 1-3 meters high or even taller, simple, densely leafy, the stems rather
slender; leaves elliptic-lanceolate to linear-lanceolate, 30-70 cm. long, 10-15 cm.
wide, acute to long-attenuate, narrowed to the base, green or often deep red-purple,
often green with red margins; petioles often more than half as long as the blades;
panicles terminal, large, dense or usually open; pedicels very short or none;
perianth white, purplish, or reddish, 10-12 mm. long, the segments linear-oblong;
ovules 6-10 in each cell; fruit a large, red or purple berry.

One of the most common ornamental plants throughout the low-
lands of Central America. The form most cultivated in Guatemala
is var. ferrea (Baker) Standl., in which the leaves are colored with
dark red or purple.

YUCCA L.

Large coarse plants, with a woody caudex, acaulescent or often tall and tree-
like, simple or branched; leaves crowded at the apex of the caudex or its branches,
linear-lanceolate, thick and rigid or rarely thin and soft, usually spinose at the
apex, the margins entire or separating into coarse fibers; flowers white or cream,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 89

n very large, terminal panicles, pedicellate, pendulous; perianth long-persistent,
;he segments distinct or nearly so, lance-ovate, rather fleshy, more or less con-
nivent to form a subglobose flower; stamens 6, hypogynous, erect, much shorter
;han the perianth, the filaments rather stout; anthers small, sessile at the apex of
;he filament, introrsely dehiscent; ovary sessile, 3-flowered, the cells imperfectly
2-celled, the style columnar, short, divided at the apex into 3 lobes that are stig-
matose within; ovules numerous in each cell; fruit fleshy and pulpy or spongious
and indehiscent, often dry and loculicidally dehiscent; seeds strongly compressed,
black.

About 25 species in southern and especially southwestern United
States, Mexico, and West Indies. Only the following one is known
in Central America, where it probably is not native.

Yucca elephantipes Regel, Gartenflora 8: 35. 1859. Y. guate-
malensis J. G. Baker, Ref. Bot. 5: pi. 313. 1872 (type collected in
Guatemala by Ehrenberg). Izote; Palmera (Pete"n, fide Lundell);
Cukil, Quiil, Co'quil (Alta Verapaz, Kekchi); Pasquiy (Chimalte-
nango, fide Tejada); Pasqui (Totonicapdn, fide Tejada).

Common in cultivation or in hedges and thickets throughout
the lowlands and ascending commonly in the mountains to middle
(elevations (about 1,500 meters); often seen up to 2,700 meters or
perhaps even higher; Pete*n; Alta Verapaz; Jalapa; Escuintla;
Huehuetenango; San Marcos; probably in all the departments.
'British Honduras; generally distributed in cultivation in Central
j America, southward at least to Costa Rica, also in Mexico.

Plants usually treelike, with a thick columnar crown and a few short thick
branches, the trunk and lower parts of the branches naked, the bark rather rough;

i leaves dagger-like, stiff and hard, a meter long or usually much shorter, rough-
margined, very sharp-pointed; flowers white or creamy white, campanulate, about
4 cm. long; fruit fleshy, oblong-ovoid, with white or yellowish flesh and a papery

; core. (Fig. 19.)

Known in Costa Rica as "itabo," an Indian name, but almost
'everywhere else in Central America as "izote," a word of Nahuatl
derivation. The Maya name reported from Yucatan is "tuc." In
Guatemala the plant is commemorated geographically by the caserio
of El Hizote in the Department of Santa Rosa (the "H" is quite
superfluous). The tree is an ornamental one, and partly for that
reason often is planted about houses, especially for tall hedges. In
some regions, as about Coatepeque, it is planted thickly on steep
high roadside banks to prevent erosion. It is seen frequently, too,
in cemeteries. While most of the plants are perhaps 3-4 meters
high, often they are much taller and possibly as much as 10 meters
high. Many very tall and often much branched individuals are

FIG. 19. Yucca ekphantipes. A. Habit; X Vso. B. Perianth removed to
show stamens and pistil; X 3. C. Fruit; X Ji. D. Flower, with one of perianth
segments removed; X 1. E. Portion of inflorescence; X 1 A. F. Leaf; X Ji-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 91

seen about Coban, Retalhuleu, and Patulul. On the Volcan de
Agua izotes are planted above the highest pea fields, at perhaps
2,700 meters. Although the large panicles are a handsome sight
when the flowers are open, it is but rarely that they are seen well
developed, for almost all the panicles are cut when the flowers begin
to open, and the succulent flowers are used for food. They have a
slightly bitter but quite agreeable flavor, and constitute a palatable
vegetable. Usually they are dipped in egg and fried, or mixed with
meat stews, and sometimes they are eaten raw in combination with
other raw salad vegetables. The panicles are sold commonly in
the markets, and often they are taken to distant markets of regions
where the plant is scarce or unknown, for sale, as to Puerto Barrios.

ZYGADENUS Michaux

Plants with horizontal rhizomes, sometimes also with tunicated bulbs, the
stems erect, simple below, often branched above; leaves radical or inserted near
the base of the stem, linear and elongate; flowers racemose, the racemes simple
or paniculate; perianth persistent, the segments sometimes connate at the base
into a short turbinate tube, subequal, rotate-spreading, often bearing 2 glands
within near the base; stamens 6, inserted at the base of the perianth segments,
shorter than the perianth, the filaments filiform, connivent about the style, recurved
at the apex; anthers small, subglobose, extrorsely dehiscent and orbicular-peltate
after dehiscence; ovary ovoid, trilobate, each lobe prolonged into a slender style
which is introrsely stigmatose; ovules numerous in each cell; capsule ovoid or
oblong, acuminate, septicidally separating into 3 carpels that are introrsely
dehiscent; seeds oblong to almost linear, often somewhat curved, brown or blackish.

About fifteen species, one in Siberia, the others North American.
Guatemala is the southern limit of distribution for the genus.

Zygadenus volcanicus Benth. PI. Hartw. 96. 1842. Anticlea
\vokanica J. G. Baker, Journ. Linn. Soc. 17: 482. 1880.

Alpine meadows or on mossy rocks in alpine Juniperus forest;
often on limestone, 3,000-3,700 meters; Sacatepe"quez (type from
Volcan de Agua, at 3,300 meters, Hartweg 626); Solola (Los En-
cuentros); Huehuetenango (Sierra de los Cuchumatanes). Southern
Mexico.

Plants glabrous, arising from small bulbs, the outer bulb coats almost black;
leaves 30-40 cm. long, glaucous-green, mostly 6-12 mm. wide; scapes leafy below,
branched above and forming large open bracted panicles, the flowers pedicellate;
perianth segments white or whitish, ovate, 6-8 mm. long, very obtuse, with
2 confluent glands within near the base; capsule about 15 mm. long and 5-6 mm.
wide.

One Guatemalan collection (Skutch 1101, from the Cuchu-
matanes) has been referred by Walsh to Z. elegans Pursh, a United

92 FIELDIANA: BOTANY, VOLUME 24

States species, apparently in error. Certain United States species
of this genus are known to be poisonous to stock, sometimes causing
serious losses.

SMILACACEAE. Cat-brier Family

Vines, woody or herbaceous, the stems prickly or unarmed; leaves alternate,
petiolate, the blades reticulate-veined and with conspicuous longitudinal nerves
arising from the base, articulate with the petioles, these sheathing and often bearing
a pair of tendril-like appendages; flowers dioecious, in axillary pedunculate umbels;
perianth segments 6; stamens usually 6, the filaments distinct, inserted at the base
of the perianth segments; anthers 2-celled, basifixed, introrsely dehiscent; ovary
3-celled or rarely 1-celled, the ovules 1-2 in each cell, pendulous; style very short
or none, the stigmas 3 or rarely only 1 ; fruit a small, red or black berry, containing
1-6 seeds; endosperm osseous.

A small family of about three genera, in tropical and temperate
regions of the whole earth.

SMILAX L.

Reference: E. P. Killip & C. V. Morton, A revision of the Mexican
and Central American species of Smilax, Carnegie Inst. Wash.
Publ. 461: 257-297. pis. 1-11. 1936.

Herbaceous or woody vines, the stems armed with prickles or unarmed, arising
from woody or fleshy tubers or from long creeping rhizomes; leaves usually coria-
ceous, entire or lobate; perianth segments distinct; pedicels borne on a globose or
conic receptacle, inserted in small pits among minute bractlets; staminate flowers
with or without an abortive ovary; pistillate flowers usually smaller than the
staminate, with an ovary and usually several abortive stamens.

More than 200 species, widely distributed in both hemispheres,
most abundant in tropical regions. Several additional species occur
farther south in Central America. The species, or rather some of
them, are of importance as being the source of the drug sarsaparilla
(Zarzaparrilla). Although the plants of this genus are widely
dispersed in Guatemala and often abundant, few of them, as a rule,
are in flower or fruit, and for this reason good specimens are fewer
than they should be. On the North Coast the name "cuculmeca"
is used for some of the species, and the name "zarzaparrilla" (from
which the English "sarsaparilla" is derived) is in common use.
Tejada reports the name "quix" as in use for some of the species in
Huehuetenango.

Plants more or less pubescent, sometimes glabrate at maturity but with at least

a few hairs persistent on the petioles, peduncles, or pedicels; always unarmed.

Branches obtusely quadrangular, glabrate at maturity; staminodia 6 in the

pistillate flower S. subpubescens.

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 93

Branches terete (except the lowest), usually abundantly pilose even at maturity.
Stems densely lanate-tomentose; leaves densely tomentose beneath, tripli-

nerved S. velutina.

Stems pilose or subtomentose, the hairs coarser and at least part of them
spreading; leaves hirsute with long or short hairs, 7-nerved, the nerves

all arising from the base of the blade S. mollis.

Plants glabrous throughout, often armed with prickles.
Staminate flowers 2.8 mm. long or less.

Leaves with conspicuous reticulate veins, membranaceous or chartaceous or,
if coriaceous, small, often aculeate on the nerves beneath; branches

angulate and often flexuous S. spinosa.

Leaves with obscure veins, coriaceous, large, unarmed; branchlets terete,
straight.

Berries globose S. Lundellii.

Berries elongate, acute at each end S. munda.

Staminate flowers 4 mm. long or larger.
Peduncles of the pistillate umbels shorter than the subtending petioles,

subterete S. lanceolata.

Peduncles of the pistillate umbels longer than the petioles, almost always

conspicuously compressed.
Anthers shorter than the filaments; leaves unarmed. Petioles articulate

at or below the middle.
Branchlets terete or irregularly subangulate. Leaves drying blackish.

S. jalapensis.
Branchlets quadrangular.
Leaves drying blackish, 7-nerved, the larger often subcordate at the

* base S. jalapensis var. Botterii.

Leaves drying pale green, 5-nerved, never subcordate at the base.

S. Standleyi.

Anthers longer than the filaments.
Stems terete; petiole articulate below the middle of the free portion;

leaves acute at the base; unarmed S. panamensis.

Stems sharply or obtusely quadrangular, at least below; petiole articulate
above the middle of the free portion ; lower leaves cordate or hastate
at the base, often aculeate.
Berries red; stems obtusely quadrangular, subterete above.

S. arislolochiae folia.

Berries black; stems acutely quadrangular throughout S. Regelii.

Smilax aristolochiaefolia Mill. Gard. Diet. ed. 8. No. 7. 1768.
S. medico, Schlecht. & Cham. Linnaea 6: 47. 1831. S. ornata Lem.
'ill. Hort. 12: pi. 439. 1865. Cocomeca.

Pete*n (Uaxactun); probably extending into Alta Verapaz.
Southern Mexico and British Honduras.

Lower stems obtusely quadrangular, armed with large, flat, slightly curved
prickles 13 mm. long or less, the upper stems obtusely quadrangular or subterete,
pale, smooth or nearly so, sparsely aculeate or unarmed; petioles up to 5 cm. long,
the free portion articulate above the middle; lower leaves ovate to oblong, as much
as 28 cm. long and 14 cm. wide, rounded and mucronate at the apex, deeply cordate
or hastate at the base, usually armed on the nerves with stout yellowish prickles,
the upper leaves much smaller, usually cordate at the base, entire, pale yellowish
green when dry, glabrous, 7-nerved, the veins elevated; staminate pedicels num-

94 FIELDIANA: BOTANY, VOLUME 24

erous, 5-11 mm. long, much shorter than the peduncle, slender, glabrous; perianth
segments oblong, 4 mm. long, spreading; filaments 1-1.3 mm. long, the anthers
1.7-2 mm. long; pistillate peduncles 22 mm. long or less, much exceeding the sub-
tending petiole, compressed, the pedicels 8 mm. long or less; perianth segments
3.5 mm. long; staminodia 6; fruiting peduncles as much as 4 cm. long, the pedicels
4-10 mm. long; berries red, globose, 5-8 mm. in diameter.

The name "escoca" is reported from British Honduras, probably
Maya. This species is believed to be one of the important sources
of commercial sarsaparilla.

Smilax jalapensis Schlecht. Linnaea 18: 451. 1844. S. Botterii
A. DC. Monogr. Phan. 1: 89. 1878. S. jalapensis var. Botterii Killip
& Morton, Carnegie Inst. Wash. Publ. 461: 280. 1936. Kixcul
(Coban, Kekchi).

Damp or wet forest or thickets, 1,200-2,700 meters; Alta Verapaz;
Zacapa; Guatemala; Chimaltenango; Quiche"; Huehuetenango; San
Marcos. Southern Mexico.

Branches and branchlets terete and usually straight, sparsely armed with
straight yellowish prickles, the younger branchlets often with numerous bristle-
like prickles; petioles 2 cm. long or less, articulate about half way between
the tendrils and the base of the blade; lower leaves ovate, 12 cm. long and 7 cm.
wide or smaller, short-acuminate, subcordate at the base, the upper leaves smaller,
ovate-lanceolate, rounded at the base, all rather thin and glabrous, blackening
in drying, entire, 7-nerved; staminate flowers numerous in each umbel, the pedicels
13 mm. long or less, the perianth segments linear or linear-lanceolate, 5-6 mm.
long, acuminate; filaments 2-3 mm. long, the anthers equaling or shorter than the
filaments; pistillate peduncles longer than the subtending petiole, about 15 mm.
long at anthesis and as much as 28 mm. in fruit, strongly compressed, the pedicels
3-4 mm. long; perianth segments 3-3.5 mm. long; staminodia 3 or 6; berries
globose, 8 mm. in diameter; seeds red.

In the typical form of the species the stems are terete; in var.
Botterii they are obtusely quadrangular. The variety is represented
in Guatemala, but the characters are not very conspicuous or well
marked.

Smilax lanceolata L. Sp. PI. 1031. 1753. S. domingenis Willd.
Sp. PI. 4: 783. 1806.

Moist forest or thickets, ascending from sea level to about 1,200
meters; Alta Verapaz; Baja Verapaz; Izabal; Zacapa; Escuintla;
Sacatepe"quez. Mexico and British Honduras to Honduras, Salvador,
Costa Rica, and Panama; West Indies.

Stems terete, glabrous, the lower ones sparsely armed with stout recurved
prickles 5-6 mm. long, the upper ones unarmed; petioles 16 mm. long or less,
articulate at about the middle of the free portion; leaves ovate-lanceolate or ovate,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 95

9 cm. long and 5 cm. wide or smaller, rather thick, dark green and lustrous above,
paler and dull beneath, abruptly acuminate, acute at the base, entire, unarmed,
5-nerved; staminate pedicels 4-7 mm. long; perianth segments ligulate, 4.5-6.5 mm.
long, acute; filaments 3-4 mm. long, the anthers 1.2-2 mm. long; pistillate peduncles
subterete, 7 mm. long or less, much shorter than the subtending petioles; pedicels
4-7 mm. long; perianth segments oblong-lanceolate, 4 mm. long; staminodia 3;
fruiting pedicels 4-10 mm. long; berries dull red or brown, globose, 5-10 mm. in
diameter.

Called "tietie" and "china-root" in British Honduras, and "zarza"
and "corona de Cristo" in Honduras. In Costa Rica known as
"bejuco de canasta," the tough flexible stems being utilized for
aking baskets.

Smilax Lundellii Killip & Morton, Carnegie Inst. Wash. Publ.
461: 265. pi. 2. 1936. Diente de chucho (San Marcos) ; Zarza.

Damp forest or thickets, ascending from sea level to about
1,300 meters; Pete"n (type from Sabana Zis, Lundell 3190); Alta
Verapaz; San Marcos. British Honduras.

A large vine, the lower branches stout, terete, striate, glabrous, armed with
very stout spreading compressed prickles, the upper branches unarmed, pale;
petioles 1-2.5 cm. long, articulate at or above the middle; leaves oblong-lanceolate
or lanceolate, the lower as much as 27 cm. long and 13 cm. wide, the upper ones
much smaller, acute, at the base obtuse or acute, the lower blades often subcordate
at the base, thick and hard, unarmed, glabrous, 5-nerved, drying a rather dull
brownish green, the veins inconspicuous; staminate pedicels 1 cm. long or less,
about equaling the peduncle; perianth segments green, oval, 2 mm. long; filaments
0.8 mm. long, the anthers somewhat shorter; fruiting peduncles 5-11 mm. long,
much shorter than the subtending petiole, compressed; fruiting pedicels 7-10 mm.
'long, exceeding the peduncle; berries globose, 4-6 mm. in diameter, bluish black.

Smilax mollis Humb. & Bonpl. ex Willd. Sp. PI. 4: 785. 1806.
. mollis var. acuminata A. DC. Monogr. Phan. 1: 68. 1878.

Thickets and forest, ascending from sea level to about 3,000
meters; Pete"n; Alta Verapaz; Izabal; Chiquimula; Chimaltenango;
Quezaltenango; San Marcos. Southern Mexico to Panama.

A small or large vine, unarmed, the stems terete, pilosulous or subtomentose;
petioles 18 mm. long or less, densely pubescent, the free portion articulate at or
above the middle; lower leaves ovate-oblong or broadly oval or ovate, 18 cm. long
and 10 cm. wide or smaller, the upper leaves much smaller, ovate to oblong, acute
or apiculate, cordate at the base, rather thin, entire, deep or pale green, sparsely
hirsutulous above when young, soon glabrous and shining, persistently hirsutulous
beneath on the veins, 7-nerved, the nerves all arising from the base of the blade;
staminate peduncle 4 cm. long or less, much exceeding the subtending petiole,
densely short-hirsute, the pedicels 3-4 mm. long; perianth segments oblong-linear,
about 4 mm. long, sparsely pilose or glabrous except for an apical tuft of hairs;

96 FIELDIANA: BOTANY, VOLUME 24

filaments 2-3.5 mm. long, the anthers 1-1.2 mm. long; pistillate peduncles 3 cm.
long or less, terete or slightly compressed, usually longer than the subtending
petioles, the pedicels 3-5 mm. long; perianth segments narrowly oblong, 3.2 mm.
long; staminodia 3; berries red or yellow, globose, 4-8 mm. in diameter.

Called "pate" in Honduras, a word of Nahuatl derivation signi-
fying "medicine." It is stated that in that country the large yam-
like roots are used as a fish poison.

Smilax munda Killip & Morton, Carnegie Inst. Wash. Publ.
461: 265. pi. 3. 1936.

On wooded bluffs, 75 meters; Pete"n (Rio Cancue"n, Steyermark
45937). British Honduras (type from the Rio Grande, Schipp 1181).

A large vine, sometimes 15 meters long, the stems terete, glabrous, the lower
ones armed with a few short straight prickles, the upper branchlets unarmed;
petioles 3 cm. long or less, the free portion articulate above the middle; larger
leaves ovate-lanceolate, as much as 17 cm. long and 9 cm. wide, most of the leaves
much smaller, oblong to narrowly elliptic, acute at the base, coriaceous, unarmed,
entire, 5-nerved, the veins inconspicuous; pistillate peduncles 4-7 mm. long,
slightly shorter than the petiole, compressed, the pedicels 5-8 mm. long, exceeding
the peduncle; perianth segments green, 1-2 mm. long, the outer obovate-oblong,
the inner smaller, oblong; staminodia 3; fruiting peduncles 1 cm. long or less, the
pedicels to 13 mm. long; berries at first red, turning black, ellipsoid, as much as
17 mm. long and 5-7 mm. thick, acute at each end.

Smilax panamensis Morong, Bull. Torrey Club 21: 441. 1894.
Baja Verapaz (Panzal). Honduras; Costa Rica; Panama.

Lower stems ^stout, terete, smooth, glabrous, pale brown, armed with large
straight broad-based prickles 2 cm. long or less, the upper branches unarmed;
petioles 3 cm. long or less, the free portion articulate near the base; leaves ovate-
oblong or the upper lance-oblong, the largest 19 cm. long and 9.5 cm. wide, thick-
chartaceous, short-acuminate, at the base acute or obtuse, entire, glabrous, un-
armed, concolorous, 7-nerved; staminate peduncles 2 cm. long or less, compressed,
the pedicels 5-8 mm. long; perianth segments ovate-lanceolate, 4-6 mm. long,
green, acute; anthers 2-2.7 mm. long, slightly or much longer than the filaments;
pistillate peduncles 1.5 cm. long or less, compressed, the pedicels to 8 mm. in
length; perianth segments blackish when dry, 5 mm. long; staminodia 6; fruiting
peduncles 2.5 cm. long or less, the pedicels 7-15 mm. long; berries probably red,
7.5-10 mm. in diameter.

Smilax papyracea Duham. was reported from the Volcan de
Fuego by Hemsley on the basis of a specimen collected by Godman
and Salvin. The report probably relates to one of the species listed
here, but it is impossible to determine which one without examination
of the specimen.

Smilax Regelii Killip & Morton, Carnegie Inst. Wash. Publ.
461: 272. 1936. S. grandifolia Regel, Ind. Sem. Hort. Petrop. 16.

FIG. 20. Smilax Regelii. A. Habit of portion of flowering stem and leaf
attached to stem in position; X M- B. Leaf of another shape; X M- C. Stamen;
X 6. D. Staminate flower; X 6. Pistillate flower; X 6. F. Portion of stem
showing prickles; X l /i.

98 FIELDIANA: BOTANY, VOLUME 24

1856, not Buckl. 1843. S. ornata Hook. Bot. Mag. 115: pi 7054.
1889, not Lem. 1865. S. utilis Hemsl. in Hook. Icon. 26: pi 2589.
1899, not Wright, 1895. Zarzaparrilla; Bejuco de corona.

In forest or thickets, ascending from sea level to 1,500 meters
or more; Pete"n; Izabal; Jalapa; Santa Rosa; Chimaltenango; Que-
zaltenango; San Marcos. British Honduras to Honduras.

Often a very large vine, as much as 15 meters long, the lower stems acutely
quadrangular, armed, chiefly on the angles, with large, broad, compressed, straight
or recurved prickles 1 cm. long, the upper branches acutely quadrangular, some-
times narrowly winged on the angles, aculeate; petioles of lower leaves up to 7 cm.
long, aculeate, the free portion articulate at or above the middle; lower leaves
often very large, as much as 30 cm. long and 20 cm. wide, ovate to narrowly oblong,
rounded or short-acuminate at the apex, shallowly or deeply cordate or hastate
at the base, the upper leaves much smaller, oblong-lanceolate or oval, acute at
the base, chartaceous, glabrous, often armed beneath with short recurved pale
prickles, the larger leaves 7-nerved, the smaller 5-nerved, bright green or rather
pale green when dried; staminate peduncles 6.5 cm. long or less, longer than the
subtending petioles, the pedicels 7-12 mm. long; perianth segments lanceolate,
3.5-5 mm. long; filaments 1.2 mm. long or less, the anthers 2-2.5 mm. long;
pistillate peduncles up to 10 cm. long but usually shorter, compressed; fruiting
pedicels 9-19 mm. long; fruits globose, 1.3 cm. or less in diameter, black. (Fig. 20.)

Called "zarza" and "zarzaparrilla" on the Atlantic coast of
Honduras. This is believed to be one of the principal sources of
sarsaparilla, a product furnished by various species of Smilax but
by relatively few of them, the roots of some species being apparently
useless commercially. Small quantities of the dried roots are exported
from Guatemala to the United States and Europe. In medicine
sarsaparilla is used for its stimulant and sudorific properties. In-
troduced into Spain about 1540, it was long a favorite remedy for
venereal diseases. It still is employed for the same purpose, and for
rheumatism, scrofulous diseases, and some cutaneous affections. It
is much used in the United States for flavoring soft drinks.

Smilax Regelii f. albida (Killip & Morton) Standl. &
Steyerm., comb, nov., (S. Regelii var. albida Killip & Standl. Car-
negie Inst. Wash. Publ. 461: 273. 1936) is a form with white berries.
It has been collected near Tela, on the Atlantic coast of Honduras.

Smilax spinosa Mill. Gard. Diet. ed. 8. No. 8. 1768. S. mexicana
Griseb. ex Kunth, Enum. PI. 5: 167. 1850. S. Gaumeri Millsp. Field
Mus. 1: 357. 1898 (type from Yucatan, Gaumer 687). Madre de
zarzaparrilla; Zarza; Zarzaparrilla macho; Bejuco de la vida.

Damp or wet thickets or forest, ascending from sea level to about
2,800 meters, most plentiful at low elevations; Pete"n; Alta Verapaz;

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 99

Izabal; Zacapa; Jutiapa; Santa Rosa; Guatemala; Suchitepequez;
Retalhuleu; Quezaltenango; Huehuetenango. Mexico and British
Honduras to Panama.

Lower stems terete, armed with stout prickles or unarmed, the upper branches
usually 4-6-angulate, often flexuous; petioles short, rarely more than 1 cm. long,
aculeate or unarmed, the free portion usually articulate near the middle but often
either above or below the middle; lower leaves ovate or broadly elliptic, up to
14 cm. long and 8 cm. wide but usually much smaller, acute to rounded and
mucronate at the apex, subcordate at the base, the upper leaves smaller, lanceolate
to oval, acute or rounded at the apex, acute at the base, rather thin, concolorous,
glabrous, entire, often aculeate on the nerves beneath, 5-nerved; staminate pe-
duncles 8 mm. long or less, shorter than the subtending petioles, compressed, the
capillary pedicels 6-13 mm. long, exceeding the peduncle; perianth segments
ovate-oblong or oblong, 2.8 mm. long or less; filaments shorter or longer than the
anthers; pistillate peduncles to 9 mm. long, shorter than the subtending petiole,
compressed, the pedicels 5-9 mm. long; perianth segments oblong or oblong-
lanceolate, 1.5-2.8 mm. long; staminodia 3 or 6; berries black, globose, 4-12 mm.
in diameter.

Known in Salvador by the names "bejuco de corona," "zarza,"
and "espuela de gallo." Maya names reported from Yucatan are
"xcoceh," "xcocehac," "coceeh," and "coceh." In Huehuetenango
a beverage is made by boiling the root and flavoring the decoction
with sugar and cinnamon. The decoction is also considered there to
be a preventive of malaria.

Smilax Standleyi Killip & Morton, Carnegie Inst. Wash. Publ.
461:280. 1936.

Damp forest and thickets, 200-1,800 meters; Alta Verapaz;
Chimaltenango; Suchitepequez; Quezaltenango; Huehuetenango.
Pacific lowlands of Costa Rica (Guanacaste).

A small or large vine, the lower stems quadrangular, slender, pale green, armed
with sparse short recurved prickles, the upper stems very slender, pale green,
usually unarmed; petioles 2.5 cm. long or less, articulate below the middle of the
free portion; leaves ovate-oblong, 24 cm. long and 12 cm. wide or usually much
smaller, thin, pale green when dried, unarmed, acute or short-acuminate, obtuse
to subcordate at the base, entire, 5-nerved; staminate peduncles 2.5 cm. long or
less, longer than the petioles, compressed, the pedicels 9 mm. long or less, very
numerous, slender; perianth segments green, lanceolate, 5 mm. long; filaments
longer than the anthers; pistillate peduncles 2 cm. long or less, the pedicels 8-9
mm. long; perianth segments lanceolate, 5 mm. long; staminodia 3; fruiting
peduncles 3 cm. long, the pedicels 1 cm. long; berries red, 8 mm. in diameter,
globose.

The young leaves sometimes are marked with blotches of whitish
or gray.

100 FIELDIANA: BOTANY, VOLUME 24

Smilax subpubescens A. DC. Monogr. Phan. 1: 69. 1878.

Damp thickets or forest, 1,500-2,500 meters; Alta Verapaz;
Zacapa; El Progreso; Chimaltenango; Suchitepequez; Quiche"; Hue-
huetenango; Quezaltenango; San Marcos. Southern Mexico; Hon-
duras; Costa Rica.

A large or small vine, unarmed, the stems obtusely quadrangular or subterete,
rufescent-tomentose when young, almost glabrous at maturity; petioles 7 cm. long
or less, tomentose when young, the free portion articulate above the middle;
lower leaves broadly ovate, as much as 24 cm. long and 19 cm. wide but usually
much smaller, abruptly short-acuminate, deeply cordate at the base, the upper
leaves ovate, acuminate, rounded or subtruncate at the base, rather thin, entire,
densely reddish-tomentose on both sides when young, soon glabrescent, lustrous
above, dull beneath, 9-11-nerved; staminate peduncles 3.5 cm. long or less, usually
shorter than the subtending petioles, compressed, reddish-tomentulose; pedicels
15 mm. long or less; perianth segments narrowly oblong, 5-6 mm. long, sparsely
or densely tomentulose; filaments 2.5-4 mm. long, the anthers 1.6-2 mm. long;
pistillate peduncles about 1 cm. long, compressed, shorter than the subtending
petioles, the pedicels 6 mm. long; perianth segments 4 mm. long; staminodia 6;
fruiting peduncles to 2.5 cm. long, the pedicels 18 mm. long or less; berries bright
orange, subglobose or often slightly narrowed at the base and apex.

One Guatemalan collection of the species has been reported incor-
rectly as Smilax rotundifolia L., a species of the United States.

Smilax velutina Killip & Morton, Carnegie Inst. Wash. Publ.
461: 283. 1936.

Moist forest or thickets, sometimes in pine forest, ranging from
sea level to about 1,500 meters; Alta Verapaz; Izabal; San Marcos(?) ;
Huehuetenango. Chiapas and British Honduras.

A small or large vine, unarmed, the stems terete, densely and softly yellowish-
pilose, becoming glabrate; petioles 2.5 cm. long or less, the free portion articulate
toward the base; leaves oblong to broadly ovate or elliptic, the largest 15 cm. long
and 9 cm. wide, the upper much smaller, apiculate or short-acuminate, at the
base cordate or subcordate, entire, 7-nerved, subtomentose above when young,
soon glabrate, tomentose beneath; staminate peduncles 4 cm. long or less, terete,
the pedicels only 3-4 mm. long; perianth segments linear, 6 mm. long, pubescent
outside; filaments 4 mm. long, the anthers 1.2 mm. long; fruiting peduncles 6-15
mm. long, shorter or longer than the subtending petiole, terete, densely pilose,
the pedicels about 6 mm. long, pubescent; berries black, globose, 6 mm. in diameter.

This species was formerly confused with S. tomentosa HBK., and
has been reported from British Honduras under that name.

HAEMODORACEAE

Perennial herbs, glabrous or often tomentose or pilose, with short, often tuber-
ous rhizomes; leaves crowded at the base or on the lower portion of the scape,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 101

often distichous, linear or ensiform; inflorescence terminal, various in form, often
paniculate or cymose, the bracts mostly small; flowers perfect, regular or nearly
so; perianth corolla-like, persistent or circumscissile and deciduous, the six seg-
ments free or united to form a tube, distinctly biseriate, induplicate-valvate or
slightly imbricate; stamens 6, inserted on the tube or on the lobes of the perianth,
all perfect or 1-3 of them reduced to staminodia, sometimes 3 and opposite the
inner perianth lobes; filaments filiform, long or short, free or shortly united at the
base, the anthers ovate to linear, often sagittate, erect or versatile, 2-celled, the
cells opening by a longitudinal slit or sometimes by terminal pores; ovary inferior
or superior, normally 3-celled, the style filiform or rarely short or none, stigmatose
at the apex or rarely with 3 short stigmas; ovules 1 to many, affixed to an axial
placenta, anatropous; fruit capsular or sometimes fleshy and more or less baccate,
usually loculicidally 3-valvate, sometimes indehiscent; endosperm fleshy.

The family is a rather small one, with only a few genera and
species in America, most of its representatives being Australian and
Asiatic. In Central America a single genus and species are known.

XIPHIDIUM Aublet

Rather coarse perennials with short or elongate, stout rhizomes, the scape
simple below the inflorescence, leafy at the base; leaves broadly linear, flat, succu-
lent, thin when dried, with numerous slender nerves, equitant; flowers small,
white, glabrous, secund and pedicellate along the simple branches of the panicle;
perianth segments distinct, subequal, oblong, spreading; stamens 3, affixed to the
bases of the inner segments, the filaments short, the anthers oblong, erect; ovary
free, 3-celled, the style filiform, stigmatose at the apex; ovules numerous in each
cell, pendulous; fruit a fleshy globose capsule, loculicidally 3-valvate; seeds num-
erous, small, subglobose or slightly angulate, verruculose.

The genus consists of probably a single species.

Xiphidium caeruleum Aubl. Fl. Guian. 33. pi. 11. 1775. X.
floribundum Swartz, Prodr. Veg. Ind. Occ. 17. 1788.

Wet or moist forest or thickets, often on shaded banks, chiefly
at 1,200 meters or less; Alta Verapaz; Izabal; Chiquimula; Santa
Rosa; Escuintla; Guatemala; Chimaltenango (Yepocapa); Suchite-
pequez; Retalhuleu; Quezaltenango; Huehuetenango. Southern
Mexico to British Honduras and Panama; West Indies and South
America.

Plants usually 40-70 cm. high, sometimes taller, glabrous except for the
weakly pilose inflorescence; leaves ensiform or linear, usually 20-40 cm. long,
1.5-5 cm. wide, long-attenuate; panicles thyrsiform, the scape usually bearing
below the panicle a few adventitious buds that fall off and develop into new plants,
the branches of the panicle spreading or more often ascending, or even reflexed,
rather remotely flowered; perianth segments oblong or obovate, obtuse, 6-8 mm.
long, glabrous; fruit subglobose, commonly 7-8 mm. in diameter, varying from
green to dull red or finally purple-black. (Fig. 21.)

FIG. 21. Xiphidium caeruleum. A. Habit; X Vs. B. Flower; X 5. C. Fruit
in position with persistent perianth; X 4.

102

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 103

Called "palma del norte" in Salvador. A rather ornamental
plant, the foliage suggesting somewhat that of the genus Iris. It
is sometimes planted for ornament in local gardens. Although
essentially a plant of the tierra caliente, on the Pacific coast it
does not extend upon the plains, but often grows in abundance in
the lower foothills, sometimes covering densely paredones along
roadsides or steep banks at the bases of cliffs on stream borders.
The small white flowers are neither showy nor handsome. The
specific name is a somewhat misleading one. Aublet states that part
of the petals are blue at the apex, but no such coloration has been
observed in Central American plants.

AMARYLLIDAGEAE. Amaryllis Family

Perennial herbs, rarely frutescent, with rhizomes or tubers or sometimes
with woody, simple or branched caudices, the scapes often tall but sometimes low
or almost obsolete; leaves various in form, thin or fleshy, the nerves conspicuous
or obscure, the leaves sometimes very thick and long-persistent, their margins
often armed with spines; inflorescence terminal, the flowers perfect, variously
arranged; bracts mostly scarious, sometimes colored; flowers regular or nearly
so, the perianth usually marcescent, rarely persistent, the segments free or more
or less connate to form a tube, the segments or lobes 6, distinctly biseriate, those
of the 2 series equal or unequal; stamens generally 6, inserted upon the perianth
tube or at the bases of the segments, the filaments filiform, free or sometimes
connected by a membrane; anthers oblong, linear, or rarely ovoid-globose, erect
or versatile, 2-celled, the cells dehiscent by longitudinal slits; ovary inferior,
3-celled or rarely 1-celled (not in Guatemalan genera), the placentae axial; style
filiform or rarely columnar, usually undivided, the stigma terminal, small and
capitate, rarely of 3 branches; ovules numerous in each cell, biseriate, rarely only
1-2, anatropous; fruit capsular, globose to oblong, loculicidally 3-valvate or some-
times indehiscent; seeds usually numerous, sometimes few or only 1-2, sessile or
on very short funicles, globose to angulate or compressed, the testa usually black;
endosperm fleshy.

About seventy genera are recognized, widely distributed in
tropical and warmer regions of the earth. All the known Central
American genera (and most of the species) are represented in Guate-
mala.

Plants scandent, with leafy stems Bomarea.

Plants not scandent, the stems not leafy.
Plants with tunicated bulbs.

Scapes 1-flowered Zephyranthes.

Scapes bearing more than a single flower.

Filaments connected by a thin cuplike membrane. Flowers white.
Perianth lobes linear; leaves usually strap-shaped, not conspicuously

petiolate Pancratium.

Perianth lobes ovate; leaves elliptic, abruptly contracted at the base into
a slender petiole Eucharis.

104 FIELDIANA: BOTANY, VOLUME 24

Filaments distinct, not connected by a membrane. Flowers often red or
pink.

Flowers mostly sessile or nearly so; seeds few, not compressed . . Crinum.

Flowers long-pedicellate; seeds strongly compressed Hippeastrum.

Plants never with tunicated bulbs.

Leaves thin, membranaceous, conspicuously nerved, not fleshy; flowers bright
yellow; capsule indehiscent or circumscissile.

Capsule circumscissile; flowers slender-pedicellate Hypoxis.

Capsule indehiscent; flowers sessile Curculigo.

Leaves thick and hard or often succulent, not nerved; flowers not bright

yellow.
Perianth tube elongate, longer than the lobes, curved; scape leafy.

Polianthes.

Perianth tube short, much shorter than the lobes, not curved; stems merely
bracted.

Perianth rotate, the tube almost obsolete Furcraea.

Perianth funnelform, with an evident tube Agave.

AGAVE L.

References: William Trelease, The Agaveae of Guatemala, Trans.
Acad. Sci. St. Louis 23: 129-152. pis. 6-35. 1915; Agave, in Standl.
Contr. U. S. Nat. Herb. 23 (Trees and shrubs of Mexico) : 107-142.
1920; Alwin Berger, Die Agaven 1915.

Plants mostly large and coarse, the rhizome short and subterranean, or some-
times erect and short-columnar; leaves all radical or densely clustered at the apex of
the caudex, usually thick, rigid, and fibrous, rarely thin and withering, the margins
usually armed with spines, the tip of the leaf terminating in a sharp spine; scape
terminal, often of gigantic size, the flowers spicate or more often cymose and
forming a large thyrse; perianth subfunnelform, the tube usually short, ampliate
in the throat, the segments linear or oblong, erect or spreading, usually yellowish
white; stamens inserted in the perianth throat at the base of the lobes, longer than
the lobes, the filaments filiform or at the base membranaceous-complanate; anthers
linear, large, dorsifixed at the middle; ovary globose to oblong, often fleshy, con-
tracted into a beak, 3-celled; style filiform from a short conic base, the stigma
small, capitate, more or less trilobate; ovules numerous in each cell, biseriate;
capsule globose to ovoid or oblong-cylindric, often 3-sulcate, coriaceous, often
crowned by the persistent perianth, loculicidally dehiscent from the apex; seeds
numerous, thin and flat, densely crowded, the testa black.

An American genus (some of the species naturalized in the Old
World), with 300 or more species. They are relatively few in Central
America, the majority being Mexican. About 170 are known from
that country, and they are rather numerous also in the West Indies.
A few additional species are known from Salvador, but chiefly in
cultivation, and two have been described from Panama and Costa
Rica. In Central America the plants are not common south of
Guatemala, except in the arid interior regions of Honduras.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 105

The agaves constitute one of the most remarkable of all endemic
groups of American plants, and one of great utility and ornamental
value. These strange plants, so unlike anything in Europe or
adjacent regions, naturally attracted attention of the early explorers.
At a very early date, living plants, easily preserved, were carried
back to Europe, where they became one of the wonders of Mediter-
ranean gardens and of northern hothouses. In Europe, because of
unfavorable growing conditions, the plants often lived for many
years without blooming; hence there developed a belief that they
bloomed only at the age of 100 years, whence the customary English
name of century plant. It is unnecessary to state that in regions
where they are native, the plants require but few years to attain
full development. When once the flower stalk does arise, its growth
is extremely rapid, sometimes a foot or more in a single day. The
plants die after flowering, but each one usually leaves about its
base suckers that develop into new plants. In Europe the agaves
are sometimes known as American aloes, because of their very slight
resemblance to some of the species of the Old World genus Aloe.

In Mexico the agaves are of great industrial importance as a
source of various qualities of fiber, and of the beverages known as
pulque, mescal, and tequila, the last two of which are highly intoxi-
cating. Although species suitable for extraction of these beverages
are available in Guatemala, we cannot find a record that their sap
was ever used by the Indians for preparing intoxicating drinks, and
at the present time it would not be permitted. Some of the local
species are of great importance as fiber plants. The fiber varies in
quality according to the species, but rope and coarse textiles probably
are made from the leaves of all species having leaves of suitable
length. The roots contain saponin, and often are used as a soap
substitute under the name "amole," but much less frequently in
Guatemala than in Mexico.

In Guatemala the plants are of great importance for fences or
hedges, and the larger species, such as A. tecta, are particularly
useful for this purpose. Especially on the plains about Quezalte-
nango there are long miles of agave hedges (often mixed with tall
opuntias) that give a definitely Mexican aspect to the landscape.
Wild agave plants are now confined chiefly to rocky or steep hill-
sides unsuitable for agriculture, but centuries ago they must have
been much more plentiful. When the land was cleared for agricul-
ture, the plants were dug out and either dried and burned or else
replanted along the margins of the fields to mark the boundaries or,

106 FIELDIANA: BOTANY, VOLUME 24

after the conquest, to keep out stock. Even today this transplanting
of the maguey may be observed occasionally in the Guatemalan
highlands. About Huehuetenango one sometimes sees a unique type
of fence made by planting on top of low earth walls small agave
plants of the species having short broad leaves.

Agaves are grown on a large scale in Alta Verapaz for their fiber,
but elsewhere also they are important, and the fiber, raw or manu-
factured, is one of the important articles in all the markets. Large
amounts of it are used for rope and twine, hammocks, halters, carry-
ing nets, carrying bags, and many other articles, even for the strings
of violins and guitars at times. The flower stalks are said to attain
sometimes a height of 10 meters. About Atitlan notches are cut
in them, so that they may be used for ladders. In some regions they
are hollowed and used for conducting water, some of these water
lines being said to have a length of a kilometer. The sticky sap of
the leaves is added to whitewash to make it adhere. The leaves of
both Agave and Furcraea are often gnawed during the dry season
by mules and horses, perhaps also by cattle and sheep when forage
is scant. At Jutiapa the senior author was informed that the
seed pods of some species of maguey are used as a fish poison.
There, too, the young and more tender leaves are sometimes cooked
and eaten, especially when there is a scarcity of maize. Some of
the Mexican agaves have been introduced into the gardens of
Guatemala City and other parts of the country, but most of the plants
seen in local gardens are Guatemalan species.

The following account of Guatemalan species is based upon the
treatment by Trelease, who visited the country and spent consider-
able time in studying the plants in the field. It must be confessed
that the following account is far from satisfactory, except for a few
of the best-marked species. It is believed that the total number of
species is considerably less than is listed here, but the material
available does not permit the reduction of many of the names at
the present time. The group obviously is one that offers great
difficulties in the disposition of species, and the plants need to be
studied in the living state rather than from dried specimens, in which
all the characters are distorted.

Leaves withering, or disappearing in the dry season, new leaves appearing each

season, not spine-tipped, unarmed A. brachystachys.

Leaves persistent, hard, fleshy, fibrous, tipped with a stiff spine, the margins

usually spine-armed.
Inflorescence simple, the flowers spicate. (Subgenus Littaea.)

Leaves with spine-toothed margins A. huehueleca.

Leaves with unarmed margins A. dasylirioides.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 107

Inflorescence, so far as known, branched, the flowers in cymes or headlike

clusters.

Leaves very large and trough-like, mostly about 2 meters long and 50 cm.
broad, very thick and heavy, the marginal teeth usually joined by a

distinct hard horny border A. tecta.

Leaves mostly smaller, at least much narrower, not trough-like, usually
without a continuous horny border, or this sometimes present in species
with very short and broad leaves.

Leaves rather hard-fibrous, sword-like or dagger-like, very narrow in propor-
tion to their length.
Terminal spine of the leaf stout-conic; leaves gray-green or somewhat

glaucous A. Donnell-Smithii.

Terminal spine very slender and tapering, subulate.

Leaves green or dark green, almost flat A. sisalana.

Leaves gray-green or glaucous, deeply concave A. nivea.

Leaves fleshy, mostly lanceolate (sometimes very narrowly so) to obovate.
Plants very large, confined to gardens; leaves lanceolate, mostly more than
a meter long; panicles open, the flowers loosely clustered at the ends
of the branches; introduced species.

Terminal spine of the leaf acicular A. picta.

Terminal spine of the leaf conic A. americana.

Plants smaller, native species; leaves lanceolate to obovate, usually
smaller; panicles usually narrowly oblong, the flowers densely
clustered at the ends of the branches.
Leaves short and broad, mostly obovate, very rigid and stiff, usually

erect or incurved except during and after anthesis.
Terminal spine of the leaf very tortuous or twisted. . . .A. tortispina.
Terminal spine straight or nearly so.

Spine with a very narrow and deep groove on the upper surface,

its sides more or less incurved A. Seemanniana.

Spine with a rather flat groove, its sides not incurved. .A. Hurteri.
Leaves relatively narrow, elongate, usually not incurved.

Terminal spine of the leaf almost flat on the upper side, scarcely if
at all grooved A. sicaefolia.

Terminal spine evidently grooved on the upper side, the groove
often deep and narrow.

Groove of the terminal spine rounded and rather open.
Teeth of the leaf margins widely separated, red-brown.

A. lagunae.

Teeth of the leaf margins close together, chestnut.
Teeth of the leaf margins abruptly contracted into a slender

curved cusp A. Kellermaniana.

Teeth of the leaf margins broadly triangular, not abruptly
contracted A. samalana.

Groove of the terminal spine deep and very narrow, the margins
often incurved.

Teeth of the leaf margins large and stout, from elevated bases,
usually abruptly contracted into a stout cusp.

Terminal spine of the leaf stout-conic A. opacidens.

Terminal spine of the leaf long-acicular, slender .A. tenuispina.

Teeth of the leaf margins small and weak, their bases little if

at all elevated, not conspicuously contracted into a cusp.

Leaves about 60 cm. long, only about 6 cm. wide, their teeth

often easily deciduous A. minarum.

108 FIELDIANA: BOTANY, VOLUME 24

Leaves 60-100 cm. long, mostly 10-15 cm. wide, their teeth

persistent.
Leaves numerous, strongly ascending, about 60 cm. long

and 15 cm. wide A. Thomasae.

Leaves few, spreading, about 100 cm. long and 10 cm. wide.

A. Deamiana.

Agave americana L. Sp. PL 461. 1753. Maguey.

Planted occasionally for ornament about Guatemala City and
elsewhere. Probably native of Mexico, and cultivated there as well
as in many parts of Central America; naturalized in many parts of
the Old World, especially about the Mediterranean.

Plants very large, the leaves mostly 1-2 meters long and as much as 30 cm.
wide, usually upcurved toward the end, gray-green, very thick and heavy; terminal
leaf spine stout-conic, about 3 cm. long and 7 mm. thick, often recurved, narrowly
grooved below the middle, scarcely decurrent, the marginal teeth narrowly tri-
angular, hardened into the tops of fleshy protuberances between which the margin
is somewhat concave; inflorescence very tall, often 5-8 meters high, branched and
rather open; perianth 8.5-9 cm. long; ovary cylindric, 4-4.5 cm. long; perianth
segments 2.5-3.5 cm. long, linear or linear-lanceolate, obtuse, bright yellow-green;
anthers yellow-green, 3 cm. long; capsule 4 cm. long; seeds about 8 mm. long and
6 mm. wide.

Part of the plants cultivated in Guatemala belong to var. margi-
nata Trelease (in L. H. Bailey, Stand. Cycl. Hort. 1: 235. 1914),
in which the leaves have white or yellow margins. This species is
perhaps better known and more plentiful in southern Europe than
it is in America, for there it has become thoroughly naturalized in
some regions. It often is grown for ornament in the United States
in greenhouses, or in the South out of doors.

Agave brachystachys Cav. Descr. 453. 1802. Manfreda
brachystachys Rose, Contr. U. S. Nat. Herb. 8: 20. 1903. Amole;
Ixmaxin (San Sebastian, Huehuetenango) ; Lirio verde (fide Aguilar) ;
Amol chipalcu (Huehuetenango).

Brushy hillsides or in meadows, most often in pine-oak forest,
850-1,800 meters; Chiquimula; Jalapa; Jutiapa; Santa Rosa; Chimal-
tenango; Solola; Quiche"; Huehuetenango. Mexico; Honduras;
Salvador.

Plants acaulescent, arising from a thick underground rhizome; leaves 12-15
or fewer, lance-linear, soft and thin, 25-35 cm. long, 2-4 cm. wide, green or pale
green, often lying flat on the ground, frequently spotted with brown-purple, the
thin margins entire or nearly so, not spiny, the tip of the leaf unarmed; inflorescence
very slender, 1-2 meters tall, with a few remote bracts, the flowers sessile in mostly
elongate and interrupted spikes, few; ovary elongate-cylindric; perianth 3.5-4 cm.
long, somewhat curved, the lobes slightly longer than the tube, pale green; fila-

FIG. 22. Agave brachyslachys. A. Habit; X Viz- B. Portion of inflorescence
and single detached leaf; X H- C. Flower; X 1. D. Capsule; X 1. E. Seed;
X 4.

109

110 FIELDIANA: BOTANY, VOLUME 24

ments dull lavender; capsule oval, 2-2.5 cm. long, in age becoming whitish, sub-
terete. (Fig. 22.)

In general appearance this is very unlike the normal type of
Agave species, and there is much justification in giving generic rank
to the species of this relationship that have been referred to Man-
freda. The fleshy roots are said to be used in Guatemala, especially
by the Indians of the highlands, for washing textiles. They are
used also for shampooing the hair because it is believed locally that
they prevent baldness, something not very common among the
Indians. The Indians are said to employ the roots also as a barbasco
or fish poison. Although widely scattered in oak and pine forest,
we have not found this plant plentiful anywhere, and usually it
occurs as isolated individuals. The name "cebolla de cerro" is
recorded from Honduras, and the flower stalks are said to be used
there as arrow shafts, presumably by some of the Indians.

Agave Deamiana Trelease, Trans. Acad. Sci. St. Louis 23: 139.
pi. 13. 1915.

Slopes of ravine, 1,200 meters; endemic; Guatemala (type from
Fiscal, C. C. Deam 6154a).

Plants acaulescent, producing few or no offshoots; leaves grayish or slightly
glaucous, widely spreading, narrowly oblanceolate, attenuate, somewhat concave,
about a meter long and 10 cm. wide, minutely roughened, especially beneath;
terminal spine purplish brown, somewhat lustrous above, minutely granular
beneath, slender-conic or acicular, sometimes flexuous, involutely grooved from
about the middle, with acute edges, decurrent for about its own length, scarcely
intruded into the green tissue, 4 cm. long, 3-4 mm. thick; marginal teeth chestnut-
colored, 1-2 cm. apart, 2-4 mm. long, straight or curved (usually downward),
the slender triangular cusp lenticular-dilated into the straight margin; inflorescence
unknown.

This species grows about Fiscal in association with Hechtia,
probably on the dry hills northeast of the railroad station. We
have examined two collections of this species (Deam 6199 and 6201)
determined by Trelease in the United States National Herbarium.

Agave dasylirioides Jacobi & Bouch^, Hamb. Gart. Zeit. 21:
344. 1865.

Described from plants cultivated in Europe; said to have been
collected by Warscewicz at the foot of a volcano (Santa Maria?)
near Quezaltenango; cultivated in Europe.

Plants acaulescent; leaves very numerous, linear, ascending or recurved, 60-
100 cm. long, at the base 2.5-4 cm. wide, narrower above, flat, leathery below, thin
toward the apex, green or grayish green, smooth, the margins unarmed but some-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 111

what roughened, the terminal spine small, brown, compressed; inflorescence 1.5-2
meters tall, simple, spicate, often recurved, bearing numerous dry bracts, densely
many-flowered, the flowers sessile or nearly so; perianth 4 cm. long, green, the
segments lanceolate, subobtuse, 14-16 mm. long, 5 mm. wide, bright green; anthers
brownish or golden yellow; capsule 2.5 cm. long, 1 cm. thick; seeds semiorbicular,
3 mm. wide.

Very distinct from other local species in its long narrow unarmed
leaves, which are only 1-1.5 cm. wide at the middle. Possibly it
may occur on the dry barren rocky mountain sides above Quezal-
tenango, which we have not explored carefully, but it is somewhat
strange that it has not been introduced into cultivation in Quezal-
tenango if growing wild in that region.

Agave Donnell-Smithii Trelease, Trans. Acad. Sci. St. Louis
23: 144. pis. 23, 24- 1915. Maguey.

Brushy rocky slopes, moist quebradas, or moist thickets, often
planted in hedges or for ornament, 200-2,200 meters; endemic;
Zacapa (La Fragua) ; Santa Rosa (Barberena) ; Escuintla (type from
Escuintla, J. D. Smith 2085); Guatemala (Fiscal; planted at Guarda
Viejo and elsewhere); Chimaltenango (between Chimaltenango and
San Martin Jilotepeque) ; Huehuetenango (hills north of Chiantla) ;
Retalhuleu; Suchitepequez.

Plants acaulescent, producing numerous offshoots; leaves light green and
often whitened above, glaucous beneath, ascending or suberect, sword-shaped,
moderately concave, 75-100 cm. long or more, about 8 cm. wide, attenuate, smooth;
terminal spine garnet, becoming chestnut, lustrous, smooth, stoutly awl-shaped,
terete, flat or shallowly grooved near the base, not decurrent or intruded into the
green tissue, 12-15 mm. long, 4 mm. thick; marginal teeth colored like the terminal
spine, 1-3 cm. apart, 2-3 mm. long, upcurved, triangular, lenticularly dilated
into the straight margin; inflorescence tall, paniculate, the pedicels 5-10 mm.
long, the flowers greenish yellow, 4-4.5 cm. long; ovary 2-2.5 cm. long, about
equaling the perianth, oblong-fusiform.

This seems to be a fairly common species of the central mountains,
especially in hedges. The type collection was originally determined
and reported as Agave rigida J. Miller. According to Trelease,
A. Donnell-Smithii is related to the Mexican A. tequilana Weber,
the common source of the liquor mezcal de Tequila.

Agave huehueteca Standl. & Steyerm. Field Mus. Bot. 23: 4.
1943. Maguey.

Dry rocky exposed hillsides, mostly or wholly on limestone, 1,700-
2,200 meters, Huehuetenango; type collected along Aguacatan road
east of Huehuetenango at km. 13-14, Standley 82039; collected also

112 FIELDIANA: BOTANY, VOLUME 24

on hills north of Chiantla and at the crossing of Rio San Juan
Ixtan, east of San Rafael Pe"tzal; endemic.

Leaves forming a very dense rosette, numerous, oblong-ovate or obovate,
about 35 cm. long and 10 cm. wide or often larger or smaller, suberect or ascending
and usually somewhat incurved, more or less concave, thick, hard, fibrous, grayish
green or glaucescent, very rigid; terminal spine conic or elongate-conic, stout,
2.5-4 cm. long, 6 mm. thick, grayish brown, usually dull but sometimes lustrous,
terete above, broadly sulcate below, decurrent into the leaf tissue; margin of the
leaf with a broad continuous gray border, the large coarse teeth 5-11 mm. long or
even larger, compressed, grayish, very broad at the base, often curved or recurved
or even flexuous; inflorescence 3-5 meters tall, pole-like, simple, spicate, erect,
the spike very dense and many-flowered, the bracts numerous, almost setaceous
from a broad base, the flowers almost sessile, clustered; ovary 12 mm. long, 4 mm.
thick; perianth tube obconic, 5 mm. long, the lobes narrowly oblong, dark purplish
green, 2-2.5 cm. long, obtuse; stamens long-exserted, the filaments dark purplish
red, the anthers almost 2 cm. long; capsule globose-oval, about 2 cm. long and 12
mm. broad, very shallowly sulcate, green; seeds semiorbicular, 4 mm. long.

A handsome plant, plentiful locally along the southern base of
the cordillera of Huehuetenango, but not common. It may well
be that the Guatemalan plant can be identified with some species
described from Mexico, but we have not been able to place it among
the Mexican species. It is the easternmost species known of the
subgenus Littaea. This species is noteworthy for the fact that each
blooming plant usually is encircled by a ring of suckers or hijos
almost as large as the parent plant.

Agave Hurteri Trelease, Trans. Acad. Sci. St. Louis 23: 136.
pis. 8-10. 1915. A. pachycentra Trelease, op. cit. 135. pi. 7. 1915.

Maguey.

Usually on dry, open, often rocky or brushy hillsides, sometimes
in pine and oak forest, ascending from about 1,000 meters (probably
at times even lower) to 3,700 meters (Sierra de los Cuchumatanes) ;
endemic; El Progreso (type of A. pachycentra from Cruz, Trelease 2);
Jalapa; Totonicapan; Quezaltenango (type from Zunil, Trelease 3);
San Marcos; Huehuetenango.

Plants acaulescent, often producing offshoots; leaves grayish or glaucous,
ascending or somewhat incurved, forming a large dense rosette, oblong-obovate
or broadly oblong-oblanceolate, 35-80 cm. long, mostly 10-20 cm. wide, acute,
somewhat concave, smooth; terminal spine chestnut, often lustrous above, stout-
conic or sometimes slender, 2-4 cm. long and as much as 8 mm. thick, sometimes
even longer, straight or nearly so, more or less intruded below into the green tissue;
teeth colored like the spine, 5-10 mm. long, 1-2 cm. apart, straight or usually
curved, abruptly cuspidate; inflorescence 2-5 meters tall, oblong-paniculate, the
flowers densely massed in large subglobose clusters, these almost sessile or stout-
pedunculate; flowers on pedicels 1 cm. long or less, whitish or pale yellow, 6-7 cm.
long; ovary 2.5-3 cm. long; perianth tube obconic, 12-15 mm. long, the segments

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 113

2.5 cm. long; capsule oblong, about 5 cm. long and 2 cm. thick, neither stipitate
nor beaked; seeds 8 mm. long and 5 mm. wide.

The species was dedicated to Mr. G. Hurter of Quezaltenango.
It has a remarkably wide distribution, at least altitudinally, its
extreme upward limit being in the Sierra de los Cuchumatanes, on
the low cedar-clad hills rising above the alpine plains of that region,
where it is abundant and constitutes a conspicuous element of the
landscape. It is hard to believe that these plants should belong to
the same species that is found as low as 1,000 meters. We have
collected numerous specimens that are referred here. While it is
easy to indicate minor differences between some of these, it has been
found impossible to separate them in a key by characters that appear
plausible; consequently we have preferred to treat them all as forms
of a single not too variable species. For this we have chosen the
name A. Hurteri, rather than A. pachy centra published on an
earlier page, because the former is illustrated and was described
from complete, rather than sterile, specimens. The plant is a highly
ornamental one because of its compact, often almost globose and
somewhat cabbage-like clusters of leaves. It abounds in Huehue-
tenango, especially, but is common also in some parts of Totonicapan,
especially about Momostenango, growing usually on exposed rock
outcrops, and it is plentiful in many localities in hedge-rows. The
flowers often are full of wasps, but these leave the plant when the
stalk is cut and falls to the ground.

Agave Kellermaniana Trelease, Trans. Acad. Sci. St. Louis,
23: 142. pi 18. 1915.

Dept. Guatemala, the type from sides of ravines near Fiscal,
1,110 meters, C. C. Deam 6239.

Plants acaulescent; leaves very glaucous, elongate-lanceolate, about 100 cm.
long and 8-10 cm. wide or larger, acuminate, rather concave, minutely granular
beneath; terminal spine chestnut-colored, somewhat lustrous, slender-conic,
granular below, round-grooved from above the middle with rather blunt edges,
decurrent on the margins for 1-2 times its length, somewhat intruded dorsally
into the green tissue, 3-3.5 cm. long, 4-5 mm. thick; marginal teeth 1-2.5 cm.
apart, 3-5 mm. long, upcurved, the cusps rather slender, lustrous purple-chestnut,
with abrupt bluish bases; inflorescence unknown.

Agave lagunae Trelease, Trans. St. Louis Acad. Sci. 23: 143.
pi. 21. 1915.

On gravel, talus, and cliffs, Pacific slope, 700-1,500 meters;
Guatemala (type from Amatitlan, above the lake, Trelease 10);
Escuintla (between Palin and Escuintla); endemic.

114 FIELDIANA: BOTANY, VOLUME 24

Plants acaulescent, with few offshoots; leaves very glaucous, spreading, lanceo-
late, acuminate, moderately concave, minutely granular on the lower surface,
about 30-40 cm. long and 8-10 cm. wide; terminal spine garnet-colored, lustrous,
granular below, conic, often slightly flexuous, round-grooved from near the apex,
with acute short-decurrent edges, somewhat intruded dorsally into the green
tissue, 2-3 cm. long, 3-5 mm. thick; teeth colored like the spine, lustrous, 2-4 cm.
apart, 3-6 mm. long, upcurved or recurved, the cusps rather slender, mostly unci-
nate, lenticularly dilated into the margin or on the tops of fleshy prominences
between which the margin is almost straight; inflorescence unknown.

Agave minarum Trelease, Trans. Acad. Sci. St. Louis 23: 139.
pi. 14. 1915. Maguey.

El Rancho, the type from Sierra de las Minas, 600 meters, Keller-
man 5129; Zacapa; at about 200 meters; perhaps also at Jutiapa,
900 meters, in dry thickets.

Plants acaulescent, with few or no offshoots; leaves yellow-green, loosely
spreading, lanceolate, acute, smooth, about 60 cm. long and 6 cm. wide; terminal
spine brown, lustrous above, conic, smooth, almost straight, round-grooved above,
becoming involute as the leaf dries, decurrent for somewhat more than its own
length, somewhat intruded dorsally into the green tissue, about 4.5 cm. long and

5 mm. thick; marginal teeth easily detachable, 5-10 mm. apart, the intervening
margin straight; inflorescence 2-3 meters tall, loosely few-branched above, the
flowers relatively lax at the ends of the branches, the pedicels 2-10 mm. long;
flowers yellow, 4 cm. long; ovary 2 cm. long, oblong-fusiform; perianth tube conic,

6 mm. long, the lobes 1.5 cm. long; filaments twice as long as the perianth
lobes.

Agave nivea Trelease, Trans. Acad. Sci. St. Louis 23: 143. pi. 22.
1915. Maguey.

Brushy hillsides, rocky slopes, damp thickets, 400-2,000 meters;
endemic; El Progreso (type from El Rancho, in hedge-rows, Trelease
11); Zacapa; Jalapa; Jutiapa; Solola(?); Huehuetenango.

Plants acaulescent, producing numerous offshoots; leaves mostly very glaucous,
narrowly lanceolate, ascending or suberect, very concave, usually minutely granular
beneath, sometimes 2 meters long and 15 cm. wide, often smaller; terminal spine
chestnut-colored or black, somewhat lustrous, slender-conic, very shallowly
grooved near the base, 1.5-2 cm. long, 3 mm. thick, scarcely decurrent or intruded
into the leaf tissue; teeth colored like the leaves, rather lustrous, 3-5 cm. apart,
about 3 mm. long, usually upcurved, the cusps often curved or flexed, slender,
broadened below and often from somewhat raised, fleshy bases between which
the margin is almost straight; inflorescence said to be an open panicle of yellowish
flowers.

Agave opacidens Trelease, Trans. Acad. Sci. St. Louis 23: 140.
pi. 16. 1915.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 115

El Progreso (type from Cruz, Trelease 5); Huehuetenango (fre-
quent at 1,700-1,900 meters, in hedges, open rocky slopes, or oak
forest); endemic.

Plants acaulescent, producing few or no offshoots; leaves glaucous, lanceolate,
acuminate, roughened beneath, about 75 cm. long and 8-10 cm. wide; terminal
spine dull brown, turning grayish, granular below, stoutly triquetrous-conic, almost
straight, involutely grooved from above the middle with acute edges, decurrent
for 1-2 times its length, somewhat intruded dorsally into the green tissue, 5-6 cm.
long, 3-6 mm. thick; marginal teeth gray, rather dull, 2-5 cm. apart, 5-8 mm.
long, almost straight, heavily triangular, lenticular-dilated into more or less
elevated prominences of the otherwise almost straight margin; inflorescence about
5 meters tall, the upper half oblong-paniculate; bracts broadly triangular, ap-
pressed, the pedicels about 1 cm. long; flowers yellow, 4-5 cm. long; ovary 2.5-3
cm. long, fusiform; perianth tube open-obconic, 1 cm. long, the lobes 12-15 mm.
long, 5 mm. wide; capsule oblong, stipitate, 4-4.5 cm. long, 2 cm. broad; seeds
about 6 mm. long and 5 mm. wide.

Although coming from a region far removed from the type locality,
the Huehuetenango specimens seem to agree well with the original
description and illustration.

Agave picta Salm-Dyck, Bonplandia 7: 88. 1859. A. ingens
Berger, Hort. Mortol. 12: 360. 1912.

Planted occasionally for ornament in parks and gardens, as at
Guatemala, Antigua, Huehuetenango, Retalhuleu, and elsewhere.
Doubtless native of Mexico, but probably unknown in a wild state.

Plants acaulescent or nearly so; leaves numerous, curved upward and outward,
dark green, thick, lanceolate, about 2 meters long and 15-35 cm. wide, or often
much smaller, attenuate, very convex beneath, the marginal teeth rather small
and mostly straight, 2-4 cm. apart, reddish brown or grayish, the terminal spine
4-5 cm. long, slender, acicular, straight; inflorescence sometimes 10 meters tall,
the stalk sometimes 15 cm. thick, the inflorescence rather openly paniculate but
oblong; flowers similar to those of A. americana.

Agave samalana Trelease, Trans. Acad. Sci. St. Louis 23: 142.
pis. 19, 20. 1915.

Quezaltenango (type collected on mountain sides along Rio
Samald near Esperanza, between Santa Maria de Jesus and Zunil,
Trelease 7).

Plants acaulescent, scarcely producing offshoots; leaves glaucous, especially
beneath, numerous, oblanceolate, acuminate, somewhat concave, minutely rough-
ened beneath, about 60 cm. long and 15 cm. wide; terminal spine reddish, darkening
in age, somewhat lustrous, granular below, slender-conic or acicular, almost straight,
round-grooved from above the middle with acute edges, decurrent for about its
own length, more or less intruded dorsally into the green tissue, 3.5-5.5 cm. long,
3-4 mm. thick; marginal teeth chestnut-colored, dull, 5-20 mm. apart, 1-5 mm.

116 FIELDIANA: BOTANY, VOLUME 24

long, straight or slightly upcurved, triangular, rather lenticularly widened into
the nearly straight, denticulate margin; inflorescence 3-4 meters tall, the upper
half narrowly paniculate, with very short branches; capsule oblong, 5-6 cm. long,
2 cm. broad; seeds about 7 mm. long and 5 mm. wide.

Judging from the original habit photograph of this species, it is
one of the agaves growing along the very steep hillsides above the
road leading down from Zunil to Santa Maria. Agaves are very
plentiful in this region, at least on slopes too steep for planting
with corn (which means almost perpendicular), but although we
have made numerous Agave collections in this region, none of them
are certainly identifiable with A. samalana.

Agave Seemanniana Jacobi, Abh. Schles. Ges. Vaterl. Cult.
1868: 154. 1868. Maguey.

Dry, rocky, open or brushy slopes, 400-2,200 meters; Baja
Verapaz; Zacapa; Chiquimula; Jalapa; Jutiapa; Guatemala; Solola.
Also in Nicaragua, the type from Segovia.

Plants acaulescent, producing few or no offshoots; leaves glaucous, spreading
or somewhat incurved, oblanceolate-oblong or obovate, acute or acuminate,
minutely roughened toward the apex, about 35 cm. long and 8 cm. wide; terminal
spine purplish brown, somewhat lustrous above, minutely granular below, slender-
conic or acicular, sometimes somewhat flexuous, involutely grooved from below
the middle with acute edges, decurrent for its own length or less, scarcely intruded
into the green tissue, 2-3 cm. long, 2-4 mm. thick; marginal teeth brown or becom-
ing gray, 1-2 cm. apart, 2-3 mm. long or somewhat larger, rather straight but the
lower ones directed downward, the slender cusps triangular, abruptly dilated to
the tops of fleshy prominences between which the slightly concave margin some-
times bears slender dark denticles; inflorescence about 2-4 meters tall, or some-
times higher, the panicle oblong, the flowers densely crowded into short-stalked
subglobose clusters; capsule broadly oblong, deeply trisulcate, about 3.5 cm. long
and 1.5 cm. broad.

The seed pods of this species are said to be used in Jutiapa for
poisoning fish, and in the same region it is stated that the young
and more tender leaves are sometimes roasted and eaten. Such
use of Agave leaves for food was formerly common among the Indians
of the southwestern United States. The flesh is said to contain a
large amount of sugar, thus making it palatable. In Huehuetenango
also the leaves are gathered for food, as the authors have observed
in the mountains above Chiantla. A. Seemanniana has been intro-
duced into cultivation in Europe, and even carried to Australia.
Several minor varieties have been described from cultivated plants.

Agave sicaefolia Trelease, Trans. Acad. Sci. St. Louis 23: 141.
pi. 17. 1915.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 117

Dry open hillsides and in hedges, ascending from about 300
to 3,000 meters; El Progreso (type from El Rancho, Trelease 6);
Totonicapan (San Francisco el Alto to Momostenango) ; endemic.

Plants acaulescent, often forming dense clumps; leaves gray-green or glaucous,
numerous, crowded, lanceolate, acuminate, almost flat to deeply concave, slightly
granular beneath toward the apex, 60-75 cm. long and 7-8 cm. wide or even larger,
rather thick but flexible; terminal spine purplish chestnut to grayish brown, rather
lustrous, granular below, triquetrous, almost straight, flattened on the upper sur-
face or very openly grooved, with acute short-decurrent margin, somewhat in-
truded dorsally and ventrally into the green tissue, 3-4 cm. long, 4 mm. thick;
marginal teeth colored like the spine, somewhat lustrous, 1.5-3.5 cm. apart,
3-5 mm. long, mostly upcurved, the cusps rather slender, sometimes uncinate,
lenticular-dilated at the base, the rather concave intervening margin sometimes
with 1 or more minute brown teeth; inflorescence unknown in details, but in the
dry state 3-4 meters tall, the flowers in very dense and large, short-stalked clusters,
forming a narrow panicle.

The Momostenango plant grows in a region remote from the type
locality, but leaf material of it agrees well with the original diagnosis
and illustration. This is a large and conspicuous plant, prominent
on the hillsides where it grows.

Agave sisalana Perrine, U. S. Sen. 25th Congr. Sess. 2. Doc.
300: pis. 1,2, 1*. 1838. Maguey.

Planted abundantly in some regions, and, often escaping, seen
in many localities in hedges or fence-rows; Pete"n; Alta Verapaz;
Baja Verapaz; El Progreso; Santa Rosa; Escuintla; Suchitepequez.
Native of the Yucatan Peninsula, but cultivated in many other
tropical regions upon a small or large scale.

Plants acaulescent or with a short caudex (exposed after the leaves have been
cut); leaves sword-shaped, about 1.5 meters long and 10 cm. wide, lightly glaucous,
becoming green, almost flat, the terminal spine chestnut-colored, conic, sometimes
lightly recurved, shallowly round-grooved toward the base, not decurrent upon
the green tissue, 2-2.5 cm. long, 4-5 mm. thick, the margins unarmed or with a
few rudimentary teeth; flowers 4.5-6 cm. long, the ovary and lobes of about equal
length; anthers 2.5 cm. long; bulblets often produced in large numbers.

Called "mescal" in Honduras; the Maya name of Yucatan is
"yaxci." The typical form of the species has almost unarmed leaves;
f. armata Trelease (Contr. U. S. Nat. Herb. 23: 118. 1920) has the
leaf margins rather closely beset with small spiny teeth; both occur
in Guatemala. This plant is of great commercial importance, and
is the basis of the principal industry and the prosperity of Yucatan.
Its fiber is the only one wholly satisfactory for manufacture of the
twine used in binding bundles of wheat when that grain is harvested.
Although the plant (its product is known in the United States as

118 FIELDIANA: BOTANY, VOLUME 24

"sisal" or "sisal hemp") has been introduced into various parts of
the tropics and planted more or less extensively, Yucatan continues
to be the chief source of the fiber. Sisal has been planted in small
amounts in various parts of the Guatemalan lowlands, but the
principal and only large plantation is one in Alta Verapaz near Santa
Cruz. This produces sufficient fiber, it is said, for manufacture of
a large part of the coffee sacks used in Guatemala. Rope also is
made of it, and the rope of Coban is sold all over Guatemala. It is
impossible to say what part of the rope and textiles sold in Guatemala
are made of sisal and what part of Furcraea, for the two sometimes
are grown together, and it is not certain that ordinarily any dis-
tinction is made between them.

Agave tecta Trelease, Trans. Acad. Sci. St. Louis 23: 145.
pis. 26, 27. 1915. Maguey.

Abundant in hedges in many regions in the western highlands,
and not observed in a truly wild state, 1,500-2,600 meters; Chimal-
tenango; Totonicapan; Huehuetenango; Quezaltenango (type from
Quezaltenango, Trelease 17) ; probably also in Quiche" and San Marcos,
although we have no notes or specimens to authenticate its occur-
rence there; reported by Trelease also from Sacatepe"quez and Guate-
mala; endemic.

Plants very large and coarse, acaulescent, producing offshoots; leaves grayish
green or rather dark green, lanceolate or broadly lanceolate, ascending and then
often outcurved, acuminate, very concave and trough-like, smooth, 1-2 meters
long, 50 cm. wide or narrower, very thick; terminal spine chestnut-colored, becom-
ing gray, dull, smooth, conic or subacicular, often slightly flexuous, shallowly
grooved below the middle, 4.5-6.5 cm. long, 5-7 mm. thick, long-decurrent, some-
what intruded dorsally into the green tissue; marginal teeth colored like the spine,
often lustrous, 4-7 cm. apart, about 8 mm. long, recurved, the cusps triangular,
often curved, abruptly dilated into low bases 15 mm. wide or more, the smooth
intervening margin almost straight; inflorescence 5-6 meters tall, the panicle
ellipsoid, rather openly branched, the bracts broadly triangular, densely imbricate
below and sometimes completely covering the scape, those of the panicle smaller
and spreading, the thick pedicels 1 cm. long or less; flowers yellow, about 7 cm.
long, the ovary 3-3.5 cm. long; perianth tube obconic, 1-1.5 cm. long, the lobes
about 2 cm. long; filaments long-exserted; capsule oblong, about 6 cm. long and
3 cm. broad.

This is one of the few easily recognizable Agave species of Guate-
mala, distinguished by its massive leaves of trough-like form. It
is perhaps the most conspicuous plant of the plains about Quezal-
tenango, where there are many miles of hedges composed of it and
Opuntia, imposing in appearance but none too handsome, at least
during the dry months when the leaves are more than ordinarily

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 119

gray with dust. The species is related to the Mexican group of
agaves that supply pulque, and it may have been introduced into
Guatemala from Mexico. While this is possible, it is less probable,
because the size of the plants would make their transportation from
such a distance difficult. It seems more likely that the plant was
originally native in the Quezaltenango region and perhaps else-
where in Guatemala, but that the wild plants have all been reset
in the hedges. It is recorded by Fuentes y Guzman (Recordacion
Florida 1: 289. Madrid, 1882; fide Trelease) that two centuries ago
excellent pulque was produced at Almolonga or Ciudad Vieja in
Sacatep4quez and at San Gaspar. Quite possibly this was the result
of the influence of the Mexican mercenaries who took part in the
conquest of Guatemala. Trelease is of the opinion that Agave tecta
must have been the species used for the purpose. Of course, it is
conceivable that there may have been large plantations formerly
about Antigua and that the plants were destroyed when pulque was
no longer demanded or permitted. From the few plants of Nopalea
now surviving about Antigua, one would never suppose that in the
valley there were formerly hundreds or thousands of acres of Nopalea
grown for production of cochineal. The concave leaves of Agave
tecta are sometimes lapped end over end to make troughs for conduct-
ing water short distances. Their irregular form makes them rather
unsatisfactory for this purpose.

Agave tenuispina Trelease, Trans. St. Louis Acad. Sci. 23: 140.
pi. 15. 1915. Maguey.

At about 1,200-1,500 meters; El Progreso (type from Cruz, on
mountain sides, Trelease 4); Jalapa; Huehuetenango.

Plants acaulescent, with few or no offshoots; leaves glaucous, forming a rather
compact rosette, narrowly lanceolate, acuminate, smooth, about 70 cm. long and
as much as 20 cm. wide, stiff; terminal spine dull brown, acicular, smooth or some-
what roughened at the base, slightly undulate or straight, involutely grooved from
above the middle with rather blunt edges, decurrent for hardly half its length
and not intruded into the green tissue, 6-7 cm. long, 3 mm. thick; marginal teeth
brown, somewhat lustrous, 2-4 cm. apart, 5-10 mm. long, curved in either direction,
rather narrowly triangular, somewhat deltoidally dilated into fleshy prominences;
inflorescence unknown.

The determination of the Huehuetenango material is perhaps
questionable, but it agrees well enough with the original description
and illustration of the species.

Agave Thomasae Trelease, Trans. Acad. Sci. St. Louis 23: 138.
pi. 12. 1915.

120 FIELDIANA: BOTANY, VOLUME 24

Described from a plant cultivated at Quezaltenango, said to have
come from the mountains, Trelease 19, in April, 1915; a collection
from hillsides near Zunil, 2,200 meters, probably represents the same
species.

Plants acaulescent, producing offshoots; leaves green or slightly grayish,
ascending with an outcurved tip, spatulate-oblanceolate, acute or acuminate, not
very thick, flexible or rather stiff, concave, smooth above, finely roughened beneath,
about 60 cm. long and 15 cm. wide or sometimes larger; terminal spine chestnut-
colored, somewhat roughened, acicular, almost straight, involutely grooved below
the middle with acute edges, little decurrent or intruded into the green tissue,
about 2.5 cm. long and 2 mm. thick; marginal teeth colored like the spine, 5-10
mm. apart, 1-3 mm. long, almost straight, the slender cusp slightly dilated at the
base, the straight intervening margin with numerous minute needle-like teeth;
inflorescence unknown.

A plant that probably represents this species has been noted in
patios at several places in Quezaltenango. The leaves are much
less armed, except apically, than in most of the other species. The
species was named for Sra. Dora Thomas, who owned the type plant.

Agave tortispina Trelease, Trans. Acad. Sci. St. Louis 23: 135.
pi. 6. 1915.

El Rancho (type from Cruz, Trelease 1).

Plants acaulescent, producing few offshoots; leaves very blue-glaucous, ascend-
ing in a close rosette, broadly oblong-oblanceolate, acute, concave, minutely
granular beneath, 35-60 cm. long, 15-20 cm. wide; terminal spine gray, dull or
slightly lustrous, more or less granular, stoutly half-conic or triquetrous, strongly
undulate and twisted, very openly grooved from near the apex with acute edges,
usually carinate dorsally and ventrally, decurrent for about its own length, more
or less intruded into the green tissue, 5-6 cm. long, 8 mm. thick; teeth colored
like the spine, rather dull, 2.5-5 cm. apart, 5-10 mm. long, more or less curved,
especially downward, heavily triangular, abruptly dilated into fleshy prominences,
the intervening margin somewhat concave; inflorescence unknown.

At first glance this would seem to be a distinct species, but there
is some question whether the undulate and twisted form of the
terminal spine is a dependable character, rather than a freak. It
is doubtful whether the plant is really different from A. Seemanniana.

The following species were described by Berger almost im-
mediately after publication of the Trelease paper. All were based
upon young plants growing in Germany, sent from Guatemala by
W. Eichlam. No illustrations were published, and the identity of
the plants named is uncertain, although it is to be presumed that
they are synonymous with some of the species described above.

i STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 121

Agave Caroli-Schmidtii Berger, Agaven 199. 1915.

Based on plants sent from Guatemala by W. Eichlam and grown
at Erfurt, Germany.

Plants acaulescent, without offshoots; leaves erect, ovate-spatulate, strongly
narrowed toward the base, broadest above the middle, short-pointed, flat but with
somewhat incurved margins, smooth, or somewhat roughened near the base,
bright gray, almost white beneath; terminal spine 2.5-3 cm. long, short-decurrent,
somewhat intruded beneath into the green tissue, blackish brown, lustrous;
marginal teeth on elevated projections, spreading, the cusp narrow-deltoid, up-
curved or down-curved, blackish brown, 8-10 mm. long.

Agave Eichlami Berger, Agaven 200. 1915.

Described from a plant grown at Erfurt, Germany, sent from
Guatemala by W. Eichlam.

Plants acaulescent, without offshoots; leaves erect, lance-spatulate, acuminate,
concave above, bright gray, especially beneath, smooth or rough above, very
rough beneath; terminal spine straight, long-decurrent, roughened dorsally,
blackish brown, about 3 cm. long; marginal teeth with a lenticular base, the cusp
deltoid, curved downward or less frequently upward, 8-10 mm. long, blackish
brown, 5-8 mm. apart, the intervening margin usually straight, the spine bases
more rarely fleshy.

Berger described also a var. inter jecta (loc. cit.), with less decurrent
terminal spine.

Agave guatemalensis Berger, Agaven 201. 1915.

Type grown at Erfurt, Germany, sent from Guatemala by
W. Eichlam.

Plants acaulescent, without offshoots; leaves erect, more or less recurved from
the middle, thick-fleshy, ovate-spatulate, acute, concave above, more or less
gray-green, smooth, rough beneath, pale gray; terminal spine strong, soon becom-
ing gray, somewhat flexuous or straight, flat below and decurrent upon the blade,
about 3 cm. long, decurrent for about 2 cm.; marginal teeth with semilunate base,
the cusp gray, deltoid and straight or curved downward, about 8 mm. long, 1-2
cm. apart.

Agave Weingartii Berger, Agaven 200. 1915.

Type cultivated at Erfurt, Germany, sent from Guatemala by
W. Eichlam.

Plants acaulescent, the leaves not very dense, lanceolate or lance-spatulate,
rather thin, acuminate, concave above, usually smooth and apple-green shaded
with gray, beneath bright gray and mostly rough; terminal spine 3 cm. long, slender,
finely roughened, lustrous blackish brown, decurrent on the green tissue for 2-3
cm.; marginal teeth on the upper fourth or fifth of the leaf, the teeth 8-20 mm.
apart, deltoid from a lenticular base, less often on fleshy projections, blackish
brown, mostly 2-5 mm. long.

122 FIELDIANA: BOTANY, VOLUME 24

BOMAREA Mirbel

Scandent herbs, often with greatly elongate stems, glabrous or pubescent,
the roots slender, often bearing at the tip large tubers; leaves narrow or broad,
many-nerved, acute, abruptly short-petiolate; flowers large and showy, usually
red and yellow, often spotted, in simple or compound umbels; perianth funnel-
form, the segments distinct, those of the 2 series dissimilar; stamens attached to
the bases of the segments, about equaling them, the filaments filiform, the anthers
oblong, basifixed; ovary 3-celled, the ovules numerous, the style filiform, the
stigma trifid; capsule turbinate or subglobose, truncate at the apex, 6-costate,
tardily dehiscent; seeds subglobose, covered with a bright red, pulpy testa.

A large American genus, with probably one hundred or more
species, most of them Andean. One or two besides those listed here
occur in Costa Rica and Panama. In Costa Rica the name Papa
de venado is given to the plants. The watery tubers are said to be
edible but almost flavorless.

Rays of the umbel 1-flowered, usually without bractlets, very densely short-
villous B. acutifolia.

Rays of the umbel mostly 2-3-flowered, usually bracteolate, glabrate or sparsely
short- villous . . . . B. hirtella.

Bomarea acutifolia (Link & Otto) Herb. Amaryll. 112. 1837.
Alstroemeria acutifolia Link & Otto, Icon. 1: 5. pi. 29. 1820. Grana-
dillo de cante (fide Aguilar).

Damp forest or thickets in the mountains, 1,800-3,500 meters;
El Progreso; Zacapa; Jalapa; Sacatepe"quez; Chimaltenango; Solola;
Quiche" ; Huehuetenango; Quezaltenango; San Marcos. Mexico;
perhaps also in Costa Rica and Panama.

A small or large vine, the soft herbaceous stems glabrous or nearly so, bearing
many leaves; leaves ovate to lanceolate, mostly 5-15 cm. long, acuminate or
attenuate, on short slender petioles, obtuse or rounded at the base, paler beneath
and glabrous or sparsely pilose; rays of the umbel 5-30, 2-5 cm. long, densely
villous with brownish hairs, naked or bracteolate at or near the base; ovary
villous; perianth 2.5-4 cm. long, the outer segments oblong, orange-red with
green tip, or sometimes greenish throughout, the inner segments cuneate-spatulate,
clawed, orange-yellow and brick-red, truncate at the apex.

Like other species of the genus, this is a handsome and showy
plant because of its vividly colored flowers. When occurring in
quantity, it affords a fine display of color, but in Guatemala, at least
during the drier months, plants of the genus seldom are found in
profusion.

Bomarea hirtella (HBK.) Herb. Amaryll. 112. 1837. Alstroe-
meria hirtella HBK. Nov. Gen. & Sp. 1 : 284. 1815. B. miniata Kunth,

FIG. 23. Bomarea hirtella. A. Upper portion of flowering stem; X M-
B. Stigma and upper part of style; X 13. C. Outer perianth segments; X 2,
D. Inner perianth segments; X 2. E. Anther and upper portion of filament; X 13.

123

124 FIELDI ANA: BOTANY, VOLUME 24

Enum. PL 5: 792. 1850. Granadillo de monte (Guatemala, fide
Aguilar) ; Bate (San Ildefonso Ixtahuacan, Huehuetenango).

Damp or wet forests or thickets, 800-3,000 meters, rarely at
lower elevations; Pete"n (La Libertad); Alta Verapaz; Zacapa;
Chiquimula; Jalapa; reported from Santa Rosa; Guatemala; Sacate-
pe*quez; Chimaltenango; Solola; Huehuetenango; San Marcos.
Mexico; Honduras; perhaps farther southward into Costa Rica.

A small or rather large vine, the slender stems usually glabrous, sometimes
sparsely villous; leaves evidently petiolate, usually elongate-lanceolate, as much
as 16 cm. long and 1.5-2.5 cm. wide, acuminate or attenuate, usually abruptly
contracted and rounded at the base, glabrous above, paler beneath and often
hirtellous or villous; umbels usually large and with elongate rays, these glabrous
or sparsely short-villous, bearing one or more green ovate bracts and mostly
2-3 flowers, some of the rays only 1-flowered; ovary commonly sparsely short-
pilose or almost glabrous; perianth 3-3.5 cm. long, the outer segments pale red,
rose-pink, or salmon, the inner ones pale green or pale pink with dark brown
spots, spatulate; capsule 2-3 cm. broad, soon opening and exposing the bright red
seeds. (Fig. 23.)

By some authors B. miniata is recognized as a distinct species,
but the characters by which it is separable from B. hirtella are not
apparent to the writers. Guatemalan material of B. hirtella has
been variously referred to B. Caldasiana Herb., B. Caldasii (HBK.)
Stand., and B. ovata (Cav.) Mirb., species not known to occur in
Central America. This plant, like others of its genus found in
Central America, usually attains its best development not within
the forest but on bushes at the outer forest edge, where it is more or
less exposed to the sun.

CRINUM L.

Plants arising from large tunicated bulbs having many coats and usually an
elongate neck; leaves persistent, strap-shaped, often broad, succulent, the scape
solid; flowers umbellate, few or numerous, white or reddish, the pedicels short
or none; spathe valves 2, broad, membranous; perianth salverform or funnel-
form, the tube long and slender, often greatly elongate, straight or somewhat
curved, the segments linear or lanceolate, subequal; stamens inserted in the throat
of the perianth tube, the filaments long and filiform, the anthers linear, versatile;
ovary 3-celled, with few, sometimes only 2, ovules in each cell; style long and
filiform, the minute stigma capitate; fruit irregular in shape, tardily dehiscent;
seeds large, green and bulblike, with very thick endosperm.

About eighty species are listed, widely distributed in the tropics
of both hemispheres, but they are poorly known for the most part,
and the true number is very uncertain. One other Central American
species has been described from Panama.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 125

Perianth funnelform, the segments often equaling the tube, linear-lanceolate;

flowers distinctly pedicellate C. longifolium.

Perianth salverform, the segments usually much shorter than the tube; flowers
sessile.

Perianth segments narrowly linear C. cruentum.

Perianth segments linear-lanceolate, usually 1 cm. wide or more . . C. erubescens.

Crinum cruentum Ker in Lindl. Bot. Reg. pi. 171. 1817.

Seen in cultivation at Quirigua (Izabal) and elsewhere, and prob-
ably native along the north coast. Reported from Mexico; British
Honduras: Atlantic coast of Honduras; Costa Rica.

Bulbs large, with only a short neck, stoloniferous; leaves sometimes a meter
long, 4-7 cm. wide, dark green, gradually narrowed to the acute apex; scape stout,
somewhat compressed, 60-90 cm. long; flowers commonly 6-8, sessile and crowded,
the spathe valves lance-deltoid, 7 cm. long; perianth tube very slender, 10-20 cm.
long, 2-3 mm. thick in the dry state, the lobes pink or white, 7-13 cm. long, 3 mm.
wide; filaments red; anthers linear, about 12 mm. long.

Crinum erubescens Ait. Hort. Kew. 1: 413. 1789. Lirio
silvestre; Reina blanca, Reina rosa (fide Aguilar).

Wet soil or shallow water, usually at the margins of streams or
lakes, chiefly at low elevations, but collected as high as 1,800 meters,
perhaps only in cultivation; Alta Verapaz; Izabal; Santa Rosa;
Guatemala; Suchitepequez; Retalhuleu. Mexico to Panama; West
Indies; South America.

Bulbs ovoid, 7-10 cm. in diameter, with a short neck; leaves numerous, thin,
60-90 cm. long, 4-7 cm. wide, rather blunt at the apex, the margins slightly rough-
ened; scape 60 cm. high or more, stout; flowers 4-12 in the umbel, sessile or essen-
tially so, the spathe valves lance-deltoid, green, about 7 cm. long; perianth tube
very slender, 10-15 cm. long, 3-4 mm. thick in the dry state, the segments half
as long as the tube or shorter, linear-lanceolate, about 1 cm. wide, usually white
within and red-purple outside, sometimes wholly white; filaments bright red, at
least above; anthers linear, about 12 mm. long.

Called "lirio" in Salvador and doubtless also in Guatemala. It
is questionable whether the plant is native much above sea level,
but it is frequent in cultivation in many of the gardens of mountain
regions. It is particularly plentiful about Tactic, Alta Verapaz,
and has run wild in that region.

Crinum longifolium (L.) Thunb. Prodr. 39. 1794. Amaryllis
longifolia L. Sp. PI. 421. 1753. C. longiflorum Herb. Amaryll. 271.
1837. A. longifolia var. longiflora Ker, Bot. Reg. pi. 303. 1818.
Lirio; Reina de las flores; Reina.

Cultivated for ornament throughout the lowlands and up to the
central highlands, at 1,500 meters or more; escaped from cultivation

126 FIELDIANA: BOTANY, VOLUME 24

and more or less naturalized in some parts of the lowlands, at least
about the sites of former dwellings. Native of South Africa, but
now generally cultivated in tropical regions.

Plants sometimes forming dense clumps, the bulb 7-10 cm. thick, narrowed
into a very long and thick neck; leaves few or numerous, ensiform, acute or acumi-
nate, sometimes 1.5 meters long but usually shorter, recurved, 4-8 cm. wide,
rather pale, often slightly roughened on the margins; flowers 6-8 or more, on short
stout pedicels, the spathe valves deltoid, 7-10 cm. long; perianth tube greenish,
10-12 cm. long, the segments linear-lanceolate, usually white within, rose-purple
outside, mostly 1-1.5 cm. wide, acute, often connivent; filaments much shorter
than the perianth lobes, the anthers linear, about 12 mm. long.

This species is much planted in the gardens of the lowlands,
and it thrives with little or no attention. It is planted frequently
also in the uplands, especially in parks. The long bulbs, projecting
far above the ground, are conspicuous, as are also the large broad
leaves. The flowers are rather showy, but scarcely handsome enough
to warrant the name of "reina de las flores" so often given in Guate-
mala to the plant.

CURCULIGO Gaertner

Plants with short, sometimes tuberous or corm-like rhizomes covered with
reticulate fibers; leaves radical, usually elongate-lanceolate, plicate- veined, small
and grasslike or sometimes large; flowers spicate, the spikes sessile among the
leaves and few-flowered or sometimes long-pedunculate, the bracts linear, the
whole inflorescence often villous; perianth 6-parted, borne at the apex of a very
long and slender, tube-like elongation of the ovary, this resembling a corolla tube,
the perianth segments subequal, spreading; stamens 6, attached at the base of
the segments, the filaments short, the anthers linear, erect, shorter than the
perianth; ovary 3-celled, usually produced into a long tube-like beak, the style
short, columnar, the 3 stigmas oblong, erect; ovules 2-many in each cell; fruit
somewhat succulent, indehiscent; seeds subglobose, black.

About a dozen species in the tropics of America, Asia, Africa,
and Australia. A single species is native in Central America.

Curculigo scorzoneraefolia (Lam.) J. G. Baker, Journ. Linn.
Soc. Bot. 17: 124. 1878. Hypoxis scorzoneraefolia Lam. Encycl.
3: 183. 1789.

Mostly in savannas or other grassy places, chiefly at 1,000 meters
or less ; Pete"n ; Alta Verapaz ; Santa Rosa ; Huehuetenango. Southern
Mexico; Honduras; Nicaragua; Costa Rica; Panama; West Indies;
South America.

Plants small and low, with oblong corms; leaves few, usually 2-3, erect, linear-
lanceolate, mostly 10-30 cm. long and 5-20 mm. wide, long-attenuate, narrowed
downward into a distinct slender petiole, thin, many-nerved, sparsely pilose;

FIG. 24. Curculigo scorzoneraefolia. A. Habit of plant; X %. B. Flower
with bracts; X 3.

127

128 FIELDIANA: BOTANY, VOLUME 24

flowers sessile, only 1 developing at a time, the false perianth tube almost filiform,
usually 1.5-2.5 cm. long, appressed-pilose; perianth yellow, the segments elliptic-
lanceolate, 1-1.5 cm. long, acute or acuminate, sparsely pilose outside. (Fig. 24.)

Curculigo capitulata (Lour.) Kuntze, Rev. Gen. 703. 1891.
Leucojum capitulatum Lour. Fl. Cochinch. 199. 1790. C. recurvata
Dryand. in Ait. Hort. Kew. ed. 2. 2: 253. 1811.

This species, native of southeastern Asia and tropical Australia,
is sometimes cultivated for ornament in Guatemala and in other
parts of Central America. It has a tendency to spread and establish
itself, and is rather well naturalized in the cemetery at Retalhuleu
and in the Jardin Botanico, Guatemala.

A coarse plant of somewhat palm-like appearance, the bright yellow, starlike
flowers in dense short-pedunculate clusters at the bases of the leaves. The erect
thin leaves are long-petiolate and as much as a meter tall, and 4-12 cm. wide.

In Salvador the plant is called "coquillo."

EUCHARIS Planch. & Linden

Plants arising from a tunicated bulb; leaves few, broad, abruptly contracted
at the base into a slender petiole; flowers few, large and showy, pure white, umbel-
late; outer spathe valves 2, ovate-lanceolate, the inner numerous, linear; perianth
tube cylindric, with a dilated throat, somewhat curved; perianth segments equal,
oblong or ovate, spreading; stamens inserted at the throat of the perianth tube,
shorter than the segments, the filaments variously appendaged downward, the
anthers versatile; ovary globose, 3-celled; ovules many in each cell, rarely 2,
superposed; style filiform, the stigma capitate, 3-lobate; capsule deeply 3-lobate,
tardily dehiscent; seeds large, bulbiform, sometimes solitary.

Half a dozen species, chiefly in the northern Andes. One species
is native in Costa Rica.

Eucharis grandiflora Planch. & Linden, Fl. Serres I. 9: 255.
1853-54. Eucaria.

Planted frequently in gardens for ornament, especially in the
lowlands, but also as high as Guatemala (1,500 meters). Native
of the Andes of Colombia.

Bulbs globose, about 5 cm. in diameter; leaves 2-4, the petiole 15-30 cm. long;
leaf blades oblong to elliptic or oval, often 30 cm. long, 10-15 cm. wide, abruptly
short-acuminate, rounded to attenuate at the base; scape terete, 35-60 cm. tall;
flowers 4-6 in the umbel, on pedicels 1-2.5 cm. long; outer spathe valves ovate-
lanceolate; perianth tube somewhat curved, 5 cm. long, the throat 12-15 mm.
broad; perianth lobes oblong-ovate, 3.5-5 cm. long, pure white, obtuse; stamen
cup of 6 quadrate connate segments, with a short lanceolate filament at the center
of the base of each; ovules about 20 in each cell.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 129

The plant is a handsome one because of its pure white flowers,
but it appears to be a rather shy bloomer.

FURCRAEA Ventenat

Reference: William Trelease, Trans. Acad. Sci. St. Louis 23: 147-
150. 1915.

Large plants with an erect, hypogaean or subterranean caudex; leaves densely
crowded at the apex of the caudex, long and narrow, thin or rather thick, the
margins spinose-dentate or subentire; panicles large, terminal, usually pyramidal,
the flowers pedicellate in the axils of bracts, solitary or fasciculate, white or
greenish white, often replaced by bulblets; perianth rotate, cleft almost to the
base, the segments equal, oval-oblong; stamens affixed to the bases of the perianth
segments and shorter than them, the filaments dilated below the middle, subulate
above, the anthers linear-oblong, dorsifixed, shallowly bilobate at the base;
ovary oblong, 3-celled, usually short-rostrate at the apex, the style columnar,
thickened below the middle, the stigma small, obscurely trilobate; ovules numerous
in each cell, biseriate; capsule oblong or ovoid, loculicidally 3-valvate; seeds
numerous, flattened.

About fifteen species, in Mexico, Central America, and western
South America. The generic name has often been written Fourcroya
or Furcroea. The leaves, as in Agave, contain a tough fiber, which
is sometimes used commercially as well as locally. The fiber of the
Brazilian F. gigantea Vent, has been exploited under the name of
Mauritius hemp. Some of the Central American species reproduce
principally by bulblets that replace the flowers. The bulblets often
are found in great quantities beneath the plants, where under
favorable conditions they take root and grow rapidly, soon forming
a dense thicket of large plants. Plants so grown are said sometimes
to flower in only three years. As with the Agaves, the species of
Furcraea are poorly understood, and the following treatment is far
from satisfactory, although based upon the fullest data now available.
Unless flowers are available, it often is difficult to determine whether
a certain plant is a Furcraea or an Agave.

Leaves without spine-like teeth, the margins somewhat roughened, the leaves

thin and flexible, scarcely fleshy; plants usually with short or elongate trunks.

Plants treelike, the trunk becoming sometimes 15 meters tall; flowers pubescent.

F. longaeva.
Plants acaulescent or with a trunk 2 meters tall or less; flowers glabrous.

F. quicheensis.

Leaves with large or small, spine-like teeth, the margins between the teeth not
roughened, smooth; plants usually acaulescent.

Leaves without teeth on the upper third or two-thirds F. samalana.

Leaves with spine-like teeth nearly or quite throughout.

130 FIELDIANA: BOTANY, VOLUME 24

Leaves flattish, not very concave, the marginal teeth red or brown, rather

distant F. guatemalensis.

Leaves very concave, the marginal teeth blackish, close together.

F. melanodonta.

Furcraea guatemalensis Trelease, Trans. Acad. Sci. St. Louis
23: 149. pis. 32-34. 1915. Maguey; Icaj (Coban, Quecchi); Mecate
(Huehuetenango) .

Open hillsides, dry rocky hills, sometimes in pine-oak forest,
probably most often seen in hedges or fencerows, 900-2,300 meters;
Alta Verapaz (perhaps only in cultivation); Zacapa; Jalapa; Santa
Rosa; Guatemala (type from Guarda Viejo, Trelease 23); Sacate-
pe"quez; Chimaltenango; Huehuetenango; Quezaltenango. Salvador,
but probably only in cultivation.

Plants acaulescent or practically so; leaves lanceolate or sword-shaped, 2
meters long or less, 10-15 cm. wide, dull green above and grayish beneath, or
sometimes rather bright green, acute, smooth or nearly so, moderately concave or
often almost flat; terminal spine of the leaf short and stout, awl-shaped, the
marginal teeth red-brown or paler, in age often chestnut, upcurved and com-
pressed, 1-3 cm. apart, about 3 mm. long, often with a very broad base; inflores-
cence large, 2-5 meters tall, openly paniculate, glabrous or nearly so; perianth
pale green or greenish white, about 3 cm. long, the segments oblong-elliptic,
acutish, about 1 cm. wide; capsules globose-cubical, short-stipitate or almost
sessile, about 5 cm. long and broad; seeds about 20 mm. in greatest breadth;
bulblets often abundant, globose-ovoid, with dark green, lustrous scales. (Fig. 25.)

This species is abundant in hedgerows in the vicinity of Guate-
mala as well as in many other parts of the central and western
departments. No data are available in general regarding use made
of the leaves, but doubtless fiber is extracted from them. In Alta
Verapaz, however, it seems that this species has been introduced
and planted rather extensively in some places. Apparently it is
planted with Agave sisalana, and perhaps no distinction is made
between the two.

Furcraea longaeva Zucc. Acad. Caes. Leop. Carol. 16, pt. 2:
665. 1833; Trelease, Trans. Acad. Sci. St. Louis 23: pi. 28.

Huehuetenango (Villa Cyrnos, Cap Martin near Nenton, May 17,
1909, herb, of Alwin Berger). Southern Mexico.

The Guatemala collection cited was found in the United States
National Herbarium and is the only herbarium material examined
by the authors. Skinner reported it for Guatemala and his drawing
accompanying one of his orchid plates is presumed to represent
this species.

FIG. 25. Furcraea gualemalensis. A. Flower, with parts in position; X !%
B. Flower opened to show position and arrangement of parts with ovary opened;
X 1H.

131

132 FIELD IANA: BOTANY, VOLUME 24

Arboreous, with a tall simple trunk sometimes 15 meters tall, the old and dead
leaves reflexed and long-persistent, the younger leaves erect or spreading, grayish,
sword-shaped, concave, subacuminate, as much as 2 meters long and 8-15 cm. wide,
minutely rough-margined; inflorescence to 5 meters in height, broadly conic,
short-stalked; perianth 2.3-3 cm. long, the outer segments elliptic-oblong, acutish
or subacute, 8-9 mm. wide, the inner segments ovate or elliptic-ovate, obtuse to
subacute, 12-15 mm. wide; ovary 2-2.5 cm. long, slightly exceeding the perianth;
capsule oblong, narrowed to the base; seeds about 6 mm. long and 4 mm. wide.

The pubescent ovary and outer surface of the perianth segments
are noteworthy characteristics of this species.

Furcraea melanodonta Trelease, Trans. Acad. Sci. St. Louis
23: 150. pi. 35. 1915. Maguey.

Dry rocky hillsides or in pine-oak forest, mostly at 1,200-1,800
meters; Baja Verapaz; El Progreso (type from Cruz, Trelease 26);
Chimaltenango; endemic.

Plants essentially acaulescent, the caudex in exposed places sometimes a
meter high; leaves sword-shaped, grayish or bluish or sometimes rather bright
green, acute, sometimes slightly roughened, 1.5 meters long or less, 7-8 cm. wide,
rather thin and almost flat or often concave; terminal spine short and stout,
blunt, grooved at the base, slightly decurrent, the marginal teeth blackish or
chestnut, 1-2 cm. apart, 5 mm. long or less, strongly upcurved, broad-based and
decurrent on fleshy elevations; inflorescence and capsules unknown.

It is doubtful whether this can be separated from F. guatemalensis,
for the characters used originally to separate the two seem variable
and not dependable.

Furcraea quicheensis Trelease, Trans. Acad. Sci. St. Louis 23:
148. pi. 29. 1915. Mecate; Mecatle; Cheche; Mecatl; Maguey.

Open mountain slopes, often on exposed white-sand mountain
sides, sometimes in oak forest, most often, perhaps, seen in fence-
rows, and much planted in some regions for hedges, chiefly at 2,400-
3,300 meters; Solola; Quiche" (type collected between Quiche" and
Totonicapan, 0. F. Cook in May, 1906); Huehuetenango; Totoni-
capan; Quezaltenango; San Marcos. Probably in Chiapas.

Plants acaulescent at first, but in age developing a thick naked trunk, this
rarely more than 1.5 meters tall, simple or sometimes with a few branches, densely
leafy at the apex; leaves very glaucous and whitish, thin and soft, in texture re-
sembling those of Yucca filamentosa, not fleshy, linear-lanceolate, 1.2 meters long
or less, mostly 6-10 cm. wide, long-attenuate, spreading or reflexed, the yellowish
margins minutely and remotely denticulate and rough to the touch, the tip of the
leaf narrowly rounded and blunt, without a terminal spine; panicles 2-5 meters
tall, rather narrow, the flowers pedicellate, the bracts much exceeding the pedicels;
these, like the peduncles, often red; perianth pale green, 3-4 cm. long, the segments

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 133

suberect or somewhat spreading; anthers pale green; ovary 2.5-3 cm. long, often
purplish red; capsule about 6 cm. long and 3 cm. broad, contracted at the base,
short-beaked, somewhat lustrous; seeds thin, black and lustrous, about 10 mm. long
and 6 mm. wide.

This is in some respects one of the most interesting of Guatemalan
plants, limited in its distribution to the highlands, where it often
forms a conspicuous feature of the landscape. Westward from
Guatemala, the plant is first seen in the bleak mountains of the
Los Encuentros region, and from there on westward it may be
observed here and there. It is most abundant in the highlands of
Huehuetenango, especially above Chiantla up to the rim of the
alpine region of Los Cuchumatanes, and in certain parts of Quezal-
tenango and San Marcos. In general appearance Furcraea qui-
cheensis suggests a Yucca more than one of the Agaves, because
of its habit and its exceedingly thin pale leaves. The pale foliage
makes the plant easily recognizable even from a long distance in
the mountains. It is hard to determine in just which regions the
plant is native, for most of the individuals occur in fencerows or
hedges, but their absence in adjoining fields probably is accounted
for by the fact that the Indians have removed them to the field
borders in order to clear the soil for cultivation, a clearing that in
most regions took place centuries ago. In some parts of the white-
sand mountains of San Marcos it seems that the Furcraea has been
planted to prevent drifting of the sand that is so hard to control
in these desolate regions. The plant is a very decorative and hand-
some one. The short trunks sometimes bear at the apex a number
of perhaps adventitious buds that develop into dense clusters of
leaves. The young leaves are at first cream-colored and remain so
for some time, or at least until fully exposed to the sun. In this state
they are much used for decorations, often like ribbon, to make bows
for trimming coronas or wreaths, or to make crosses for Palm Sunday,
like those of palm leaves used in the United States. For this purpose
the leaves are taken by cargadores from the Occidente even to
Guatemala City. The senior author saw quantities of the pale
leaves used in decorating arches across the road between San Marcos
and Quezaltenango on the occasion of the President's visit in January,
1941. The name mecatl is pure Nahuatl, and is the word from
which the term mecate, now applied in many regions of Mexico
and Central America to string of any kind, is derived. Strips of
the leaves of this Furcraea are much used in the Occidente as a
substitute for twine, and large bunches for this purpose were noted
on sale in the market of Momostenango.

134 FIELDIANA: BOTANY, VOLUME 24

Furcraea samalana Trelease, Trans. Acad. Sci. St. Louis 23:
149. pis. 30, 81. 1915. Maguey.

Brushy or rocky slopes, often in moist thickets but more often
in dry places, ascending from sea level to about 1,650 meters, only
in the Occidente; Suchitepequez; Quezaltenango (type collected in
the barranco of Rio Samala between Zunil and Santa Maria de
Jesus, Trelease 20). Salvador, perhaps only in cultivation.

Plants usually acaulescent, said to have sometimes a trunk 2 meters high, but
this probably very unusual; leaves sword-shaped, green or slightly grayish, mostly
1-2 meters long and 10-15 cm. wide, acute, almost smooth, with flat or outcurved
margins, rather thin; terminal spine very short and small, usually abortive, the
marginal teeth red-brown, usually none on the upper part of the leaf, 1-6 cm.
apart, 4 mm. long or smaller, mostly upcurved, decurrent on low fleshy elevations;
panicles 3-5 meters tall, narrow; perianth 2.5-3 cm. long, the segments greenish
white with a green central stripe; flowers often replaced by large bulblets, these
ovoid, with dull gray-green scales, leafy-tufted at the apex.

Called "magueyon" and "magueyon macho" in Salvador, where
the leaves are said to be an important source of fiber. This is
perhaps the species collected in Quezaltenango by Warscewicz and
reported by Hemsley under the name Furcraea Selloa K. Koch.

HIPPEASTRUM Herb. Amaryllis

Plants arising from tunicated bulbs; leaves linear or strap-shaped; scape
hollow, bearing an umbel of 2 or more flowers, these pedicellate, the flowers red
or whitish; perianth funnelform, commonly declinate, the tube usually short,
rarely long, the segments subequal or the lowest one of the inner series narrower;
stamens inserted in the throat of the perianth tube, somewhat declinate, the
filaments filiform, the anthers linear or narrowly oblong, versatile; ovary 3-celled;
ovules many in each cell, superposed; style long, declinate, the stigma capitate or
3-fid; capsule globose, loculicidally 3-valvate; seeds usually compressed, with a
thin black testa.

About forty species, all apparently native of South America.
Because of their handsome large flowers, many of the species have
long been favorites in cultivation, and many fine forms with excep-
tionally large and beautifully colored flowers have been developed
by hybridization or otherwise. Some of these "improved" forms
may be seen in cultivation in Guatemala, especially about Coban.

Perianth tube very slender, commonly 7-12 cm. long H. solandriflorum.

Perianth tube funnelform, usually 2.5 cm. long or shorter H. puniceum.

Hippeastrum puniceum (Lam.) Urban, Symb. Antill. 4: 151.
1903. Amaryllis punicea Lam. Encycl. 1: 122. 1783. A. equestris

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 135

Ait. Hort. Kew. 1: 417. 1789. H. equestre Herb. App. 31. 1821.
Amarilis; Azucena roja.

Cultivated generally in gardens, often in large quantities, for
ornament, in the lowlands and far up into the highlands; one of
the most popular garden flowers of Guatemala and other parts of
Central America. Native of South America, but cultivated in
almost all regions of the earth, either out of doors or as a house plant.

Bulbs globose or ovoid, stoloniferous, 4-5 cm. long or larger, the outer coats
brown; leaves strap-shaped, 30-60 cm. long, 2.5-5 cm. wide, gradually narrowed
to the apex; scape terete, pale, 40-60 cm. high, the umbel usually 2-4-flowered;
spathe valves lanceolate; pedicels 3-7 cm. long; perianth commonly 10 cm. long
or larger, usually bright red or orange-red, the segments obovate or elliptic-obovate;
stamens shorter than the perianth.

Called "amapola" and "bucaro" in Salvador. Large quantities
of the flowers are offered for sale in Guatemalan markets. The
flowers are most plentiful about March, toward the end of the dry
season, and they often appear when no leaves have yet developed.
The plant frequently escapes in the lowlands, and sometimes is
found in thickets, especially about Coban and in the Pacific boca-
costa. It grows luxuriantly in the lowlands where it receives little
or no attention, in great contrast to the carefully tended house
plants of the North.

Hippeastrum solandriflorum Herb. App. 31. 1821. Amarilis.

Cultivated commonly for ornament, especially in the Coban
region. Native from Colombia to the Guianas and northern Brazil.

Bulbs ovoid, 7-10 cm. in diameter, with a short neck; leaves strap-shaped,
35-50 cm. long, 2.5-3.5 cm. wide; scape somewhat 2-edged, 35-60 cm. tall or more,
the umbel 2-4-flowered, the pedicels 2.5-3.5 cm. long; spathe valves lanceolate,
5-7 cm. long; perianth narrowly funnelform, the limb 15-25 cm. long, the slender
greenish tube commonly 7-12 cm. long; perianth segments greenish white, usually
with obscure darker stripes and often tinged, especially outside, with pink or
purple; anthers linear-oblong.

This is one of the favorite garden flowers of Coban, where it is
often grown in abundance for market. Large quantities of the hand-
some flowers appear in the Coban market during Holy Week, when
they are carried as decorations in the many religious processions,
and appear upon the altars of the churches.

HYPOXIS L.

Reference: A. Brackett, Revision of the American species of
Hypoxis, Rhodora 25: 120-147. 1923.

136 FIELDIANA: BOTANY, VOLUME 24

Small plants, often pilose, with corms or short rhizomes; leaves linear, grass-
like; scapes very slender and few-flowered; perianth 6-parted, the segments equal
or nearly so, distinct, withering-persistent; stamens inserted on the bases of the
perianth segments, the filaments short, the anthers erect, sagittate or entire;
ovary 3-celled, the style short, with 3 erect stigmas; ovules numerous, in 2 rows
in each cell; capsule thin- walled, circumscissile below the apex; seeds subglobose,
tuber culate or echinate.

About fifty species, in temperate and tropical regions of both
hemispheres. No other species are known from Central America.

Mature seeds black; outer coat of the seed firm and close, covered with crowded
low obtuse tubercles H. decumbens.

Mature seeds brown; outer coat of the seed of loose texture, rugose and con-
tracted into rather remote minute tubercles H. rugosperma.

Hypoxis decumbens L. PI. Jam. Pugill. 11. 1759. H. decumbens
var. major Seub. in Mart. Fl. Bras. 3, pt. 1: 51. pi. 7, fig. 1. 1847.
H. racemosa Bonn. Smith, Bot. Gaz. 14 : 30. 1889 (type from Coban,
Alta Verapaz, Turckheim 33). Cebollin de coyote (fide Aguilar).

Mostly in moist open grassy places, sometimes on dry rocky
hillsides or in pine forest, 3,500 meters or lower; Pete*n; Alta Verapaz;
Baja Verapaz; Zacapa; Jalapa; Guatemala; Sacatepe'quez; Chimal-
tenango; Solola; Quiche"; Huehuetenango. Mexico to British Hon-
duras; Honduras; Costa Rica; Panama; West Indies; South America.

Plants arising from cylindric or ellipsoid corms 1-2 cm. thick; leaf sheaths
not separating into fibers, the blades 10-40 cm. long, 2-12 mm. wide, sparsely
pilose or glabrate; peduncles filiform, 2-12 cm. long, 1-4-flowered; perianth
segments lanceolate, acute, 4-10 mm. long; fruiting pedicels 2 cm. long or less;
capsule clavate, cylindric, or oblong-ellipsoid, usually densely pilose, 6-17 mm.
long; seeds black, dull or slightly lustrous, covered with low tubercles. (Fig. 26.)

Var. major (of which H. racemosa is a synonym) is a more robust
form with relatively wide leaves. A collection from Huehuetenango
(Standley 82627) is very close to H. hirsuta (L.) Cov. in its seed
characters. It may be remarked that the American species of
Hypoxis are all closely related and separated by minute characters.

Hypoxis rugosperma Brackett, Rhodora 25: 142. fig. 11. 1923.

Santa Rosa, Dept. Santa Rosa, 900 meters, Heyde & Lux 2934
(cited by Brackett). Known otherwise only from the type, collected
at Guadalajara, Jalisco, Mexico.

Corms globose or subcylindric, 6-15 mm. thick; outer leaves separating into
tufts of brown fibers, the leaves 27 cm. long or less, 2-5 mm. wide, pilose; peduncles
pilose, 4-12 cm. long; ovary and capsule densely pilose; perianth segments narrowly
elliptic or lanceolate, acute, 3-7 mm. long; capsule subcylindric or ellipsoid, 6-9

FIG. 26. Hypoxis decumbens. A. Habit of plant; X %. B. Flower; X 3.
C. Stamen; X 15. D. Fruiting stalk with bracts and dehiscent capsule; X 3.
E. Seed; X 30.

137

138 FIELDIANA: BOTANY, VOLUME 24

mm. long; seeds brown, the testa loose, irregularly covered with confluent, thin,
slightly jagged crests.

We have seen no material of this species.

Narcissus. A species of Narcissus with pale yellow "single"
flowers was seen in bloom in March, 1939, in an Indian garden
near San Marcos, where there were numerous plants. Undoubtedly
it is cultivated elsewhere in the same region, since garden plants
are passed on from one grower to another. No doubt the plant has
been in cultivation a long time in this isolated locality, where it
was surprising to find it, since the so-called "spring bulbs," such
as tulips and hyacinths, are believed not to thrive anywhere in
Central America. If imported, the bulbs produce flowers but once,
ordinarily, and often fail to bloom at all.

PANCRATIUM L. Spider Lily

Plants with tunicated bulbs; leaves linear or usually strap-shaped, not con-
tracted into a petiole; scapes solid; flowers usually pure white, few or many in
an umbel, fragrant, commonly sessile; outer spathe valves 2, lanceolate; perianth
tube elongate, dilated above, the segments equal, spreading, linear or lanceolate;
filaments inserted in the throat of the perianth tube, united to form a cup, usually
with an entire or bifid process between each 2 filaments; anthers linear, versatile;
ovary 3-celled, the ovules 2-many in each cell; style long, filiform, the stigma
capitate; capsule usually large and subglobose, often bulb-like and somewhat
fleshy, finally loculicidally 3-valvate; seeds angulate, usually black.

About fifty species, in the tropics of both hemispheres. By most
recent authors the American species have been treated as a distinct
genus, Hymenocallis, based upon the number of ovules. This
character, however, does not hold, and there being left for segrega-
tion of the two genera nothing but their geographic distribution, it
seems necessary to unite them, as was done by most of the older
workers upon American plants.

Perianth segments lanceolate, mostly 12 mm. wide or more; staminal cup but
little shorter than the perianth segments, its lobes large, broadly rounded or

ovate P. narcissiflorum.

Perianth segments linear, commonly 2-6 mm. wide; staminal cup less than one-
third as long as the perianth segments, its lobes, if any, small and incon-
spicuous.

Leaves strap-shaped, not contracted into a petiole P. littorale.

Leaves contracted into a distinct narrow petiole.

Perianth tube about 6 cm. long, the segments slightly longer than the tube.

P. Skinnerianum.
Perianth tube about 20 cm. long, the segments shorter than the tube.

P. guatemalense.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 139

Hymenocallis bistubata Herb. Bot. Reg. 1844: Misc. 53. 1844.
This was based upon plants sent by Hartweg to England and culti-
vated there. It may have been collected in Guatemala, or perhaps
in some other country in which Hartweg collected. A similar but
apparently distinct species has been collected in Salvador.

Pancratium guatemalense Standl. & Steyerm. Field Mus.
Bot. 23: 38. 1944.

Wet soil, mostly along the borders of streams in mixed forest,
900 meters or lower; endemic; Izabal (type collected between
Escobas and Montana Escobas, across the bay from Puerto Barrios,
Steyermark 39288).

Bulbs subglobose or broadly ovoid, 10-12 cm. in diameter; leaves at anthesis
usually 15-16, mostly distichous, gradually narrowed below into a petiole, 48-90
cm. long, 11-19 cm. wide, broadly elliptic-oblong or broadly oblanceolate, abruptly
caudate at the apex; scapes 75-100 cm. tall, the flowers about 20, sessile; valves
of the spathe membranaceous, reflexed in anthesis; flowers fragrant, 30-40 cm.
long, the tube 16-20 cm. long, 7 mm. thick; perianth segments white, linear-
lanceolate, recurved and twisted, 10-12 cm. long, 8 mm. wide, acute; stamen cup
2 cm. long, dentate; filaments white, green near the apex, 6 cm. long; anthers
linear, yellow, 2.5 cm. long; style 25 cm. long; ovary oblong, 2.5-3 cm. long, ob-
tusely angulate; ovules 4-5 in each cell.

The plant is a handsome and showy one. Bulbs taken to the
Garfield Park Conservatory in Chicago produced luxuriant plants
that bloomed profusely.

Pancratium littorale Jacq. Sel. Stirp. Amer. 99. pi. 179, f. 94.
1766. Hymenocallis littoralis Salisb. Trans. Hort. Soc. Lond. 1: 338.
1812. H. tenuiflora Herb. Amaryll. 44. 1837. H. panamensis
Lindl. Bot. Reg. 1841: Misc. 67. 1841. H. insignis Kunth, Enum.
PI. 5: 675. 1850. Lino; Lino zac (Pete"n fide Lundell).

Marshy savannas, often abundant in salt marshes and swamps,
or at the edges of streams, frequently at the borders of mangrove
swamps, also frequent in cultivation inland, and sometimes escaping
to thickets and waste ground, plentiful along both coasts, at or little
above sea level; Pete"n; Alta Verapaz; Izabal; El Progreso; Santa
Rosa; Escuintla; Suchitepequez ; Retalhuleu; Huehuetenango; San
Marcos. Mexico to British Honduras and Panama; South America;
naturalized in some parts of the Old World.

Bulbs 7-10 cm. thick; leaves few, suberect or recurved, bright green, strap-
shaped or ensiform, acute, not narrowed into a petiole, usually 60-75 cm. long and
2.5-4 cm. wide; scape somewhat compressed and 2-edged, about 60 cm. tall;
flowers usually 4-8 in each umbel, sessile, the outer spathe valves deltoid, 5-7 cm.

FIG. 27. Pancratium littorale. A. Habit of plant; X V- B. Inflorescence;
X K-

140

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 141

long; perianth tube 10-25 cm. long, very slender, the linear segments recurved,
about 10 cm. long; staminal cup broadly funnelform, about 2 cm. long, irregularly
dentate along the margin; anthers linear, 12 mm. long; ovules 4-6 in each cell;
capsule rather large, green, resembling a small bulb. (Fig. 27.)

Called "lirio monteY' in Salvador. The plant is a handsome
one, like other members of the genus, and on this account is often
planted in gardens. In the salt-water marshes along the coasts it
sometimes forms large colonies.

Pancratium narcissiflorum Jacq. Fragm. Bot. pi. 138. 1809.
P. calathinum Ker, Bot. Reg. 3: pi. 215. 1817. Ismene calathina
Herb. Bot. Reg. App. 46. 1821. Hymenocallis calathina Nichols.
Diet. Gard. 2: 165. 1885. H. narcissiflora Macbride, Field Mus.
Bot. 11: 11. 1931. Azucena blanca.

Cultivated for ornament in gardens at Tactic, Alta Verapaz,
Guatemala, and probably elsewhere. Supposed to be native in the
Andes of Peru and Bolivia; said to have been introduced into cultiva-
tion in Europe in 1794.

Plants rather coarse and stout, the bulbs globose, with a cylindric neck;
leaves few, subdistichous, bright green, strap-shaped, about 60 cm. long, 3-5 cm.
wide, acute or subobtuse; scape about 60 cm. tall, 2-edged and somewhat com-
pressed; flowers 2-5, sessile, the outer spathe valves large, ovate; perianth tube
bright green, dilated and funnelform above, 7-10 cm. long, the segments pure
white within, lanceolate, about equaling the tube; staminal cup obconic, white
striped with green, 5 cm. long, the large lobes rounded or broadly ovate, dentate;
anthers linear, 12 mm. long; style long-exserted, very slender.

The flowers are exceptionally beautiful, and have more substance
than those of other species because of the broad perianth segments
and large staminal cup. The plant was noted in cultivation only
at Tactic, and was said to be uncommon there. Strangely enough,
J. G. Baker (Handb. Amaryll. 129. 1888) states that he had seen
Guatemalan specimens, probably from cultivation, and these may
well have come likewise from Alta Verapaz.

Pancratium Skinnerianum (Herb.) Standl. & Steyerm. Field
Mus. Bot. 23: 39. 1944. Hymenocallis Skinneriana Herb. Bot. Reg.
1843: Misc. 45. 1843.

Collected in Guatemala by Skinner, the locality unknown;
introduced into cultivation in England.

Bulbs of medium size; leaves oblong, acute, about 30 cm. long and 12 cm. wide,
contracted at the base into a petiole; flowers about 6, sessile, the spathe valves
lanceolate, 3.5 cm. long; perianth tube about 6 cm. long, the segments slightly
longer; staminal cup narrowly funnelform, dentate between the filaments, the

142 FIELDIANA: BOTANY, VOLUME 24

free portion of the filament scarcely longer than the cup; anthers linear; style
longer than the stamens.

Apparently a rare plant, not obtained by any recent collectors.
POLIANTHES L. Tuberose

Rootstock tuber-bearing; leaves mostly basal, few, lanceolate or linear,
elongate, succulent but thin, the cauline leaves mostly reduced and bract-like;
flowers white or reddish, mostly in pairs and racemose, the pedicels articulate at
the apex; perianth with a long narrow funnelform tube, more or less curved, the
segments short, subequal; stamens inserted about the middle of the tube, included,
the filaments filiform, short, the anthers erect, linear, dorsifixed; ovary 3-celled,
free at the apex, the ovules numerous, superposed; style filiform, the 3 stigmas
ovate, falcate; capsule crowned by the persistent perianth; seeds flat, with a lax
testa.

About a dozen species have been described, all natives of Mexico.

Polianthes tuberosa L. Sp. PI. 453. 1753. Nardo.

Common in gardens almost throughout Guatemala, and grown
in large quantities for the larger markets. Native of Mexico, but
probably unknown in a wild state.

Plants tall and strict, usually 60-90 cm. high, the basal leaves linear, pale green,
30-45 cm. long, deeply channeled in the lower half, spotted with dark brown on
the back; cauline leaves usually 8-12, smaller and shorter than the basal ones;
flowers double, waxy white, forming a long lax spike; perianth 3.5-6 cm. long,
the segments oblong-lanceolate, spreading, much shorter than the tube.

The tuberose is one of the favorite garden flowers of Guatemala,
perhaps because of its intense but often rather cloying fragrance.
Large bunches of the flower spikes are generally to be seen in the
Guatemala market, where the flowers are colored with synthetic
dyes in vivid shades of blue, yellow, pink, and green. The plant
presumably has been in cultivation in Mexican gardens, at least,
for many centuries. It is often grown in the United States.

Sprekelia formosissima (L.) Herb. App. 35. 1821. Amaryllis
formosissima L. Sp. PI. 293. 1753. This plant has been noted by
the writers only in cultivation in Coban. J. G. Baker reports it
as a native of Mexico and Guatemala. It is known to be native in
Mexico, but there is no evidence that it is native also in Guatemala.
This may well be the case, however, for the blooming period is short,
and the plant on that account could easily be missed. It is a bulbous
plant somewhat suggestive of Hippeastrum, with a single large
horizontal flower at the apex of the scape. The perianth is about

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 143

10 cm. long, with narrow, bright crimson segments. Leaves are
absent when the flowers open.

ZEPHYRANTHES Herb.

Plants with tunicated bulbs; leaves mostly linear and almost grasslike, con-
temporary with the flowers or developing after them; scape elongate, slender,
hollow, bearing a single flower, this red, white, or yellow; spathe membranous,
tubular below, bilobate above; perianth funnelform, erect or slightly inclined, the
tube short or elongate, the segments subequal; stamens inserted in the throat or
the middle of the perianth tube, slightly declinate, somewhat unequal; filaments
filiform, the anthers linear-oblong, versatile; ovary 3-celled, the ovules numerous,
superposed; style filiform, the stigma usually trifid; capsule subglobose, loculicidally
bivalvate; seeds flat or compressed, black.

About forty species, in tropical and temperate America. Only
the following ones occur in Central America.

Perianth white, 3.5-5 cm. long Z. tubispatha.

Perianth bright pink, or sometimes white tinged with pink, but then 6-7 cmT long.

Perianth 2.5-3 cm. long Z. Lindleyana.

Perianth usually 6-7 cm. long.

Ovary long-stipitate Z. carinata.

Ovary almost sessile Z. brevipes.

Zephyranthes brevipes (J. G. Baker) Standl. Field Mus. Bot.
4: 199. 1929. Z. carinata var. brevipes J. G. Baker ex Bonn. Smith,
Enum. PI. Guat. 3: 80. 1893, nomen nudum. Flor de Mayo.

Open grassy places, 1,200-1,800 meters, endemic; Alta Verapaz,
the type from Santa Cruz, 1,380 meters, J. D. Smith 1671; Chimal-
tenango; doubtless also in other departments.

Bulbs about 2 cm. in diameter, with an elongate neck; leaves equaling the
scape, 1.5-5 mm. wide; scapes slender, about 25 cm. high, the spathe 2 cm. long;
stipe of the ovary only 3-5 mm. long; perianth pink or white tinged with pink,
6-7.5 cm. long, the segments narrow, oblanceolate, 1 cm. wide or narrower.

Some of the specimens have been referred, almost certainly in
error, to Z. sessilis Herb. The species is said to be abundant in
some of the cultivated fields of Chimaltenango during the rainy
months.

Zephyranthes carinata Herb. Bot. Mag. pi. 2594. 1825. Ata-
mosco carinata Standl. in Standl. & Calderon, Lista PI. Salv. 51.
1925. Mananitas (Escuintla); Jacinto; Ceboy-ac (Coban, Kekchi;
the "ceboy" is the Spanish cebolla).

Frequent in gardens, planted for ornament, also in open fields
or thickets, especially of the Coban region; perhaps not native in

FIG. 28. Zephyranthes carinata. A. Habit of plant; X M- B. Flower;
X %. C. Stigmas and upper part of style; X 4. D. Stamen, showing anther and
upper part of filament; X 3.

144

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 145

Guatemala; collected more or less wild in Alta Verapaz and Quezal-
tenango, and cultivated generally. Mexico; Costa Rica; West
Indies.

Bulbs ovoid, about 2.5 cm. in diameter, with an elongate neck; leaves 15-30
cm. long, 2-5 mm. wide, shallowly channeled on the upper surface, often absent
at time of flowering; scapes 10-30 cm. long, the spathe 3-5 cm. long; stipe of the
ovary usually well over 1 cm. long; perianth rose-pink, the segments obovate,
usually 6-7 cm. long, 12-20 mm. wide. (Fig. 28.)

The plant seems to lie dormant through much of the dry season,
and blooms during the spring months. It bursts into flower quickly
if supplied with water. Baker states that this is the commonest
species in cultivation in Europe.

Zephyranthes Lindleyana Herb. Amaryll. 174. pi. 35, /. 5.
1837. Guatemala, at 1,350 meters; reported also from Chimalte-
nango; sometimes cultivated for ornament. Southern Mexico;
Honduras.

Bulbs globose, 1.5 cm. in diameter, with a short neck; leaves 15-20 cm. long,
usually present at anthesis, 1-6 mm. wide; scapes slender, 10-30 cm. long, the
spathe 2.5-3 cm. long, trifid at the apex only; perianth deep rose-pink, 2.5-3 cm.
long, the segments cuneate-obovate, about 12 mm. wide; stigma 3-fid.

Called "jacinto" in Salvador.

Zephyranthes tubispatha (Ker) Herb. App. 96. 1821. Ama-
ryllis tubispatha Ker, Bot. Mag. pi. 1586. 1813.

Pete"n (La Libertad, perhaps only in cultivation, but quite
possibly native on the savannas). West Indies; Colombia and
Venezuela.

Bulb globose, 2.5 cm. in diameter, with a short neck; leaves few, appearing
with the flowers, 30 cm. long or less, 2-5 mm. wide; scapes slender, about 15 cm.
long, the spathe 2.5-3 cm. long, cut only at the apex; stipe of the ovary much
elongate, usually exceeding the spathe; perianth 3.5-5 cm. long, white, slightly
tinged with green, with no red or pink, the tube almost obsolete, the segments
obovate, about 12 mm. wide; style deeply 3-fid; capsule 12 mm. broad.

The plant has been reported from Pete'n under the name Z. citrina
J. G. Baker.

DIOSCOREACEAE. Yam Family

References: R. Knuth, Dioscoreaceae, Pflanzenreich IV. 43: 99.
1924. C. V. Morton, Notes on Dioscorea, with special reference to the
species of the Yucatan Peninsula, Carnegie Inst. Wash. Publ.
461:241-253. 1936.

146 FIELDIANA: BOTANY, VOLUME 24

Chiefly perennial herbaceous vines, often arising from large tuberous roots;
leaves alternate or opposite, petiolate, usually cordate, palmately 3-13-nerved
and also reticulate-veined, entire or palmately lobate or foliolate, the petiole articu-
late at the base; flowers small, sessile or pedicellate, solitary or fasciculate, some-
times capitate-congested, arranged in spikes or racemes, generally unisexual,
dioecious or rarely monoecious, regular; bracts minute or none; racemes or spikes
axillary, often paniculate; perianth 6-parted, 2-seriate, the staminate deeply
parted, campanulate or explanate, the lobes subequal; stamens central, inserted
at the base of the perianth segments or on them, sometimes 6 and perfect, some-
times 3 perfect with 3 alternating staminodia, or only 3 and all perfect, free or
connate into a column; anthers generally small, subglobose or oblong, or the
cells distinct but contiguous, or the anthers didymous; pistillate perianth 6-parted,
persistent; staminodia minute or 3 or 6, sometimes none; ovary inferior, linear
or oblong, triquetrous, 3-celled; style 3, very short, distinct or connate, the stigmas
3, very short, entire or 2-fid; ovules 2 in each cell or rarely more, superposed,
pendulous, anatropous or almost amphitropous; fruit capsular, 3-valvate, rarely
by abortion monocarpic and winged above, sometimes baccate; seeds compressed
or globose, the testa appressed, usually broadly winged; endosperm almost carti-
laginous, the embryo enclosed in the endosperm, small.

Genera about eight, the species widely distributed, chiefly in
tropical regions. Only one genus is represented in continental
North America.

DIOSCOREA L.

Vines, usually herbaceous and twining, arising from underground tubers or
thick rhizomes; leaves alternate or opposite; flowers small, regular, unisexual,
dioecious; staminate perianth campanulate to rotate or tubuliform, the lobes
equal, linear to lanceolate or rounded; anthers small, the cells contiguous or
somewhat separated, free or rarely united; pistillate perianth 6-parted, persistent,
the staminodia minute and 3 or 6, or none; ovary linear or oblong, triquetrous,
3-celled, the 3 styles very short, reflexed, entire or 2-fid; ovules usually 2 in each
cell, sometimes more numerous, laterally attached below the apex; capsule tri-
quetrous, 3-winged, the wings vertical, loculicidally dehiscent; seeds compressed,
winged.

Species 400-500, almost all confined to the tropics, in both
hemispheres. Other species are known from Central America.

Leaves lobate or 3-foliolate.

Leaves 3-foliolate D. tenebrosa.

Leaves merely lobate.

Leaves deeply 3-5-lobate, the lobes all acuminate; cultivated plants.D. trifida.
Leaves hastately 3-lobate, the lateral lobes short, obtuse or rounded, native

plants D. Aguilarii.

Leaves entire.

Stems conspicuously winged D. alata.

Stems not winged.

Mature fruit very asymmetric, samaroid, one of the wings broad and well
developed, the others abortive and very narrow D. cyphocarpa.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 147

Mature fruit symmetric, not samaroid, all the angles or wings subequal.
Staminate flowers sessile or essentially so, often glomerate in small dense

clusters or heads (see D. hondurensis).
Stems usually bearing large aerial bulbs or bulblets in the leaf axils;

plants cultivated or rarely escaping D. bulbifera.

Stems not bearing bulblets in the leaf axils.

Seeds winged on the lower side only. Fertile stamens 6; perianth

puberulent D. Bartlettii.

Seeds winged all around.

Fertile stamens 3, alternating with 3 staminodia.
Staminate flowers solitary.

Leaves mostly 6-8 cm. long; perianth segments 3-4 mm. long.

D. dicranandra.
Leaves mostly 3 cm. long or shorter; perianth segments 1.5 mm.

long D. yucatanensis.

Staminate flowers glomerate.

Leaf blades mostly about 4 cm. long D. cyanisticta.

Leaf blades mostly 7-12 cm. long.

Cells of the anthers separated D. polygonoides.

Cells of the anthers coherent D. malagalpensis.

Fertile stamens 6.

Staminate flowers in cymose clusters; rudiment of the style large.
Leaves ovate-cordate, 7-9-nerved, glabrous. . .D. Bernoulliana.
Leaves orbicular-cordate, 11-nerved, pubescent beneath.

D. cymosula.
Staminate flowers in fasciculate heads; rudiment of the style

obsolete or nearly so.

Stamens unequal, 3 of them longer than the others . D. floribunda.
Stamens equal or nearly so.

Leaves stipulate D. Friedrichsthalii.

Leaves not stipulate.

Leaf blades abruptly cuneate-decurrent, elongate triangular-
cordate D. Tuerckheimii.

Leaf blades not decurrent upon the petiole, usually broadly

cordate D. macrostachya.

Staminate flowers conspicuously pedicellate.

Seeds winged only on the lower side. Stamens 3 D. convolvulacea.

Seeds winged all around.
Fertile stamens 6.

Flowers solitary D. belizensis.

Flowers cymulose D. Nelsonii.

Fertile stamens 3, sometimes with 3 alternating staminodia.

Staminodia none D. Standleyi.

Staminodia 3, alternating with the stamens.

Bracts of the Staminate inflorescence about 6 mm. long.

D. densiflora.
Bracts of the staminate inflorescence 2-3 mm. long. .D. Carionis.

Dioscorea Aguilarii Standl. & Steyerm. Field Mus. Bot. 22:
133. 1940. Name de ratdn.

Known only from the type, Dept. Guatemala, about 1,500
meters, Jose Ignacio Aguilar 90.

148 FIELDIANA: BOTANY, VOLUME 24

Plants very slender, the stems glabrous or shortly spreading-pilosulous; leaves
rather thin, on slender petioles 2-3.5 cm. long, the petiole short-pilose; leaf blades
hastate-lobate, 4.5-8.5 cm. long, 2.5 cm. wide near the base, glabrous above,
rather densely hirsute beneath, the middle lobe narrowly lance-oblong, 14-22 mm.
wide at the base, narrowly long-attenuate, 3-nerved, the basal lobes as much as
2 cm. long, somewhat rectangular, divergent, rounded on the angles, the outer
sides of the lobes subtruncate or broadly concave, 2-nerved; staminate spikes
slender and elongate, tortuous, geminate, long-pedunculate, the floriferous portion
2-5 cm. long, the rachis pilose with short spreading hairs; flowers sessile, in few-
many-flowered spikelets as much as 7 mm. long, these rather dense; perianth
segments scarcely 1 mm. long, oval, obtuse, glabrous; stamens 6, with very short
filaments.

Dioscorea alata L. Sp. PI. 1033. 1753. Name.

Native of tropical Asia or Africa, cultivated in most tropical
regions for its edible roots; planted commonly in Guatemala, espe-
cially in the lowlands, and sometimes at higher elevations, as in
the Department of Guatemala; sometimes escaping from cultivation
to second-growth thickets, as at Puerto Barrios.

Plants stout, glabrous, the stems narrowly 4-winged, sometimes producing
bulblets in the leaf axils; leaves long-petiolate, ovate-cordate or sagittate-cordate,
with a broad or narrow, usually very deep basal sinus, mostly 10-20 cm. long,
acuminate or caudate-acuminate; staminate spikes branched, elongate, the
spikelets verticillate, the flowers sessile; perianth 1.5 mm. in diameter, pale green,
the segments unequal; fertile stamens 6, very short; capsule lustrous, 3 cm. broad.

Sometimes called "name bianco" in Honduras; "macal," "max-
cal," "akilmaxcal" (Yucatan, Maya). This is one of two or three
yams of Old World origin that are planted commonly in Central
America, chiefly along the Atlantic coast where there is Carib or
West Indian influence. They doubtless were introduced first to the
West Indies by African slaves or slave ships, and later probably were
carried to the continent by Caribs or perhaps by the Spaniards. In
Guatemala yams are scarcely known to the majority of the people
on the Pacific coast, many of whom do not understand the term
"name," which probably is of African origin. The roots, however,
are sold in the markets of central Guatemala, most of them brought
probably from the lower parts of the Atlantic slope. Yams some-
what resemble large sweet potatoes, and often reach a great size.
The outside of the hard tuberous root bears many black hair-like
roots, while the inside usually is white. It is cooked and eaten like
a potato, generally boiled. When on the table, a yam is firm and
solid, and to many people not attractive, although in flavor it
probably is not much inferior to a white potato. The roots are so
large that often they are cut into pieces for sale, and for cutting

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 149

them an axe or machete is needed. In the United States the name
"yam" (derived obviously from name) is often given to large sweet
potatoes (Ipomoea Batatas), which do not resemble very closely
true yams.

Dioscorea Bartlettii Morton, Carnegie Inst. Wash. Publ. 461:
242. 1936. Cocomeca; Cocomeca blanca (Pete"n).

Moist or wet thickets and forest, often or perhaps always on
limestone, chiefly at 1,000 meters or less; Pete"n (type from Uaxactun,
H. H. BartUtt 12425); Alta Verapaz; Izabal. Tabasco; British
Honduras.

Rhizome woody, epigaean, semi-ellipsoid, very large, covered with dark-colored
bark-like plates; stems sinistrorsely twining, minutely puberulent, spiny near
the base; leaves alternate, long-petiolate, the petioles 7 cm. long or less; leaf blades
ovate, entire, the larger ones about 16 cm. long and 14 cm. wide, chartaceous,
abruptly acuminate or caudate-acuminate, broadly cordate or almost truncate
at the base, 7-nerved, glabrous; staminate inflorescences as much as 20 cm. long,
usually much branched, the branches densely puberulent; flowers sessile, aggregate,
2-3-nate or rarely solitary, purple; perianth 1.5 mm. long, puberulent, the lobes
ovate, subobtuse; fertile stamens 6, the filaments very short, the anther cells
contiguous, discrete; style rudiment none; pistillate inflorescence spicate, some-
times branched, as much as 23 cm. long; sterile stamens 6; styles 3, very short;
capsule oblong, as much as 4.5 cm. long and 2 cm. broad, subligneous; seeds 2.5
cm. long, winged only on the lower side.

It is of some interest to find that the name "cocolmeca," of
Nahuatl origin, is used in Pete"n. This name is given in south-
western Mexico to other species of the genus that arise from epigaean
tubers whose surface is covered with plates in such a manner as to
suggest a turtle shell. There the sap of the young shoots is used
in the preparation of pozonque, a beverage made from chocolate and
soft corn, and this juice beaten into a foamy mass. The large root
is used there also as a barbasco, for stupefying fish, and perhaps
contains the alkaloid dioscorine that has been found in other species.
Asiatic species of Dioscorea are sometimes used likewise as fish
poisons, and some of them produce roots that are poisonous to man
if eaten, at least when raw.

Dioscorea belizensis Lundell, Contr. Univ. Mich. Herb. 6: 5.
1941.

Moist or wet thickets, 900 meters or less; Alta Verapaz (?);
Zacapa. British Honduras, the type from Carib Reserve, Stann
Creek District, P. H. Gentle 2998.

150 FIELDIANA: BOTANY, VOLUME 24

A small or large vine with thick hypogaean roots, the stems often stout;
leaves on petioles 10.5 cm. long or less, the petioles glabrous; leaf blades thick-
chartaceous, glabrous above, puberulent or glabrous beneath, broadly ovate, as
much as 22 cm. long and 17 cm. wide, cuspidate-acuminate or rounded and cuspi-
date at the apex, deeply cordate at the base, 11-nerved, the veins often impressed
on the upper surface; staminate inflorescences solitary, as much as 150 cm. long,
the racemes paniculate; bractlets broadly ovate, cuspidate, 2 mm. long or less;
pedicels 1 mm. long or longer, the flowers dark red, 3 mm. long; perianth segments
broadly ovate; fertile stamens 6.

Dioscorea Bernoulliana Prain & Burkill, Kew Bull. 1916: 192.
1916.

Moist or wet thickets or forest, 1,000 meters or less; Pete"n;
Izabal. Collected also between "Escomillas" and Palo Hueco (de-
partment uncertain), Bernoulli & Cario 847. British Honduras.

Plants glabrous, the stems sinistrorsely twining; leaves on petioles 4 cm. long
or less, ovate-cordate, acutely acuminate, broadly and shallowly cordate at the
base, membranaceous or rather firm, 7-9-nerved, about 8 cm. long and wide;
staminate racemes solitary or 2-4-nate, simple, as much as 16 cm. long; flowers
cymulose, the cymules on very short peduncles, 2-4-flowered, the bracts 1 mm.
long; perianth 1.5 mm. broad, the outer segments oblong, the inner ones slightly
broader; stamens 6, the filaments adnate to the base of the segments, the anthers
suborbicular; ovary rudiment trigonous, half as long as the filaments; capsule
glabrous, dark brown, about 14 mm. long and 17 mm. broad; seeds orbicular,
completely surrounded by a thin, dark brown wing 2-4 mm. wide.

Dioscorea bulbifera L. Sp. PL 1035. 1753. Papa del aire;
Name montaraz; Papa voladora.

Probably native of tropical Asia and perhaps also of Africa; often
cultivated in other tropical regions; planted in Guatemala for
ornament or as a curiosity, or sometimes for food, at low and middle
elevations.

Plants glabrous, the stems producing numerous large or small bulblets, these
succulent, somewhat compressed, sometimes 10 cm. broad, smooth or somewhat
verrucose; leaves alternate, long-petiolate, rounded-cordate-ovate, 7-15 cm. wide,
cuspidate-acuminate, about 9-nerved, membranaceous; stipules present, mem-
branaceous, oblong-lanceolate; staminate spikes slender, simple, 3-10 cm. long,
sometimes paniculate, the flowers solitary, approximate, variable in size; perianth
segments lanceolate, spreading; stamens 6, minute, the anthers oblong-elliptic;
style rudiments 3, subulate; pistillate spikes simple, 10-25 cm. long, fasciculate,
the flowers erect and often appressed to the rachis; capsule 2.5 cm. long and 1.5 cm.
broad, oblong, rounded at each end or sometimes subacute at the apex.

The Maya name of Yucatan is reported as "bauiac." In central
Guatemala this plant is grown principally as a curiosity, but it is
said to be grown rather commonly for food in the Oriente, often
planted with "huisquil" (Sechium) and allowed to climb over the same

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 151

supports with it. The large fleshy roots are cooked and eaten, and
the often very large and heavy, greenish bulblets are used in the
same manner.

Dioscorea Carionis Prain & Burkell, Kew Bull. 1916: 193. 1916.

Moist or wet forest or thickets, 400-1,800 meters; endemic; El
Progreso; Zacapa; Escuintla; Suchitepequez; Quezaltenango; San
Marcos.

A slender vine, glabrous or nearly so, sometimes finely and sparsely puberulent,
the stems sinistrorsely twining; leaves long-petiolate, ovate-cordate or rounded-
cordate, sometimes 15 cm. long and 13 cm. wide, abruptly acuminate or cuspidate-
acuminate, membranaceous, rather deeply and broadly cordate, 7-9-nerved, some-
times puberulent beneath on the nerves; flowers pale green, the stamina te racemes
axillary, solitary or fasciculate, as much as 20 cm. long, the flowers solitary, the
pedicels 2 mm. long, puberulent; bracts ovate-lanceolate, 3 mm. long or less;
perianth 5 mm. broad, the segments ovate-lanceolate, acute, 3-4 mm. long, sub-
equal; stamens 3, the anthers suborbicular; staminodia 3; capsule about 13 mm.
long and 12 mm. broad, brownish.

Dioscorea convolvulacea Schlecht. & Cham. Linnaea 6: 49.
1831. D. capillaris Hemsl. Biol. Centr. Amer. Bot. 3: 354. 1884.
D. capillaris var. glabra Hemsl. loc. cit. D. esurentium Uline, Bot.
Jahrb. 22: 429. 1897 (type locality given erroneously as Guatemala).
D. convolvulacea var. glabra Uline ex Knuth, Pflanzenreich IV. 43:
99. 1924. D. convolvulacea var. esurentium Uline ex Knuth, loc.
cit. Madre de maiz.

Moist or wet thickets or forest, 2,000 meters or less, most common
at rather low elevations; Alta Verapaz; Zacapa; Chiquimula; Jalapa;
Santa Rosa; Escuintla; Guatemala; Quiche". Southern Mexico;
British Honduras to Costa Rica; Trinidad.

Plants arising from small or large, tuberous roots, the stems short-hirtellous
or glabrous, the plants often glabrous throughout or nearly so; leaves long-petio-
late, rounded-ovate or ovate-cordate, sometimes 30 cm. long and wide but usually
much smaller, acuminate or caudate-acuminate, shallowly or rather deeply cordate,
7-13-nerved, glabrous above or nearly so, glabrous beneath or often rather densely
pubescent or puberulent; staminate inflorescences long and slender, simple or
branched below, the rachis puberulent, the flowers solitary, green or purplish, on
pedicels 2-3 mm. long; perianth 3-4 mm. broad, the segments spreading or finally
reflexed, oblong, obtuse; fertile stamens 3, divaricate, the anthers minute; capsule
oblong or oblong-elliptic, glabrous, 12-14 mm. long and 9 mm. broad or some-
what larger; seeds winged on the lower side.

Probably the most common species of the genus in Guatemala,
abundant in some localities, flowering during the wet months. The
material is somewhat variable, but the varieties that have been

152 FIELDIANA: BOTANY, VOLUME 24

proposed seem to be of little importance. This and perhaps some
of the other similar wild species of Dioscorea are of considerable
economic importance in Guatemala, and they are well known,
especially in the Occidente, under the name "madre de maiz." In
some years because of unfavorable weather or insect attacks the
corn crop fails in regions of Guatemala, a very serious event indeed,
considering that maize is the most important food of the country.
In recent years the national government has done much to alleviate
the resultant suffering, but in former times local famines resulted.
In such periods the starving people had to go to the fields and forest
to find any plants that could be used as emergency foods. Very
important on the Pacific slope was madre de maiz, whose tuberous
roots were grated and made into a kind of tortilla or tamal. It is
said that they are an unappetizing food, but that they serve to
relieve hunger and at least prevent death by starvation. This species
has been reported from Guatemala as D. propinqua Hemsl. and as
D. hirsuta var. glabra Uline.

Dioscorea cyanisticta Bonn. Smith, Bot. Gaz. 20: 10. 1895.
Tzahuacax (Huehuetenango).

At 1,400-2,100 meters; Huehuetenango (Sierra de los Cuchu-
matanes, between Chacula and Canquintic, Steyermark 51779;
determined by Morton). Costa Rica.

Plants slender, glabrous or nearly so; leaves membranaceous, small, pellucid-
lineolate, broadly ovate-cordate, mostly 4 cm. long or shorter, acuminate or long-
acuminate, deeply cordate at the base, with 11 or fewer nerves, glandular beneath
at the base; staminate spikes mostly simple, clustered in the leaf axils or forming
a lax panicle, subsessile, 10-15 cm. long, the rachis scabrous-puberulent; flowers
in clusters of 2-3, rarely solitary, equaling the elongate-lanceolate bract, sessile
or nearly so, the glomerules approximate; perianth campanulate, 3 mm. in diameter,
the segments oblong-ovate, spreading, twice as long as the tube, 1-nerved; fertile
stamens 3, short, the anthers globose, extrorse; staminodia 3, filiform, equaling
the fertile stamens.

Dioscorea cymosula Hemsl. Biol. Centr. Amer. Bot. 3: 355.
pi 90. 1884.

At 1,500 meters, Santa Rosa, Buena Vista, Heyde & Lux 6391
(recorded from Guatemala as D. composite, Hemsl.). Mexico;
Panama.

Stems rather stout, puberulent; leaves long-petiolate, membranaceous, orbicu-
lar-cordate, 14 cm. wide or less, acute or apiculate, 1 1-nerved, whitish-pubescent,
especially beneath, the petiole as much as 7.5 cm. long; staminate racemes elongate,
sometimes 30 cm. long, simple; flowers sessile or nearly so in small cymules, 3-4

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 153

mm. broad, the cymules 2-6-flowered; bracts narrow, longer than the flowers;
perianth segments almost free, lance-oblong, subobtuse, villosulous outside, erect-
patent; stamens 6, inserted at the base of the segments, the anthers elliptic;
style rudiment rather large, conic; pistillate flowers with 6 sterile stamens, the
ovary densely tomentose; capsule elliptic, densely tomentose, 2.5 cm. long and
2 cm. broad or perhaps larger.

The single Guatemalan collection cited was reported by Knuth.
The specimens, deposited in the United States National Herbarium,
were examined by the authors.

Dioscorea cyphocarpa Robinson ex Knuth, Notizbl. Bot. Gart.
Berlin 7: 209. 1917. D. tacanensis Lundell, Lloydia 2: 78. 1939
(type from Volcan de Tacana, Chiapas, at 1,000-2,000 meters,
E. Matilda 2416).

Doubtless on Volcan de Tacana in San Marcos. Southern and
western Mexico.

Plants glabrous or nearly so, the stems slender or rather stout; leaves mem-
branaceous, on petioles 10 cm. long or less, ovate-cordate, 12.5 cm. long or shorter,
acuminate, deeply cordate at the base, glabrous, 11-13-nerved; staminate spikes
5 cm. long, simple, rather remotely flowered, solitary or binate, pedunculate, the
rachis puberulent, the pedicels 3-4 mm. long; perianth 3-4 mm. broad, cam-
panulate-rotate, the lobes lanceolate, subobtuse; stamens 3, erect, central; style
rudiment none; pistillate spikes in fruit 6 cm. long or more, solitary, simple,
pedunculate, the flowers in fruit on long filiform pedicels; capsule 2 cm. long and
1.5 cm. wide or smaller, cuneately narrowed at the base, very asymmetric, 2 of
the cells sterile and with very narrow wings, the third cell with a broad wing;
seeds 7-8 mm. long, 5 mm. wide.

Dioscorea densiflora Hemsl. Biol. Centr. Amer. Bot. 3: 356.
1884.

Moist or wet thickets or forest, 1,300 meters or less; Alta Verapaz;
Izabal. Southern Mexico; British Honduras; Honduras (Cozumel
Island).

Plants glabrous except for the puberulent racemes, the stems slender, some-
what succulent; leaves on slender petioles 3-4 cm. long, membranaceous, bright
green when dried, ovate-cordate, 8-15 cm. long or smaller, acuminate, openly
cordate, 7-nerved; staminate racemes slender, many-flowered, 20 cm. long or
shorter, solitary or 2-4-nate, simple or furcate at the base, densely flowered;
pedicels solitary, 2-3 mm. long, the bracts linear-filiform, longer than the flowers;
perianth segments membranaceous, cream-colored, lanceolate, acuminate; fertile
stamens 3, inserted in the throat of the perianth; staminodia clavate or spatulate,
equaling the fertile stamens; style rudiment 3-tuberculate; pistillate spikes simple;
perianth segments ovate-oblong; sterile stamens 6; capsule elliptic, 2.5 cm. long,
1.7 cm. broad; seeds oblong, irregularly winged on all sides, 8-10 mm. long.

154 FIELD IANA: BOTANY, VOLUME 24

Dioscorea dicranandra Bonn. Smith, Bot. Gaz. 19: 13. 1894.
D. albicaulis Uline, Bot. Jahrb. 22: 425. 1896 (type from Cerro
Gordo, Santa Rosa, Heyde & Lux 3869).

Moist forest or thickets, sometimes in oak forest, 800-2,050
meters; endemic: Alta Verapaz; Santa Rosa; Escuintla.

Plants herbaceous, glabrous, the stems rather stout and succulent, sinistrorsely
twining; leaves on slender, rather short petioles, membranaceous, ovate-cordate
or broadly cordate, 6-8 cm. long and wide, long-acuminate, broadly and shallowly
cordate, 7-nerved; staminate inflorescences simple or sometimes sparsely branched
below, as much as 20 cm. long, solitary, the flowers 2-4 mm. apart, sessile; perianth
rotate, the segments rather thick, purplish or when dried brownish, lance-oblong,
obtuse, 3-4 mm. long, twice as long as the bracts; fertile stamens 3; staminodia 3,
clavate; style rudiment none.

Dioscorea floribunda Mart. & Gal. Bull. Acad. Brux. 9, pt. 2:
391. 1842.

Moist or wet thickets or forest, sometimes in oak forest, 1,500
meters or less; Baja Verapaz; Izabal; Santa Rosa; Suchitepe"quez;
Huehuetenango. Southern Mexico.

Plants glabrous, the stems slender or stout, sinistrorsely twining; leaves alter-
nate, on petioles 7 cm. long or less, rather thick and firm, broadly ovate or tri-
angular-ovate, medium-sized, long-acuminate or caudate-acuminate, shallowly or
deeply cordate, 9-nerved; staminate spikes approximate, usually paniculate, the
rachises very slender, the flowers solitary or 2-3-nate, sessile or nearly so, dark
green, 2-3 mm. long, the perianth membranaceous, tubular, the segments erect,
shorter than the tube, elliptic, obtuse; fertile stamens 6, central, 3 of them equaling
the tube, the other 3 much shorter; anthers didymous; style rudiment none;
pistillate spikes rather short, solitary; perianth distinctly stipitate, campanulate;
sterile stamens 6.

Dioscorea Friedrichsthalii Knuth, Pflanzenreich IV. 43: 169.
1924.

Known only from the type, said to have been collected in Guate-
mala by Friedrichsthal, the locality not indicated.

Plants glabrous, the stems slender; leaves alternate, on petioles about 4 cm.
long, papyraceous, brownish when dried, 13 cm. long, 10 cm. wide, obtuse or
acuminate, deeply cordate, 7-9-nerved, the basal lobes rounded; pistillate racemes
10-15 cm. long, pedunculate; capsule quadrangular-rounded, broadly cuneate-
narrowed at the base, broadly incised at the apex, coriaceous, brown, 22 mm.
long and broad, somewhat lustrous; seeds winged, 10 mm. long, 8 mm. wide,
fuscous, the wings 2-3 mm. wide.

There is no certainty that this plant was collected in Guatemala,
since the "Guatemala" heading on Friedrichsthal's labels is not
dependable, and the type may well have come from Honduras,
Nicaragua, or Costa Rica.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 155

Dioscorea hondurensis Knuth, Repert. Sp. Nov. 38: 120.
1935.

Known only from the type, upper Moho River, British Honduras,
M. E. Peck 549.

Stems stout; leaves alternate, on petioles about 7 cm. long, firm-papyraceous,
cordate-ovate, 16 cm. long and 11 cm. wide, acute, rather deeply cordate, sub-
decurrent on the petiole, 9-nerved; pistillate spikes sometimes 50 cm. long, flori-
ferous to the base; capsule obtriquetrous, as much as 4.5 cm. broad and 3 cm. long,
almost retuse at the apex, borne on a pedicel 1 cm. long.

The species is known only from fruiting specimens, and on that
account it is not included in the key to species presented on the
preceding pages.

Dioscorea macrostachya Benth. PI. Hartweg. 73. 1839. D.
composite, Hemsl. Biol. Centr. Amer. Bot. 3: 354. 1884. D. astro-
stigma Uline, Bot. Jahrb. 22: 3. 1896 (type from Volcan de Fuego,
Sacatepe"quez, J. D. Smith 2565). Peng (Huehuetenango).

Moist or dry thickets or forest, 1,500 meters or less; Alta Verapaz;
Izabal; Jalapa; Santa Rosa; Sacatepe"quez ; Quezaltenango; San
Marcos ; Huehuetenango. Southern Mexico to Salvador and Panama.

Plants glabrous or nearly so, the stems stout or slender, usually dextrorsely
twining; leaves alternate, long-petiolate, membranaceous or chartaceous, 20 cm.
long and 18 cm. wide or usually much smaller, abruptly acute or cuspidate-
acuminate, shallowly or deeply cordate, 7-9-nerved; staminate inflorescences
simple or branched, 15-30 cm. long, the flowers fasciculate-glomerate, the fascicles
short-stipitate, 2-3-flowered, dense, the flowers sessile or nearly so; perianth rotate,
1.5-2 mm. broad, the segments somewhat carnose, rounded and obtuse, dark
brownish when dried; fertile stamens 6, subcentral, the anthers subsessile; pistil-
late spikes simple; sterile stamens 6, minute; stigmas subsessile, thick, divergent,
furcate or stellate-bifurcate at the apex; capsule oblong or oval, 2.5 cm. long, the
angles marginate; seeds oblong, winged on all sides. (Fig. 29.)

Called "camotillo" in Honduras, where the root is used as a fish
poison; "cuculmeca" (Salvador).

Dioscorea matagalpensis Uline, Bot. Jahrb. 22: 432. 1897.
Moist or wet thickets or forest, 750 meters or less; Pete"n; Zacapa.
Yucatan Peninsula of Mexico; British Honduras; Nicaragua.

Plants slender, the stems sinistrorsely twining; leaves on long slender petioles,
membranaceous, green when dried, rounded-cordate, 10 cm. long and 7 cm. wide
or smaller, acuminate or narrowly long-acuminate, deeply cordate at the base,
7-nerved, glabrous; staminate spikes 10 cm. long, simple or sparsely branched,
the flowers in 3-4-flowered glomerules; perianth 1.5 mm. broad, the segments
ovate-oblong, obtuse, equal; fertile stamens 3, the anthers small, the cells not

FIG. 29. Dioscorea macrostachya. A. Habit of portion of staminate plant;
X %. B. Habit of portion of pistillate plant; X % C. Staminate flower, half
of it viewed from within; X 5. D. Staminate flower, viewed from above; X 3.
E. Pistillate flower; X 6. F. Seed; X 1. G. Staminate flowers in position,
viewed from side; X 4. H. Capsule; X 1. I. Stamen; X 18. J. Pistillate flower,
viewed from above; X 3.

156

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 157

discrete; sterile stamens 3, spatulate, all the stamens inserted on the perianth
tube above its base; style rudiment none; pistillate racemes solitary, simple, 8-12
cm. long; perianth subrotate, fuscous, 1.5 mm. broad, the segments ovate-orbicular,
obtuse, membranaceous; sterile stamens 6; capsule glabrous, about 14 mm. long
and 11 mm. broad; seeds winged on all sides.

Dioscorea Nelsonii Uline ex Knuth, Notizbl. Bot. Gart. Berlin
7: 202. 1917.

Moist or wet, mixed forest, sometimes in pine forest, 1,000-1,500
meters; Solola; San Marcos; determinations of both collections are
questionable. Southern and western Mexico.

Stems stout, sinistrorsely twining, glabrous except sometimes at the nodes;
leaves large, long-petiolate, membranaceous or chartaceous, orbicular-cordate,
18 cm. long and wide or smaller, 11-nerved, glabrous, rather shallowly cordate
at the base, acuminate; racemes elongate, the flowers in 3-5-flowered fascicles,
short-pedicellate or almost sessile; staminate perianth 2.5 mm. long, the segments
spreading, narrowly oblong; fertile stamens 6, subincurved, slightly shorter than
the perianth, inserted at the base of the segments, the anthers oblong, introrse;
style rudiment rather large; pistillate perianth obscurely puberulent; capsule
elliptic, acute at the base and apex, about 33 mm. long and 22 mm. broad.

Dioscorea polygonoides Humb. & Bonpl. ex Willd. Sp. PL 4:
795. 1805.

Wet to dry thickets or forest, 2,000 meters or less; Chiquimula;
Santa Rosa; Escuintla; Retalhuleu. Southern Mexico; Panama;
West Indies; northern South America.

Plants glabrous, the stems sinistrorsely twining; leaves on long or short
petioles, membranaceous, usually pale green when dry, cordate-ovate, mostly 8-12
cm. long and almost as wide, cuspidate-acuminate, deeply or shallowly cordate,
7-9-nerved; staminate spikes simple or with a few branches, sometimes much
elongate, the rachis scaberulous, the flowers sessile, glomerulate, the glomerules
rather remote, 3-5-flowered; perianth membranaceous, green, glandular-puncticu-
late outside, turbinate-rotate, the lobes short-ovate, often auriculate at the base;
stamens 6, inserted on the perianth tube, 3 of the stamens fertile, the anthers
extrorse, with distinct cells; sterile stamens 3, entire or 2-fid at the apex, slightly
shorter than the fertile ones; style rudiments 3, distinct, minute; pistillate racemes
simple, sterile stamens 6, with 3 of them antheriferous; stigmas 2-lobate at the
apex; capsule glabrous, broadly elliptic, 2-2.5 cm. long, 1.7-2.2 cm. wide, or
sometimes smaller; seeds winged on all sides.

Dioscorea Standleyi Morton, Carnegie Inst. Wash. Publ. 461:
252. 1936.

At 500-1,500 meters; Chiquimula (Cerro Tixixi, north of Jocotan,
Steyermark 31622; determined by Morton). Mountains of Costa
Rica.

158 FIELDIANA: BOTANY, VOLUME 24

Stems glabrous; leaves alternate, long-petiolate, ovate-cordate, 12 cm. long
and 10 cm. wide or smaller, papyraceous, acuminate, rather shallowly cordate at
the base, 9-nerved, glabrous; staminate spikes solitary or binate, 14-15 cm. long,
glabrous, the flowers arranged in 2-3-flowered cymules, the pedicels 2-4 mm. long;
staminate perianth green, glabrous, the segments linear-oblong, 2 mm. long;
stamens 3, the anthers globose, extrorse, the cells distinct; style rudiment none;
pistillate inflorescence racemose, spike-like, solitary, simple, 11 cm. long or less,
the flowers on very short pedicels, the bracts lanceolate; perianth stipitate, the
segments linear-oblong, 1.5 mm. long; staminodia none; ovary glabrous; capsule
(immature) 15 mm. long, 5 mm. wide; seeds winged on all sides.

Dioscorea tenebrosa Morton, Carnegie Inst. Wash. Publ. 461:
247. 1936.

Known only from the type, Pansamala, Alta Verapaz, 1,100
meters, Turckheim 1241.

Stems sinistrorsely or dextrorsely twining, glabrous; leaves alternate, on
petioles 3 cm. long or less, membranaceous, glabrous, 3-parted, the middle segments
lanceolate, up to 8 cm. in length and 3.5 cm. wide, acuminate, narrowed and cuneate
at the base, 3-nerved, the lateral segments shorter, the largest 5 cm. long and
2.5 cm. wide, somewhat oblique and falcate; staminate inflorescences racemose,
solitary or binate, simple, 10 cm. long or less, the flowers solitary, approximate,
the pedicels 1-1.5 mm. long, glabrous; perianth rotate, 5-6 mm. broad, the seg-
ments almost free, oblong, glabrous; stamens 3, inserted on the margin of a fleshy
disk, the anthers sessile; style rudiment none.

This has been recorded from Guatemala as D. trifoliata HBK., a
South American species.

Dioscorea Tuerckheimii Knuth, Notizbl. Bot. Gart. Berlin 7:
203. 1917.

Moist or wet thickets or forest, Alta Verapaz (known only from
the type, Cubilgiiitz, Turckheim 7787).

Plants glabrous; leaves alternate, slender-petiolate, papyraceous, elongate-
cordate-triangular, about 10 cm. long and 6 cm. wide, gradually narrowed to the
acuminate apex, very shallowly cordate or subtruncate at the base, the blade
usually abruptly short-decurrent upon the petiole, 9-nerved; staminate spikes
5-7 cm. long, solitary, dense, the bracts lance-acuminate; flowers fasciculate,
the fascicles mostly 2-3-flowered, the pedicels 0.5-3 mm. long; perianth campanu-
late-globose, 1.5 mm. broad, somewhat fleshy, the segments broadly ovate, often
broader than long, rounded at the apex or almost retuse; fertile stamens 6, attached
to the base of the perianth tube, the filaments short; style rudiment conoid;
capsule about 2 cm. long and 1.5 cm. broad, glabrous.

This has been reported from Guatemala as D. propinqua Hemsl.

Dioscorea yucatanensis Uline, Field Mus. Bot. 1: 416. 1899.
Reported with some doubt by Morton from Duck Run, El Cayo
District, British Honduras, H. H. Bartlett 11560. Yucatan.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 159

Plants slender, glabrous; leaves alternate, on petioles 1-5 cm. long, mem-
branaceous, orbicular-cordate, very small, 7-nerved; staminate spikes solitary,
much longer than the leaves, the flowers minute, sessile, subsolitary, dark purple;
perianth 1.5 mm. in diameter, the segments membranaceous, oblong; fertile stamens
3, short, subcentral, alternating with 3 spatulate staminodia, erect-divergent, the
anthers minute, introrse; style rudiment none; pistillate spikes few-flowered, twice
as long as the leaves in age, 7-10-flowered; sterile stamens 3, alternating with 3
minute ones; ovary glabrous.

IRIDACEAE. Iris Family
Reference: J. G. Baker, Handbook of the Irideae, London. 1892.

Perennial herbs, with elongate or bulb-like rootstocks, rarely annuals with
fibrous roots or perennials with a cluster of tuberous-thickened roots; leaves
usually equitant, 2-ranked; flowers perfect, regular or irregular, small or large,
solitary or in clusters from spathe-like bracts; perianth of 6 lobes or distinct
segments, the tube adnate to the ovary, the segments 2-seriate, convolute in bud,
withering-persistent; stamens 3, inserted on the perianth opposite the outer
segments, the filaments distinct or partially united, the anthers 2-celled, extrorse;
ovary inferior, 3-celled; ovules numerous, anatropous; styles distinct, entire or
parted, sometimes petaloid; fruit capsular, loculicidally 3-valvate; seeds numerous,
in 1-2 rows in each cell; embryo straight, the endosperm carnose or corneous.

Genera about 55, with 800 species or more, widely distributed
in tropical and temperate regions. No other genera are known to
be native in Central America.

Plants arising from rootstocks or from a cluster of fibrous or fleshy-thickened

tuber-like roots, never from bulb-like corms; leaves not plicate.
Plants without rootstocks, arising from a cluster of fibrous or fleshy tuber-like

roots; flowers small, blue, white, or yellow Sisyrinchium.

Plants with short or elongate rootstocks; flowers usually large.

Stems broadly winged Neomarica.

Stems not winged.

Leaves with a conspicuous costa. Flowers yellow, about 2 cm. long.

Trimeza.

Leaves not with an evident costa, the nerves all about equally prominent.
Style branches opposite the stamens and the outer perianth segments;

cultivated plants, sometimes naturalized Iris.

Style branches alternate with the anthers; native plants. . .Orthrosanthus.
Plants arising from bulb-like corms covered with membranaceous or firm tunics;

leaves often plicate.
Leaves not plicate; cultivated plants, rarely becoming naturalized.

Perianth 3-4 cm. long; spathes about 8 mm. long; perianth orange-red.

Tritonia.

Perianth much larger; spathes large; perianth variously colored. . .Gladiolus.
Leaves evidently plicate.

Style branches opposite the stamens and outer perianth segments.

Perianth divisions subequal Nemastylis.

Perianth divisions unequal.

160 FIELDIANA: BOTANY, VOLUME 24

Perianth about 7 cm. long, the inner segments large, spreading, panduri-

form, the outer ones orange-red and yellow Tigridia.

Perianth 3-3.5 cm. long, the inner segments small, appressed, ovate, the

outer ones blood-red Rigidella.

Style branches alternate with the anthers.

Perianth segments very unequal, the inner ones small; style-arms not
linear-subulate; spathes sessile. Perianth white or bluish white.

Cipura.

Perianth segments subequal; spathes usually pedunculate; style-arms linear
to subulate.

Filaments united Gelasine.

Filaments free.

Leaves broad, strongly plicate; inflorescence pseudolateral; tunics

purple or purple-brown; very thick Eleutherine.

Leaves very narrow, or, if broader, not strongly plicate; inflorescence
not pseudolateral, or if so, spathes 1-flowered; tunics castaneous
or blackish-brown, thin Calydorea.

CALYDOREA Herbert

Plants perennial, glabrous, arising from membranous, thin, castaneous or
blackish-brown tunicate bulbs; leaves few, very narrow, or, if broader, not strongly
plicate; spathes 1-flowered, the inflorescence terminal, not pseudolateral, peduncu-
late; perianth tube none, the segments equal or subequal, spreading, obovate-
cuneate; stamens attached at the base of the segments, alternate with the style-
arms, the filaments erect, free, the anthers linear, rolling up spirally at the tip;
ovary oblong, 3-celled; style filiform, the style-arms narrowly linear, of the same
width throughout, entire, or rarely briefly bifid at the apex, stigmatose at apex;
capsule obovoid or turbinate, exserted, loculicidally 3-valvate; seeds small, globose
or angulate.

About 8 species, all but the following South American.

Calydorea guatemalensis (Standl.) Foster, Contr. Gray Herb.
155: 46. 1945. Eleutherine guatemalensis Standl. Field Mus. Bot. 4:
200. 1929.

Moist shaded soil, sometimes in pine forest, 1,200-2,000 meters;
endemic; Alta Verapaz (type from Coban, Tiirckheim II. 1885);
Baja Verapaz; Zacapa; Chiquimula; Escuintla.

Bulb about 3.5 cm. long, with firm brown tunics; basal leaves 2-3, petiolate,
broadly linear, 30-70 cm. long, 1-2.5 cm. wide; scapes 40-70 cm. high, slender,
the bract at the base of the inflorescence similar to the basal leaves but narrower;
spathes one or more, 4.5-6.5 cm. long, the peduncles mostly 8-20 cm. long; perianth
white, turning bluish in age, 3 cm. broad or less; capsule obpyramidal-obovoid,
12-15 mm. long.

CIPURA Aublet

Rootstock a small ovoid bulb-like corm with thin loose brown tunics; radical
leaves few, plicate; scape terete, naked, bearing at the apex a dense head-like cluster
of spathes, the inflorescence subtended by a reduced plicate leaf; perianth parted

FIG. 30. Cipura paludosa. A. Habit of plant; X Vs. B. Spathe; X 4.
C. Outer (left) and inner (right) perianth segments; X 4. D. Capsule; X 4.
E. Stamen and style-branches; X 4.

161

162 FIELDIANA: BOTANY, VOLUME 24

to the ovary, the outer segments obovate-cuneate, becoming reflexed, the inner
segments shorter, connivent; stamens inserted at the base of the inner segments,
the anthers as long as the free filaments; ovary clavate, the ovules numerous,
superposed; style filiform, the branches oblong, petaloid; capsule oblong-turbinate,
membranous; seeds numerous, small, angulate.

The genus consists of a single species.

Cipura paludosa Aubl. PI. Guian. 38. pi. 13. 1775. C. cubensis
Griseb. Cat. PL Cub. 251. 1866.

Open grassy places, often in savannas or thin pine forest, some-
times a weed about dwellings or in cultivated ground, 1,500 meters
or less; Pete*n; Alta Verapaz; Izabal; Chiquimula; Suchitepequez;
Quiche" ; Huehuetenango. Mexico; British Honduras to Panama;
West Indies; South America.

Basal leaves usually 2-3 and 20-40 cm. long, 3-12 mm. wide, glabrous; scape
not winged, very slender, 15-45 cm. high; spathes cylindric, 2.5-5 cm. long,
green, the outer one short; pedicels slender, about equaling the spathe; perianth
white or tinged with lavender, very delicate and fugacious, somewhat campanu-
late, nutant, the outer segments 12-20 mm. long, the inner ones much shorter,
with yellow blotches near the base; capsule 1-2 cm. long; seeds dark brown, hard,
obtusely angulate. (Fig. 30.)

ELEUTHERINE Herbert

Plants perennial, glabrous, arising from membranous, very thick, purple or
purple-brown tunicate bulbs; leaves few, broad, strongly plicate; spathes one or
two, several-flowered, the inflorescence pseudolateral, pedunculate; perianth tube
none, the segments equal or subequal, spreading, obovate-cuneate; stamens
attached at base of the segments, alternate with the style-arms, the filaments
short, free, anthers linear; ovary oblong, 3-celled; style very short, style-arms subu-
late, entire, broadest at or near the middle, stigmatose at apex; capsule oblong,
loculicidally 3-valvate; seeds small, angulate.

Probably 2 species ranging from Mexico, Central America and
the West Indies to South America.

Eleutherine bulbosa (Mill.) Urban, Repert. Sp. Nov. 15: 305.
1918. Sisyrinchium bulbosum Mill. Gard. Diet. ed. 8: no. 3. 1768.
Sisyrinchium palmifolium L. Mant. PL 1: 122. 1767. Eleutherine
plicata Herb, ex Klatt in Mart. Fl. Bras. 3, pt. 1: 514. 1871. E.
palmifolia (L.) Merrill, Philip. Journ. Sci. 7: Bot. 233. 1912.

At about 900 meters in shady places; Quezaltenango. Mexico;
Salvador; Honduras; West Indies. South America.

Bulbs ovoid, 3 cm. or more in diameter, the tunics firm, red-brown; basal leaves
1-2, linear-lanceolate, 15-40 cm. long, 1-3.5 cm. wide, plicate; scape 10-30 cm.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 163

high, not winged, usually bearing a single 6-12-flowered spathe, this subtended
by a leaf similar to the basal ones; spathes about 1.5 cm. long, the flowers slender-
pedicellate; perianth white, 2-2.5 cm. broad, the segments obovate, spreading;
capsule about 2 cm. long.

Called "coquillo" in Salvador.

GELASINE Herbert

Plants small or large, from a tunicate corm; leaves plicate; flowers blue,
fugitive, several subtended by each spathe; perianth tube very short, the segments
equal, obovate; stamens inserted at the base of the perianth segments, the fila-
ments united to the apex to form a cylindric column; anthers basifixed, erect-
patent; ovary clavate, the ovules numerous, superposed; style short, subulate,
the stigmas linear, simple; capsule turbinate, loculicidally 3-valvate.

One other species is known, in Uruguay and southern Brazil.

Gelasine trichantha J. G. Baker, Bot. Jahrb. 8: 215. 1887.

Known only from the type, Sipicapa, San Marcos, F. C. Lehmann
1541; known to us only from description.

Corm globose, the tunics brown, membranaceous, produced above the neck
of the corm, the stem very slender, bearing a single head of flowers, 3.5-5 cm.
high; basal leaf 1 or none, linear, the cauline leaf 1, linear, erect, dilated and
clasping at the base; outer valve of the spathe firm, lanceolate, scarious above,
the inner valves membranaceous, the pedicels equaling the valves; ovary turbinate,
glabrous; perianth blue, the segments equal, oblanceolate, densely pilose; stamens
shorter than the perianth, the filaments very short, the anthers large, yellow;
style very short, the 3 branches patent, filiform, stigmatose at the apex, shorter
than the anthers.

Flowers of the genus Freesia (probably F. hybrida Hort.) are
sometimes seen on sale in the markets of Guatemala City, but the
plants are not common in Central America.

GLADIOLUS L. Gladiolus

Plants arising from tunicate corms; leaves distichous, narrow, mostly super-
posed on the stem, linear or ensiform; inflorescence spicate, the flowers 1 to each
spathe, sessile, of various colors, the valves of the spathe linear or lanceolate;
flowers very variable in size and color; perianth tube generally funnelform, the
segments of the limb more or less unequal in shape and direction; stamens inserted
in the throat of the perianth tube, contiguous and arching, the filaments short,
free, the anthers linear, basifixed; ovary 3-celled, the ovules numerous, super-
posed; seeds globose or discoid, sometimes winged.

About 150 species, all natives of the Old World. A number of
them are grown for ornament.

164 FIELDIANA: BOTANY, VOLUME 24

Gladiolus hortulanus L. H. Bailey, in Hortus 277. 1930.
Gladiola.

Grown commonly for ornament and often in large numbers for
sale in the markets, principally at middle or fairly high elevations;
sometimes found more or less wild in thickets, as about Coban,
also in cornfields of the highlands, perhaps there only the remnants
of commercial planting.

The gladioli of gardens are largely of hybrid origin and of con-
fused ancestry. The flowers sell well in the larger markets, as they
do in the United States, and the Indian gardeners of such places
as San Juan Sacatepe"quez and the vicinity of Quezaltenango often
plant good-sized patches of the bulbs. Gladioli thrive particularly
well about Coban and Tactic, as do most other cultivated flowers,
and in that region some fine large-flowered varieties may be seen,
probably introduced by German gardeners. The gladiolus is much
used for decorating wayside crosses and shrines because the flowers
last a long time without water.

IRISL. Iris

Perennial herbs, usually with thick, elongate, often horizontal rootstocks;
leaves linear or ensiform, thin or rather thick and fleshy, equitant; stems simple
or branched, the flowers often large and showy, 1-2 in each spathe, the outer
valves of the spathe large, ovate, herbaceous or membranous; perianth tube
cylindric, long, short, or obsolete, the segments of the 2 series more or less dis-
similar in shape and size, the 3 outer ones cuneate-obovate, reflexed in anthesis,
the 3 inner ones unguiculate, usually erect; stamens inserted at the base of the
outer perianth segments opposite the petaloid styles; filaments short, the anthers
linear, basifixed; ovary 3-celled, the ovules numerous; style branches petaloid,
with a terminal stigma and 2 deltoid petaloid crests; capsule oblong, trigonous
or hexagonous; seeds globose.

About 150 species, widely distributed, chiefly in temperate
regions. A good many species are native in temperate North
America, but none in tropical America. Many are cultivated for
their handsome flowers.

Iris germanica L. Sp. PI. 38. 1753. Lino.

Native of Europe, but grown in many temperate regions for its
handsome flowers; planted commonly in Guatemala at middle and
high elevations; naturalized in many places along roadsides and in
thickets.

Plants erect from a thick rootstock, the stems stout, usually branched, mostly
70 cm. high or less, bearing several large ensiform leaves; leaves glaucescent, 1.5-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 165

5 cm. wide, finely many-nerved, the basal ones usually shorter than the stems;
flowers almost sessile in the bracts, large and showy, deep violet-blue to pale
blue or often white, veined with yellow.

This iris is common in the gardens of Cobdn, and abundant in
many parts of the Occidente. It was noted as thoroughly naturalized
in the mountains of Quezaltenango and San Marcos. On the plains
of Quezaltenango there are many long dense rows of the plants,
bordering roads and separating fields. This is the plant to which
the name "lirio" is most commonly applied in Central America.

NEMASTYLIS Nuttall

Reference: Robert F. Foster, A revision of the North American
species of Nemastylis, Contr. Gray Herb. 155: 26-44. 1945.

Herbaceous perennials with simple or branched stems from an ovoid or sub-
globose bulb; leaves linear to lance-ensiform, often plicate; flowers blue, white,
or lavender, sometimes marked with irregular splotches and lines, fugitive, rotate,
1-several subtended by the spathes; perianth tube very short, the segments regular,
equal or unequal; stamens inserted near the base of the perianth segments, the
filaments free or more or less united; anthers longer than the filaments, with a
narrow connective coiling downward from the apex at maturity; ovary oblong-
ellipsoid, subclavate, or subturbinate; style filiform, equal to or usually shorter
than the 3 style branches, the style branches bifid, opposite the stamens; stigmas
apical, subcapitate or tufted, or minutely 2-parted; capsule oblong-ellipsoid,
dehiscent by 6 deltoid teeth at the apex.

About 15 species, all native of the New World. Of this number,
Foster limits his treatment, based upon Nuttall's concept of the
genus, to four North American species, of which only one, N. tennis,
reaches Guatemala. The remaining species from Guatemala treated
below by the present authors were all determined as Nemastylis
by Dr. Foster, but may eventually have to be transferred to other
genera, in accordance with his future monographic studies. He
considers the South American species previously assigned to Nema-
stylis to be better placed in other genera.

Perianth rotate; style shorter than style-arms or barely equal to them; anther-
connective narrow, anthers coiling downward from the apex with age.

N. tennis.

Perianth broadly campanulate to crateriform; style much longer than style-arms;
anther-connective broad, anthers seldom coiling downward from the apex
with age.

Depressed plant, 2-5 cm. tall; perianth segments strongly unequal. N. Seleriana.
Plant 10-40 cm. tall; perianth segments subequal.

Basal leaf solitary; perianth blue- violet without specks N. Iriflora.

Basal leaves usually 2; perianth blue or lilac marked with reddish-brown
specks N. silvestris.

166 FIELDIANA: BOTANY, VOLUME 24

Nemastylis Seleriana Loes. Verh. Bot. Ver. Brand. 58: 137.
1916. Nemastylis Lehmannii Standl. Field Mus. Bot. 4: 199. 1929
(type collected between Volcan de Tajumulco and Tacana, F. C.
Lehmann 1556).

Open moist meadows, 3,100-3,600 meters; endemic; Totonicapan;
Quezaltenango (type collected between Ziha and Calel, Seler 3155);
San Marcos.

Subacaulescent plant, 2-5 cm. tall, bulb ovoid-conic, the tunics dark brown,
the stem 1-2-flowered; leaves basal, 1-3, narrowly linear, subulate-acuminate,
3-10 cm. long, 1-4 mm. wide; spathe dilated at base, the expanded portion leaf-
like, linear-oblanceolate, subulate-acuminate, 2.5-4.5 cm. long; bracts 2, sub-
scarious, linear-oblong, 2.5-3.5 cm. long; peduncle very short, 0.5-2.5 cm. long;
pedicels 1-3 cm. long; ovary cylindric-obconic, 4-6 mm. long; perianth white
with dull purple blotches and transverse marks, the tube none, outer segments
ovate- or broadly elliptic-oblong, obtuse, 10-13 mm. long, 6 mm. wide, inner
segments 1 mm. long, abruptly unguiculate, the blade broadly ovate or sub-
orbicular-ovate, acute at apex, subtruncate or rounded at base, 5-6 mm. long,
5 mm. wide; filaments connate in a long narrow tube 3 mm. long, anthers linear-
oblong, 5-6 mm. long; style branches 2-parted, not equaling the length of the
anthers.

Nemastylis silvestris Loes. in Fedde, Repert. Spec. Nov. 16:
200. 1919. Nemastylis Bequaertii Standl. Journ. Arn. Arb. 11: 47.
1930 (type from Chichen Itza, Yucatan, J. Bequaert 109).

Open pine-oak woodland or clearings, near or slightly above
sea-level. British Honduras; Southern Mexico (Yucatan and
Chiapas).

Glabrous plant 3-4 dm. tall, bulb ovoid-conic, up to 2 cm. long, the tunics
dark or dull brown, the stem 4-6-flowered; leaves conspicuous, elongated, broadly
linear to narrowly lanceolate, the usually 2 basal ones long-attenuate, acuminate
at apex, narrowed at the base to an obscure petiole up to 8 cm. long, 2.8-4.7 cm.
long, 1.3-1.6 cm. wide, 5-nerved, cauline shorter, 0.6-1 cm. wide, inserted about
or above the middle; spathes subcymbiform, somewhat rounded at base, rather
sharply pointed at apex, 2-3 cm. long, long-pedunculate; bracts very unequal,
erect, the interior twice shorter or sometimes almost equaling the outer, 2-3 cm.
long, long-attenuate at apex; peduncle erect or ascending, slender, 4-8 cm. long;
pedicels slender, 2-2.5 cm. long, included or shortly exserted; flowers glomerulate;
ovary oblong or obconic-subovoid, 5 mm. long; perianth blue or lilac, marked
with reddish-brown specks, 9-10 mm. long, the tube almost none, the segments
obovate or cuneate-obovate, broadly rounded at the apex; filaments about 2.5 mm.
long, anthers linear, straight, 4-5 mm. long.

Nemastylis tenuis (Herb.) Baker, Handb. Irid. 112. 1892.
Nemastylis coelestina var. tenuis Herb, in Bot. Mag. 66. pi. 3779.
1840. Chlamydostylis tenuis (Herb.) Baker, in Journ. Bot. 14: 185.
1876.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 167

Open oak or pine slopes, 1,000-1,800 meters; Jalapa; Santa Rosa.
Mexico north to Chihuahua and Coahuila.

Bulb ovoid to subglobose, up to 2.5 cm. long, 2 cm. wide, the tunics castaneous
or dark brown, the stem terete, 1-5-flowered, 1-3 dm. tall, glabrous; basal leaves
2-3, linear, acute, up to 25 cm. long, 1-3 mm. wide, often scabrous on the margins,
cauline shorter, 6-9 cm. long; spathes unequal, the outer 1-2 cm. long, acute, the
inner up to 2.5 cm. long; pedicels included at anthesis or slightly exserted; ovary
ovoid or ellipsoid, 3-4 mm. long; perianth blue, segments subequal, obovate,
obtuse, up to 20 mm. long, 6-8 mm. wide; filaments entirely united, 1.5-2.5 mm.
long, anthers 6-7 mm. long; style equaling the filament column, style branches up
to 4 mm. long; capsule oblong-ellipsoid or turbinate, up to 1.5 cm. long; seeds
dark brown, more or less pyriform, 2 mm. long.

The Guatemalan specimens are referred by Dr. Foster to typical
N. tennis, which is distinguished from most of the other varieties
in having the filaments entirely united.

Nemastylis triflora Herb, in Benth. PI. Hartw. 95. 1842.

Known only from the type, oak woods in the mountains of
Comalapan, Hartweg 625.

Bulb ovoid, 1.8 cm. wide, the stem 3-flowered, 3-4.5 dm. tall; basal leaf
solitary, linear, acute, up to 30 cm. long, 5-6 mm. wide, very plicate, cauline
shorter; spathes subequal, acuminate at apex, up to 5 cm. long; perianth blue-
violet, 18-25 mm. long, the segments broadly cuneate-obovate, the larger erose;
anther about 5.5 mm. long; style branches filiform, 8 mm. long.

This species is known to the authors only from description.

NEOMARICA Sprague

Plants perennial, from short rhizomes; leaves broadly linear, ensiform, dis-
tichous, flabellate, with a prominent costa; peduncle compressed and leaf-like,
with 1 or few subsessile or pedunculate flower clusters, the peduncle long-produced
beyond the inflorescence; flowers fugitive, blue, yellow, or white; perianth tube
obsolete, the segments of the 2 rows very unlike, the outer ones obovate, spreading,
the inner ones much smaller, panduriform, convolute; stamens short, erect, the
filaments distinct; ovary clavate, 3-celled, the ovules numerous, superposed;
style subulate at the base, cyathiform with 3 angles in the upper half to which
the anthers adhere loosely; style crests lanceolate, with the transverse stigmas
at their base; capsule oblong, loculicidally 3-valvate; seeds subglobose or angled
by pressure, with a brown testa and corneous endosperm, surrounded by a fleshy
aril.

About ten species, in tropical America.

Neomarica gracilis (Herb.) Sprague, Kew Bull. 1928: 280.
1928. Marica gracilis Herb. Bot. Mag. pi. 3713. 1839. Cucqubn
(Quecchi).

FIG. 31. Neomarica gradlis. A. Habit of plant; X 1 A. B. Flower; X 1.
C. Androecium and gynoecium in position, with perianth removed; X 3.

168

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 169

Mostly in dense wet forest, 900 meters or less; Pete*n; Alta Vera-
paz; Izabal; Huehuetenango. Southern Mexico; British Honduras,
along the Atlantic coast to Panama; southward to Brazil.

Plants erect, or weak and reclining; leaves 30-80 cm. long, thin, deep green,
1.5-2.5 cm. wide, with a conspicuous elevated costa; peduncle about as long as
the leaves, flat and leaf-like, bearing a cluster of flowers far below the apex, simple,
the tip of the leaf often taking root in the soil; flower clusters sessile or nearly so,
the spathes about 3.5 cm. long, the outer valves green, lance-acuminate; open
flowers 5 cm. broad, delicate, the outer segments obovate, pale yellow, sometimes
white, with cross bars of yellow and brown near the base, the small inner segments
blue; capsule about 2.5 cm. long, the seeds subtended by a red aril. (Fig. 31.)

In Alta Verapaz the Indians administer a decoction of the
crushed roots as a supposed remedy for affections of the stomach.

ORTHROSANTHUS Sweet

Reference: Julian A. Steyermark, Orthrosanthus chimboracensis
and its varieties, Lloydia, 11: 14-20. 1948.

Plants perennial from short thick rootstocks, in general appearance like Iris;
leaves rather thick and firm, linear, equitant, many-nerved, glabrous; flower
clusters few or numerous, many-flowered, paniculate; flowers pale blue, fugitive,
the pedicels usually very short; perianth tube very short, the segments oblong,
subequal, spreading; stamens inserted at the base of the perianth, the filaments
free or connate at the very base; anthers linear, erect; ovary clavate, 3-celled, the
ovules numerous, superposed; style very short, the branches subulate, stigmatose
at the apex; capsule oblong-trigonous, loculicidally 3-valvate.

About seven species, two in the mountains of tropical America,
the others in western Australia. Only the following occurs in
Central America.

Orthrosanthus chimboracensis var. centro-americanus

Steyermark, in Lloydia, 11: 17-18. 1948. Gladiolin de monte (fide
Aguilar).

Brushy or grassy, often rocky hillsides, common in pine-oak
forest, 1,500-3,400 meters; Zacapa; Jalapa; Jutiapa; Sacatep^quez;
Suchitepequez; Solola; Quiche"; Huehuetenango; Totonicapan; Que-
zaltenango; San Marcos. Southern Mexico; Costa Rica.

Rhizome usually short and stout but sometimes elongate and rather slender;
leaves numerous, pale bluish green, 20-50 cm. long, 1 cm. wide or less; scape not
winged, 25-60 cm. high, leafy below, simple or branched, bearing few or numerous,
sessile or pedunculate spathes; bracts of the spathe 0.8-1.3 cm. long, 3-4-flowered;
perianth pale blue, very delicate, the segments 1-1.5 cm. long; capsule more or less
pubescent, 0.8-1.6 cm. long. (Fig. 32.)

FIG. 32. Orthrosanthus chimboracensis var. centro-americanus. A. Habit of
plant; X M- B. Seed; X 20. C. Capsule; X 3. D. Androecium and gynoecium
in position; X 3. E. Upper portion of style and branches; X 7. F. Stamen; X 8.

170

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 171

A rather handsome, iris-like plant, often forming large dense
colonies.

Typical 0. chimboracensis (HBK.) J. G. Baker is confined to the
Andes of South America and the volcanoes of Costa Rica. It has
glabrous ovaries and capsules, the latter larger than in the Central
American variety.

RIGIDELLA Lindley

Plants perennial from a corm having membranous tunics; leaves broad, plicate,
the stems slender, branched; pedicels protruded from the spathes; flowers fugitive,
red; perianth tube none, the segments very unequal, the outer ones oblong, conni-
vent into a cup in the lower third, then spreading or reflexed, the inner ones very
small, erect, ovate, with a narrow claw; filaments united throughout into a cylindric
column, the anthers linear, ascending; ovary clavate, 3-celled, the ovules numerous,
superposed; style slender, bifid above the middle; capsule oblong, 3-valvate toward
the apex; seeds subglobose, with a conspicuous raphe.

In southern Mexico one other species is known, R. flammea Lindl.,
from which Baker states that the Guatemalan species may not be
distinct.

Rigidella immaculata Herb, in Lindl. Bot. Reg. 27: pi. 68.

1841.

Dense or open forest of oak, pine, Juniperus, or Abies, sometimes
in moist meadows, 1,700-3,500 meters; described originally from
plants grown in England, the corms collected in the mountains of
Guatemala by Hartweg; Sacatepe*quez (Volcan de Agua); Chimal-
tenango(?); Huehuetenango. Southern Mexico.

Plants sometimes a meter high or nearly so, usually lower, arising from a
small corm covered with thin brown tunics; leaves few, broadly linear, mostly 1-3
cm. wide, conspicuously plicate and with numerous conspicuous nerves; peduncles
bearing 2 or more spathes, not winged; spathes several-flowered, 5-8 cm. long;
outer perianth segments blood-red, 3-3.5 cm. long, oblong, the inner segments
yellow, ovate, unguiculate, very small; capsule 2.5-3 cm. long.

SISYRINCHIUM L.

Plants annual or perennial, the rhizomes none or very short, the roots fibrous
or often thickened and fleshy; leaves linear or narrowly ensiform; peduncles
flattened or terete, often winged, leafy or naked; flowers blue or yellow, small,
usually several in a cluster, the clusters solitary or fasciculate, rarely spicate or
paniculate; capsules exserted from the spathe; perianth tube very short or none,
the segments oblong, subequal, spreading from above the base; stamens inserted
at the base of the perianth, the filaments more or less connate, the anthers erect
or versatile; ovary turbinate or globose, 3-celled, the ovules numerous, super-

172 FIELDIANA: BOTANY, VOLUME 24

posed; style short, subulate, the branches subulate, stigmatose at the apex; capsule
subglobose or turbinate, loculicidally 3-valvate; seeds numerous, minute.

Species number uncertain but sixty or more, in temperate and
tropical America; in tropical regions found mostly in the mountains.
The species are often difficult of segregation, and the whole genus
is badly in need of critical study. In temperate North America the
number of species has been fantastically increased, but from Central
America, fortunately, comparatively few species have as yet been
published.

Peduncles simple, each bearing a single spathe or cluster of spathes.

Perianth yellow; peduncles rather broadly winged; leaves thin and delicate.

S. tinctorium.
Perianth blue or white; peduncles not winged or scarcely so; leaves rather thick

and stiff S. Johnstonii.

Peduncles branched above, bearing 2 or more separated spathes.

Capsules about 3 mm. high; plants annual, the root fibers very slender; perianth

blue or bluish S. micranthum.

Capsules 5-20 mm. high; plants perennial, the roots usually much thickened and
fleshy; perianth yellow.

Valves of the spathe 2.5-3 cm. long; capsule 12-20 mm. long.S. convolutum.

Valves of the spathe mostly 2 cm. long or shorter; capsule usually less than
8 mm. long.

Peduncles narrowly winged; cauline leaves somewhat dilated at the base.

S. guatemalense.

Peduncles not winged but usually sharp-edged; cauline leaves not dilated
at the base S. tenuifolium.

Sisyrinchium convolutum Nocca, PL Select. Hort. Ticin. sub
pi. 1. 1800.

Moist forest or meadows, often in open pine forest, sometimes a
weed in cornfields, 1,300-3,350 meters; Zacapa; Chiquimula; El
Progreso; Jalapa; Sacatepe'quez; Quiche"; Huehuetenango. Hondu-
ras; Panama; western South America, southward to Peru.

Plants arising from a dense cluster of roots, these slightly thickened but not
tuber-like, the plants 30-75 cm. high or sometimes taller, the stems branched,
conspicuously winged; leaves blackening when dried, rather thick and firm, mostly
5-8 mm. wide; spathes several, 4-6-flowered, the pedicels short-exserted; valves
of the spathe mostly 2.5-4 cm. long, somewhat inflated and enlarged below;
perianth yellow, as much as 2.5 cm. broad, the segments 12 mm. long or more;
capsule ellipsoid, as much as 2 cm. long and 1.5 cm. broad but usually somewhat
smaller.

This has been confused with the South American S. iridifolium
HBK., whose actual status is somewhat uncertain. At any rate,
S. convolutum is a much earlier name, and may serve satisfactorily
to designate the plant here described.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 173

Sisyrinchium guatemalense (J. G. Baker) Standl. & Steyerm.
Field Mus. Bot. 23: 39. 1944. S. alatum Hook. var. guatemalense
J. G. Baker, Handb. Irid. 130. 1892 (type from mountains of Guate-
mala, the locality and collector not indicated).

Moist meadows or open woods, 750-2,100 meters; Santa Rosa;
Guatemala; Chimaltenango; Solola; Quiche"; Quezaltenango. South-
ern Mexico; Honduras; Costa Rica.

Plants arising from a dense cluster of very thick and fleshy, tuber-like roots,
generally low and 20 cm. high or less but sometimes 50 cm. high; basal leaves few,
usually blackening when dried, often 15-20 cm. long and 5 mm. wide; stems 1-
several, often tortuous, narrowly winged, bearing 2-several spathes, the spathes
long-pedunculate, 2-5-flowered, the slender pedicels partly exserted; valves of the
spathe 1.5-2 cm. long, relatively broad, somewhat dilated below, scarious-margin-
ate; perianth bright yellow; capsule subglobose, about 8 mm. long.

While treated by Baker merely as a variety of S. alatum Hook.
(S. Marchio Steud.), this is so different in general appearance from
South American material of that species that there can be little
doubt that a distinct species is represented. It may be that some
neglected older name for the plant, based on Mexican collections,
will be discovered later, but we have found none.

Sisyrinchium Johnstonii Standl. Field Mus. Bot. 17: 229. 1937.

Moist or wet, chiefly alpine meadows, 2,500-4,000 meters;
Sacatepe"quez (type from crater of Volcan de Agua, J. R. Johnston
805); Huehuetenango (Sierra de los Cuchumatanes) ; San Marcos
(volcanoes of Tajumulco and Tacana). Chiapas (Volcan de Tacana).

Plants stiffly erect from a dense cluster of small tuberous-thickened roots,
the weathered fibers of old leaves often persistent at the base of the plant; leaves
all basal, linear, thick and rather stiff, sometimes 17 cm. long but usually much
shorter, about 2.5 mm. wide, erect; scapes stiff, longer or shorter than the leaves,
bearing at the apex 1-2 sessile spathes, the spathe or pair of spathes subtended
by an erect bract; valves of the spathe about 2 cm. long; flowers 1-few, the pedicels
rather long-exserted ; perianth white with bluish lines, the segments 1 cm. long or
more.

Sisyrinchium micranthum Cav. Monad. Diss. 6: 345. pi. 191,
f. 2. 1790.

Moist or wet fields, pastures, or thickets, often on open or brushy
banks, sometimes on sandbars along streams, 300-2,400 meters;
Alta Verapaz; Baja Verapaz; Chiquimula; Jalapa; Guatemala;
Sacatepe"quez; Chimaltenango; Huehuetenango; Quezaltenango; San
Marcos. Southern Mexico; Honduras; Costa Rica; Panama; South
America.

FIG. 33. Sisyrinchium micranthum. Habit of plant; X

174

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 175

Plants very slender, annual, with a cluster of very slender, fibrous roots, erect
or ascending, often much branched from the base, the stems mostly 5-20 cm. high,
sharp-edged but scarcely winged; leaves linear, 2-4 mm. wide, the basal ones few
or numerous, much shorter than the stems, the cauline leaves shorter; spathes
few or rather numerous, solitary, long-pedunculate, 2-6-flowered, the pedicels
partly exserted; valves of the spathe 2 cm. long or less, very narrow, not dilated
below; perianth blue or white, often pale dirty bluish, the segments 6 mm. long
or less; capsule globose, usually not more than 3 mm. in diameter. (Fig. 33.)

An inconspicuous and rather weedy plant with unattractively
colored flowers.

Sisyrinchium tenuifolium Humb. & Bonpl. ex Willd. Hort.
Berol. 2: pi. 92. 1816.

Grassy open slopes or meadows, often in alpine meadows, some-
times in forest of pine and Abies, 1,600-3,500 meters; Chiquimula;
Jalapa; Sacatepe"quez; Chimaltenango; Quiche 1 ; Huehuetenango;
Quezaltenango. Mexico.

Plants erect or ascending from a dense cluster of tuberous-thickened roots,
the stems usually several, 6-30 cm. high or even taller, very slender; leaves usually
shorter than the stems, 1-4 mm. wide; spathes generally 2 or more, long-peduncu-
late, 3-4-flowered, the outer spathe valves 1-2 cm. long, often somewhat dilated
below; perianth yellow, 6-9 mm. long; capsule oblong or oval, 5-8 mm. long.

Sisyrinchium tinctorium HBK. Nov. Gen. & Sp. 1: 324. 1815.

Moist or wet fields or open wet banks, 750-3,500 meters; Alta
Verapaz; Solola; Quezaltenango; Huehuetenango. Southern Mexico;
British Honduras; Costa Rica; Panama; South America.

Plants perennial or perhaps sometimes annual, from a cluster of slender
fibrous roots; stems naked, weak, often numerous, 20-40 cm. high, broadly winged;
leaves thin and soft, 3-10 mm. wide, sometimes equaling the stems, often numerous;
spathe 1 on each stem, 4-10-flowered, the outer spathe valve 3-5.5 cm. long;
pedicels very slender and usually long-exserted ; perianth yellow, 8-12 mm. long;
capsule oval or obovoid, 8-12 mm. long, or sometimes almost 2 cm. long.

TIGRIDIA Jussieu

Rootstock an ovoid corm with brown membranous tunics; leaves linear or
lance-linear, plicate; stems terete, not winged, simple or branched; flowers fugi-
tive, often very large; perianth tube none, the segments of the 2 series dissimilar,
connivent in a cup at the base, then spreading; filaments united to the apex in
a long cylindric column, the anthers linear, erect-patent; ovary clavate, 3-celled,
the ovules numerous, superposed; style long, filiform, its branches divided into
2 slightly flattened, falcate forks; capsule clavate-oblong, shortly 3-valvate at
the apex; seeds angulate by pressure.

Species ten or fewer, one in South America, the others Mexican,
one of them extending to Guatemala and Costa Rica.

176 FIELDIANA: BOTANY, VOLUME 24

Tigridia Pavonia (L. f.) Ker in Konig & Sims, Ann. Bot.
1:246.1805. Ferraria Pavonia L. f. Suppl. PL 407. 1781. Cebollin;
the name Cochol is reported from Guatemala.

Moist forest, sometimes in oak forest, swampy thickets, or open
meadows; often a weed in cultivated ground, especially cornfields,
1,600-3,000 meters; Jalapa; Chimaltenango; Quiche"; Huehuete-
nango; Quezaltenango; San Marcos. Mexico; Salvador (probably
only in cultivation) ; introduced into Costa Rica as a weed in corn-
fields; often grown for ornament in temperate or warm regions of
other parts of the earth.

Plants erect from a rather large corm; basal leaves long-sheathing, several-
nerved, linear-lanceolate, 1-2.5 cm. wide; stems leafy, mostly 35-60 cm. high,
leafy; spathes 1-2, few-flowered, 6-10 cm. long, the flowers long-pedicellate;
outer perianth segments obovate, about 7 cm. long, orange-red above, pale yellow
heavily spotted with dull red below; inner perianth segments pale yellow, the basal
part heavily spotted with dull red, the apical portion more lightly spotted; capsule
3-5 cm. long, 1 cm. broad.

Called "flor del tigre" in Salvador; "Guatemala" (Costa Rica).
The Nahuatl name of Mexico is "oceloxochitl," "tiger" or "ocelot
flower," given because of the spots on the perianth segments. The
roots were formerly much used in the mountains of Mexico as food,
and are said to have a flavor similar to that of chestnuts. In Hue-
huetenango and doubtless elsewhere the mucilaginous sap of the
corms is used like glue for joining small articles. The plant is an
exceptionally showy and handsome one, and is often planted in
Guatemalan gardens. It flowers only during the wet months, no
traces of the plants appearing in the dry season. In some areas,
as about Chimaltenango and in Huehuetenango, it is an abundant
weed in cornfields.

TRIMEZA Salisbury

Plants perennial from a short thick erect rootstock, the peduncle bearing a
few long-stalked spathes; leaves usually broadly linear, not plicate, thin, with a
conspicuous costa; perianth tube none, the outer and inner segments very dis-
similar, the outer ones obovate, with a broad concave claw, the inner ones much
smaller, convolute, with a small deflexed blade; stamens short, erect, the filaments
free; ovary clavate, 3-celled, the ovules many, superposed; style subulate at the
base, the 3 branches ending in small tubercles or cusps, these over-topping the
small horizontal stigmas; capsule oblong, loculicidally 3-valvate.

About six species, in tropical America, only one of them in Central
America.

Trimeza martinicensis (Jacq.) Herb, in Lindl. Bot. Reg. 1844 :
Misc. 88. 1844. Iris martinicensis Jacq. Enum. PI. Carib. 12. 1760.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 177

T. lurida Salisb. Trans. Hort. Soc. 1: 308. 1812. Purga de tos frios
(Huehuetenango); Sholol-canti (Cubilguitz, Alta Verapaz).

Mostly in dense wet mixed forest, 200-1,600 meters; Alta Vera-
paz ; Huehuetenango. Southern Mexico ; West Indies ; South America.

Plants arising from small fibrous-coated corms; basal leaves linear, much
elongate, 1-2 cm. wide, thin and rather soft, with a conspicuous costa; stems
subterete, not winged, 30-80 cm. high, simple or branched, usually with a large
leaf at the base of the inflorescence; spathes few, on very long peduncles, many-
flowered, the pedicels usually partly exserted, the bracts about 2 cm. long; outer
perianth segments 2 cm. long, spreading, deep or bright yellow with purple-brown
spots in the lower third; inner perianth segments erect-ascending, with brown-
purple spots at the base and along the middle; anthers yellow, the connective
purple-brown; capsule 12-20 mm. long.

In Huehuetenango it is stated that a decoction of the root is ad-
ministered as a domestic remedy for chills.

TRITONIAKer

Rootstock a corm with fibrous or reticulate tunics; leaves mostly linear
and plicate; flowers spicate, the spikes simple or branched; valves of the spathe
oblong, brownish, emarginate; perianth tube short or elongate, dilated above, the
segments obovate or oblong, subequal or somewhat unequal; stamens unilateral,
arcuate, parallel, inserted in the perianth tube; filaments filiform, the anthers
usually versatile; ovary 3-celIed, the ovules superposed; style filiform, the branches
simple, short, spreading; capsule small, oblong, membranous, loculicidally 3-
valvate; seeds small, globose, or angulate by pressure.

About thirty species, mostly native in South Africa.

Tritonia crocosmiiflora Nichols. Gard. Diet. 4: 1887. Mont-
bretia crocosmiiflora (often written crocosmiflora and crocosmaeflora)
Hort. Fl. Mag. n. ser. pi. 472. 1881. Ccam (Coban, Quecchi).

Planted commonly for ornament at almost all elevations;
thoroughly naturalized in some localities about Cobdn, often form-
ing large colonies in thickets or on open banks.

Plants usually large and erect, a meter high or less, from a fibrous-coated
corm; leaves ensiform, often numerous, 1-2 cm. wide, with a conspicuous elevated
costa; stems not winged, leafy below, paniculately branched above, the spathes
small and calyx-like, 8 mm. long; perianth orange-red, 3-4 cm. long, the tube very
slender below, strongly curved, the lobes oblong, spreading, about equaling the
tube.

Called "zacatillo" and "iris" in Salvador. This plant is a
bigeneric hybrid, the result of a cross between two African plants,
Crocosmia aurea Planch, and Tritonia Pottsii Benth. It may be
remarked, however, that some authors consider the two generic

178 FIELDIANA: BOTANY, VOLUME 24

names synonymous. The plant is a showy and rather handsome
one, popular throughout Central America, thriving under almost
all conditions with little or no care. It is thoroughly established
in some places about Coban, possibly on the sites of former dwellings,
and appears well able to maintain itself in competition with native
vegetation.

MUSACEAE. Banana Family
Reference: K. Schumann, Pflanzenreich IV. 45. 1900.

Perennial herbs, often very large and treelike; stems at first or always very
short, much surpassed by the leaf sheaths, these often tightly rolled together
and forming a false trunk; leaves distichous or spirally arranged, large, petiolate,
the petiole vaginate, the sheath without a ligule, the blades broadly linear to
oblong, obtuse, pinnately nerved; inflorescence simple, bracteate, usually spike-
like, sometimes branched, the flowers sessile or pedicellate, bracteolate or ebracteo-
late; flowers zygomorphic, usually perfect; sepals equal or sometimes separated
into sepaloid and petaloid series, free or somewhat united, imbricate, colored;
fertile stamens 5, rarely 6, the highest often reduced to a staminode; anthers
narrowly linear, 2-celled, the cells dehiscent by longitudinal slits; ovary inferior,
3-celled; ovules solitary in each cell or several, anatropous, erect or horizontal;
style simple, the stigma deeply trilobate or simple or capitate; fruit baccate or
capsular; seeds very hard, sometimes surrounded by an aril.

Six genera, only two of which, Heliconia and Ravenala, are repre-
sented by species native in America. Only one genus is native in
Central America. The treatment of the family by Schumann,
constituting the first number of the Pflanzenreich, is now long out
of date, hence of limited value for study of American plants of the
family.

Leaves spirally arranged; flowers unisexual Musa.

Leaves distichous; flowers perfect.

Cells of the ovary 1-ovulate; capsule separating into 3 cocci; seeds not arillate;

native plants Heliconia.

Cells of the ovary several-oyulate; capsule loculicidally 3-valvate; seeds sur-
rounded by an aril; cultivated plants.

Inner equal sepals free but forming a sagittate organ; plants low, herbaceous.

Strelitzia.

Inner equal sepals not forming a sagittate organ; plants very large and tree-
like . . . . Ravenala.

HELICONIA L.

References: Robert F. Griggs, On some species of Heliconia, Bull.
Torrey Club 30: 641-664. pis. 29, 30. 1903; Some new species and
varieties of Bihai, Bull. Torrey Club 42: 315-330. pi. 19. 1915.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 179

Large or small, coarse, perennial herbs; leaves small or large, distichous,
petiolate, the petioles vaginate; inflorescence terminal, compound, with large and
usually brightly colored, cymbiform bracts subtending the several-flowered indi-
vidual inflorescences; flowers perfect; outer sepals more or less unequal, the
posterior one largest, the 2 anterior ones equal, narrower, free or united with the
interior perianth segments to form a 5-dentate boat-shaped organ; perfect stamens
5, the anthers linear, basifixed, the cells introrse; staminode petaloid, small;
ovules solitary in each cell, erect from its base; style filiform, the stigma clavate
or subclavate, 3-lobulate; capsule usually blue, sometimes by abortion 1-2-celled,
separating into usually 3 cocci; seeds obtusely trigonous, the embryo straight.

Probably fifty species, natives of tropical America. Several
others besides those listed here occur in other parts of Central
America, and the genus extends northward into southern Mexico.
The larger plants are somewhat like the banana in habit, the smallest
ones more suggestive of the genus Canna. They are confined to the
lowlands in Central America, never extending far upward on the
mountain slopes. In some parts of the tierra caliente they constitute
an important and conspicuous part of the undergrowth in the forest
or of the coarse second growth thickets, forming colonies of wide
extent. The concave bracts of those species with erect inflorescences
hold water, and it has been stated that mosquitoes breed in them.

Bracts of the inflorescence densely crowded and overlapping, appressed, concealing
the rachis. Plants very large, often 5 meters tall or more; inflorescence very
large and heavy, pendent, often a meter long or more; bracts about as broad

as long, deep red H. Mariae.

Bracts of the inflorescence not or scarcely overlapping, mostly spreading or

reflexed, not concealing the rachis, usually much longer than wide.
Inflorescence pendent, the bracts deep red.

Rachis of the inflorescence densely brown- villous; leaves green beneath.

H. rostrata.
Rachis of the inflorescence puberulent or closely tomentulose or sometimes

practically glabrous H. Collinsiana.

Inflorescence erect, the bracts variously colored.

Rachis of the inflorescence mostly 1-1.5 cm. thick; bracts close together, the
top of one usually reaching the base of the one next above, the middle
bracts mostly 3-6 cm. high at the point of attachment, the bracts rela-
tively short and broad H. Bihai.

Rachis of the inflorescence usually much less than 1 cm. thick; bracts rather
widely spaced, not approximate, the middle bracts mostly 1.5-3 cm.
high, long and narrow.

Plants small, generally about a meter high; leaves mostly 10 cm. wide or
less, sometimes broader; bracts relatively few and short.

Bracts deep red H. subulate.

Bracts yellow or orange H. psittacorum.

Plants larger, mostly more, often much more, than a meter high; leaves
usually much more than 10 cm. wide; bracts often numerous and usually
elongate.
Inflorescence persistently villous, especially on the rachis.

Bracts yellow H. spissa.

Bracts deep red H. Schiedeana.

180 FIELDI AN A: BOTANY, VOLUME 24

Inflorescence glabrous or nearly so, at least not villous.

Inflorescence elongate, the bracts broad, widely spreading or even
reflexed, not curved inward or upward. Bracts red.H. latispatha.
Inflorescence short and rather deltoid, the bracts very narrow, distinctly
curved upward and inward.

Bracts yellow 7 H . librata.

Bracts deep red H. adflexa.

Heliconia adflexa (Griggs) Standl. Journ. Wash. Acad. Sci. 17:
162. 1927. Bihai adflexa Griggs, Bull. Torrey Club 42: 325. /. 5.
1915.

Known only from Alta Verapaz, dense wet forest, 1,400-1,650
meters; type from Coban, Turckheim 11.2356.

Plants 1.5-2.5 meters tall; leaves large and elongate, commonly 15-20 cm.
wide, short-acuminate, rounded at the base, green above, slightly paler beneath
but not glaucous; inflorescence stiffly erect, long-pedunculate, somewhat deltoid
in outline, the rachis rather stout, slightly or rather strongly zigzag, densely
puberulent, with very short internodes; bracts deep red, linear, close together,
usually 12 or more, ascending and often somewhat incurved, obtuse, sparsely
puberulent or glabrate, revolute, the middle bracts 7-10 cm. long, 1.5-2 cm.
high, the upper ones much shorter; perianth 4 cm. long, yellow, sparsely pilose.

The species is well marked because of the compact inflorescence
and the long narrow bracts.

Heliconia Bihai L. Mant. PL 2: 211. 1771. H. elongata Griggs,
Bull. Torrey Club 30: 653. /. 2. 1903 (type from Rio Polichic, between
Panzos and Sepacuite", Alta Verapaz, R. F. Griggs 790). H. Champ-
neiana Griggs, op. cit. 657. pi. 30 (type collected between Panzos
and Sepacuite*, Griggs 528). Bihai Champneiana Griggs, Bull. Torrey
Club 31: 445. 1904. B. elongata Griggs, loc. cit.

Wet forest or thickets of the Atlantic lowlands, 900 meters or
lower; Pete"n; Alta Verapaz; Izabal; Huehuetenango. Southern
Mexico; British Honduras, along the Atlantic coast to Panama;
West Indies; South America; said to be naturalized in some parts
of the Old World tropics.

Plants glabrous, coarse and stout, 1.5-4 meters tall, usually with a well
developed, stout stem; leaf blades elongate-oblong, usually 20 cm. wide or more,
cuspidate-acuminate, rounded to acute at the base, green on both sides or when
young slightly glaucescent, sometimes 2 meters long; inflorescence erect, sessile
or nearly so, very thick, succulent, and heavy, oblong in outline; bracts 9-12 or
more, boat-shaped, spreading, close together, usually touching one another, the
margins apple-green, the sides bright red or orange, shading into yellow, the
middle ones 13 cm. long, the lower much longer, attenuate; flowers about 3 cm.
long, the segments bright green, linear-oblong; fruit turquoise-blue. (Fig. 34.)

FIG. 34. Heliconia Bihai. A. Inflorescence; X H- B. Habit of plant; X V-
C. Flower with bract; X 2. D. Interior perianth segments from within, showing
stamens; X 1.

181

182 FIELDIANA: BOTANY, VOLUME 24

This is perhaps the handsomest species of Central America and
certainly one of the showiest. The coloring of the bracts is more
vivid and fresh than that of most other species. The plant often
forms large dense stands in the lowlands of Alta Verapaz and Izabal,
and it is frequent in many localities all along the Atlantic coast of
Central America. There are many fine displays of it visible from
the railway in the Atlantic banana region of Guatemala.

Heliconia Collinsiana Griggs, Bull. Torrey Club 30: 648.
1903. Cachuco (Santa Rosa); Platanillo. Bihai Collinsiana Griggs,
Bull. Torrey Club 31: 445. 1904.

Wet forest or thickets, chiefly at 600 meters or less, rarely ascend-
ing to about 2,000 meters; type from banks of Rio Cahabon, 5 miles
below Cahabon, Griggs 352; Alta Verapaz; Zacapa; Chiquimula;
Santa Rosa; Escuintla; Suchitepequez; Retalhuleu; Quezaltenango;
San Marcos; Huehuetenango. British Honduras; probably extend-
ing into southern Mexico; Salvador.

Plants 2-3 meters tall or sometimes as much as 5 meters; leaves long-petiolate,
the blades narrowly oblong, often a meter long or more and as much as 40 cm. wide,
short-acuminate, rounded and often unequal at the base, usually very glaucous
and pale beneath, sometimes puberulent or short-pilose beneath on the costa,
the under surface sometimes becoming green in age; inflorescence pendent, deep
or bright red, glabrous or often densely puberulent or short-pilose, especially on
the rachis, pedunculate, the rachis flexuous or almost zigzag, often 45 cm. long,
the bracts widely spaced, lanceolate, spreading or even reflexed, long-attenuate,
the lowest 30 cm. long and about 2 cm. high at the base, the middle ones about
15 cm. long; flowers long-pedicellate, pale yellow; fruit pale yellow or reddish.

Sometimes known in Salvador by the name "hoja de sal,"
presumably because the leaves are used for wrapping salt and
other articles for market. The handsome inflorescences are sold
in the markets to be used for decorating altars, especially at Christ-
mas time, when they are carried from the lowlands up into the high-
lands. The species has been listed from Guatemala under the
name H. vaginalis Benth. Here is probably referable Guatemalan
material reported under the name Heliconia platystachys Baker,
which has broader bracts. The material here referred to H. Col-
linsiana is somewhat variable in pubescence of the inflorescence and
in color of the lower leaf surface, so that it is possible although not
probable that two species are represented.

Heliconia latispatha Benth. Bot. Voy. Sulph. 170. 1844.
Pico de gurrion (Santa Rosa) ; B^'ao (Pete"n); Platanillo; Sue (Quec-
chi).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 183

Wet forest or thickets, often abundant in second growth, ascend-
ing from sea level to about 1,400 meters, but most abundant at low
elevations; Pete"n; Alta Verapaz; Izabal; Santa Rosa; Escuintla;
Suchitepequez; Retalhuleu. Southern Mexico; British Honduras to
Panama; Colombia.

Plants glabrous or nearly so, rather stout, commonly 1.5-2.5 meters high;
leaves long-petiolate, oblong, often a meter long, mostly 20-30 cm. wide, short-
acuminate, rounded to subacute and usually oblique at the base, slightly glaucous
or green beneath; inflorescence erect, pedunculate, the bracts deep or bright red,
widely spaced, narrowly lanceolate, spreading, the lowest often dilated at the apex
into large green blades, often tinged with orange or yellow, or sometimes yellow
or orange throughout, the middle ones about 15 cm. long and 1.5-2 cm. high at
the base, long-attenuate; flowers 3-3.5 cm. long, pedicellate, the pedicels glabrous
or pilose, the perianth greenish yellow.

The inflorescences are sometimes called "cuchillos" in Salvador.
On the Pacific slope this species is particularly plentiful, growing
not only in ravines of the foothills but far out upon the plains, in
either forest or open places. Some of these habitats become very
dry in the verano but are doubtless exceedingly wet during the rainy
season.

Heliconia librata Griggs, Bull. Torrey Club 30: 649. 1903.
Bihai librata Griggs, Bull. Torrey Club 31 : 445. 1904.

Type from "Cherujija Oxec," valley of Rio Oxec, Alta Verapaz,
Griggs 696. Tabasco; Atlantic coast of Honduras.

Plants 1.5-3 meters tall; leaves long-petiolate, the larger ones a meter long
or more and about 30 cm. wide, abruptly short-acuminate, rounded and some-
what decurrent at the base, glabrous, glaucous beneath or in age merely pale
green; inflorescence erect, long-pedunculate, somewhat deltoid in outline, the
bracts close together, usually 12-16, spreading or slightly ascending, ovate,
the lowest often prolonged into a large green blade, the middle ones ovate or
ovate-lanceolate, attenuate to an obtuse tip, 10 cm. long or usually shorter, yellow
or orange; pedicels 1-1.5 cm. long, puberulent; rachis stout, almost straight, densely
puberulent.

The color of the inflorescence was unknown when the species
was described, for it was based upon aged specimens, and there is
some uncertainty regarding the identity of the material here referred
to H. librata. In Honduras the species is sometimes called "bija-
giiillo."

Heliconia Mariae Hook. f. Journ. Linn. Soc. 7: 69. 1864.
Usually in dense wet forest near sea level; Alta Verapaz; Izabal.
British Honduras, along the Atlantic coast to Panama; Colombia.

184 FIELDIANA: BOTANY, VOLUME 24

Plants very large, often as tall as a banana plant and attaining a height of
12 meters or even more, with a thick heavy trunk like that of a banana plant; leaves
long-petiolate, the blades long and very large, glabrous, pale green beneath; in-
florescence pendent, very thick and heavy, oblong, usually 30 cm. long or more
and 8-12 cm. wide, but said to attain sometimes a length of even 2 meters; bracts
very numerous, broadly ovate, obtuse, close together and densely imbricate, spread-
ing at about a right angle, deep rose-red or in age deep dark red, thinly villous-
tomentose; flowers dull red, the pedicels villous.

Called "bijagiiillo" in Honduras. This is by far the largest of all
Central American Heliconias, and unlike all of them in its very
dense and thick, heavy, pendent inflorescence. The plants are equal
in size to the tallest banana plants, and resemble them somewhat
when seen from a distance. In some parts of the Panama low-
lands the plants form such dense stands that it is impossible to force
one's way between them without use of a machete. In Guatemala
this species is conspicuous in some localities along the main rail-
road line through the banana country. The very appropriate
English name of "beefsteak Heliconia" has been suggested for
H. Mariae, since the inflorescences by their coloring do suggest a
large chunk of raw beefsteak.

Heliconia psittacorum L. f. Suppl. PI. 158. 1781. H. Ursula
L. f. loc. cit. H. aurantiaca Ghiesbr. ex Lem. 111. Hort. 9: pi. 332.
1862. H. crassa Griggs, Bull. Torrey Club 30: 646. 1903 (type
collected between Sepacuite" and Secanquim, Alta Verapaz, R. F.
Griggs 356 and 376). Bihai crassa Griggs, Bull. Torrey Club 31:
445. 1904.

Wet forest of the Atlantic lowlands, 480 meters or lower; Pete"n;
Alta Verapaz; Izabal. Tabasco; British Honduras, along the Atlantic
coast to Panama; South America.

Plants glabrous, usually about a meter tall, rarely as much as 2 meters;
leaves often numerous, the blades divaricate from the stem, narrowly oblong,
about 30 cm. long and 9 cm. wide, or often narrower or longer, abruptly acuminate,
rounded or subcordate at the often oblique base, thin, bright green; inflorescence
erect, sessile or pedunculate; bracts about 6, orange-yellow, the lowest bract
often ending in a small green blade, the others close together, 3-6 cm. long, about
1 cm. high at the base, attenuate; flowers about 15 in each bract, yellow; pedicels
1 cm. long; fruit 5 mm. in diameter or larger.

Because of its small size and rather pale coloring, this plant is
much less conspicuous than other species.

Heliconia rostrata Ruiz & Pavon, Fl. Peruv. 3: 71. pi. 305.
1803. H. pendula Wawra, Oesterr. Bot. Zeitschr. 13: 8. 1863.
Guinea de montana (Quezaltenango) ; Platanillo.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 185

Wet forest or thickets, 600-1,400 meters; Alta Verapaz; Escuintla;
Quezaltenango. British Honduras, along the Atlantic coast to
Panama; southward to Peru and Brazil.

Plants coarse and stout, usually 2-3 meters tall; leaves long-petiolate, glabrous,
oblong, 1-1.5 meters long, mostly 25-35 cm. wide, abruptly short-acuminate, green
or glaucescent beneath; inflorescence pendent, the rachis terete, rather slender,
very flexuous, densely brown- villous with rather short hairs; bracts cardinal-red
or dull red, lance-linear, widely spaced, divaricate or even somewhat reflexed,
long-attenuate, glabrous or nearly so, the middle ones about 15 cm. long and
2 cm. high at the base; flowers pedicellate, the stout pedicels villous, the perianth
pale yellow or greenish yellow.

Heliconia Schiedeana Klotzsch, Linnaea 20: 463. 1847. H.
tortuosa Griggs, Bull. Torrey Club 30: 650. pi. 29, f. 1. 1903 (type
from Sepacuite", Alta Verapaz, R. F. Griggs 17). Bihai tortuosa
Griggs, Bull. Torrey Club 31: 445. 1904. Platanillo; Xackel (Alta
Verapaz, Quecchi).

Moist or wet forest or thickets, 500-1,500 meters; Alta Verapaz;
Chiquimula; Quezaltenango; San Marcos. Southern Mexico.

Plants stout, 1.5-2.5 meters tall; leaves long-petiolate, often much elongate,
as much as 50 cm. wide but mostly narrower, glabrous, paler green beneath but
not glaucous; inflorescence pedunculate, erect, the rachis flexuous, densely pubes-
cent, the bracts widely spaced, dull red, very narrow and elongate, divaricate or
somewhat curved upward, sparsely pubescent or almost glabrous; flowers pale,
dull yellow, the ovary green; pedicels villous-pilose.

Heliconia spissa Griggs, Bull. Torrey Club 30: 652. 1903. Bihai
spissa Griggs, Bull. Torrey Club 31: 445. 1904.

Pete"n; Alta Verapaz (type collected between Cahabon and
Senahu, on steep river bank, Griggs 359) ; endemic.

Plants about 2 meters tall; leaf blades about 75 cm. long and 20 cm. wide,
oblong-oval, acute, rounded at the base, green, glabrous except beneath on the
costa, there sparsely brown-hirsute; inflorescence erect, subsessile, densely covered
with soft brown hairs except on the edges of the bracts, the rachis almost straight;
bracts widely separated, oblong-lanceolate, acute or obtuse, the lowest 17 cm. long
and 1.5 cm. wide, the middle bracts almost linear, bright yellow shading to orange;
flowers 4 cm. long, yellow, densely villous, the pedicels 1 cm. long, villous; fruit
villous.

Heliconia subulata Ruiz & Pavon, Fl. Peruv. 3: 70. pi. SOSb.
1802. H. acuminata L. Rich. Nov. Act. Acad. Nat. Cur. 15: Suppl.
26. pis. 11, 12. 1831. H. choconiana Wats. Proc. Amer. Acad. 23:
284. 1888 (type from Rio Cocon, Izabal, S. Watson).

186 FIELDIANA: BOTANY, VOLUME 24

Wet forest of the North Coast, near sea level; Izabal. British
Honduras, along the Atlantic coast to Panama; southward to
Bolivia and Brazil.

Plants glabrous, usually about a meter high, sometimes taller and as much
as 2 meters; leaves oblong, mostly 10 cm. wide or narrower and 30-40 cm. long,
bright green; inflorescence erect, on a long or short stalk; bracts deep red, few,
rather close together, the middle ones 7-9 cm. long, at the base scarcely more
than 1 cm. high, narrow, long-attenuate, the lowest bracts elongate and sometimes
bearing a small green blade at its apex ; ovary yellow, tinged at the top with grass-
green; perianth rich yellow.

MUSA L.

Reference: E. E. Cheesman, Kew Bull. 1948, no. 2: 145-153. 1948.

Large herbs with thick perennial roots, the stems often tall and thick but
consisting principally of tightly rolled leaf sheaths; leaves petiolate, arranged
spirally, long-sheathing, the blades usually oblong, very large, penninerved; in-
florescence terminal, rising among the convolute sheaths, consisting of a compound
spike; bracts large, at first completely enclosing the inflorescences; lower flowers
pistillate, the middle ones sometimes perfect or, like the upper ones, staminate;
4 of the sepals unequal, united to form a shallowly 5-lobate tube, this slit dorsally,
the fifth (dorsal) sepal free, entire or 3-lobate; perfect stamens 5, the filaments
thick-filiform, the anthers linear, introrsely dehiscent; ovary 3-celled, the cells
many-ovulate; ovules anatropous, attached horizontally; style filiform or sub-
angulate from a thickened base, the stigma subcapitate, 6-lobate; fruit baccate,
fleshy, containing few or many seeds, in cultivated plants usually seedless; seeds
subglobose or angulate, imbedded in pulp, the testa osseous.

About forty species, natives of the Old World tropics. Two of
them are cultivated on a large scale in the American tropics. M.
textilis Ne'e is exploited extensively in the East Indies for its fiber,
Manila hemp of commerce, noted for its strength and fineness.
Some of the other species are planted occasionally in Guatemala
and other parts of Central America for ornament or as curiosities.
The cultivated bananas and plantains have seedless fruits, i.e., the
seeds are abortive, but some of the less usual forms cultivated have
fruits filled with hard shot-like seeds.

Musa paradisiaca L. Sp. PI. 1043. 1753. Platano, plantain.

The plantain and banana are considered by some authors as
varieties of a single species, but commercially and economically they
are so different that it is at least convenient to consider them as
distinct species. While the banana is quite as well known in many
temperate regions as it is in the tropics, the plantain, one of the
important food plants of tropical America, is practically unknown
in North America and Europe. Plantains can be purchased at times

I STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 187

in some cities of the United States, but there is little demand for
them since few people know how to utilize them, and they are expen-
sive, a fact that would seem fantastic to people familiar with the
tropics. It is unfortunate that plantains are not available in the
United States, because they probably could become a popular
vegetable, being easy of preparation for the table, and of a flavor
that most people would consider agreeable. The fruits ordinarily are
larger than bananas, and they are green when cut, usually turning
black as they ripen, and never of a pure yellow color. Both green
and ripe plantains are prepared for the table. The most usual
method of preparing them is perhaps by boiling, when they are
not particularly appetizing, but have a slightly tart flavor. If
boiled plantains are eaten with honey, as they are often served in
Guatemala, they are much more palatable. Various desserts are
made by boiling or baking plantains with sugar, chocolate, or
various fruits. By the poor people plantains are often baked or
toasted upon a gridiron. Fried plantains are probably the most
palatable of all, and they constitute a most agreeable vegetable,
liked by most foreigners. Plantains are grown in most of the depart-
ments of Guatemala, but almost wholly at low or less than middle
elevations, principally on the plains or in the foothills. They are
much less resistant to cold than bananas. According to data supplied
by the Department of Agriculture of Guatemala, there were produced
in 1938-39 in the country 2,395,912 racimos or stems (bunches)
of plantains. About half of these came from the departments of
Santa Rosa and Escuintla, and more than 375,000 stems from San
Marcos. Other departments producing more than 100,000 stems
were Suchitepequez, Quezaltenango, and Izabal, while for Jalapa
only 576 stems were reported, and only 57,000 for Alta Verapaz.
Several varieties are recognized in Guatemala, but to the inex-
perienced eye they seem to differ little except in size.

Musa sapientum L. Sp. PI. ed. 2. 1477. 1763. Guinea; Banano,
banana. Tul (Quecchi); Ts'ahlec, Tulul (Poconchi); Haaz, Sachaaz,
Boxhaaz (Maya). M. paradisiaca var. sapientum Kuntze, Rev. Gen.
2: 692. 1891.

As in all or most other parts of Central America, the banana
in its various varieties is one of the most important of all cultivated
plants, not only as a source of food for the inhabitants but as an
article of export. For export the only variety of any importance is
the well known Gros Michel, the common banana of United States
markets, known in Guatemala generally as "banano," or sometimes

188 FIELDIANA: BOTANY, VOLUME 24

as "minimo" or "guineo de seda." It is grown to some extent in all
departments of Guatemala except Totonicapan, from sea level up
to at least 2,400 meters, but very sparingly and hardly commercially
at the higher elevations. The great majority of the fruit is produced
at or near sea level, but the plantations thrive up to probably 1,000
meters or more, and for local use the plants are cultivated consider-
ably higher. Bananas and coffee together account for about 90
per cent of the exports of Guatemala. The greater part of the
exported bananas is produced by the United Fruit Company and
its affiliates in two regions of the North and Pacific coasts, the first
with headquarters at Bananera in Izabal, and the second at Tiquisate,
in Suchitepequez, but on the border of Escuintla. Bananas produced
by private growers are shipped by the same company, but in recent
years, because of changes in methods of cultivation resulting
principally from attacks of the Panama and Sigatoka diseases, culti-
vation of bananas suitable for export has become a business requiring
such heavy investment that few private growers can afford to engage
in it. The export banana industry of Guatemala began to develop
about 1900 in the lower Motagua Valley and around Lake Izabal.
This region is still a great producer of bananas, but in very recent
years the Pacific coast region about Tiquisate has attained even
greater production. Only a few years ago large quantities of
fruit were produced by independent planters for export in the plains
and foothills of almost the whole Pacific coast, but the ravages of
disease, resulting in fruit that was perfectly edible but not export-
able have greatly circumscribed the area in which fruit is produced
for export.

For 1938-39 the production of bananas (i.e., Gros Michel) for
Guatemala is reported as 14,353,572 stems or bunches (racimos).
The leading departments in production were Escuintla, 4,830,896
stems; Izabal, 4,460,276; Suchitepequez, 2,887,640; and Retalhuleu,
1,058,244. Most of these were exported. The other departments
produced only limited amounts for local consumption.

The common banana is the one most esteemed everywhere in
Central America for eating raw. It is also cooked in various ways,
and is much in demand in all parts of the country. The fruit is
naturally much cheaper than in the United States, and it is possible
to buy a dozen of the large, handsome, delicious bananas of San
Felipe for a cent. Large quantities of the fruit are carried on the
backs of cargadores from the lowlands up into Quezaltenango and
other towns of the highlands for sale, as return loads by men who

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 189

have taken down to the tierra caliente loads of earthenware, woollen
goods, and other manufactures. Since only a few cents less than
a dime is paid for carrying such a load, transportation adds little
to the price of the fruit. While the fruit thus transported is
ripe, bananas, of course, always are cut from the plant while still
very green. In fact, those cut for local use are just as green as those
shipped to the United States. Only by infrequent accident do
bananas ripen on the plant anywhere in Guatemala or elsewhere
in Central America. Birds and mammals would consume them long
before they were fully ripe, if this practice were followed. Alta
Verapaz, or at least Coban, is quite as poorly supplied with bananas
as with most other desirable fruits, and the bananas sold in the
Coban market are usually of poor appearance and even worse
flavor. The Quecchi name for the common banana is reported as
"Mines."

Plants of the various varieties of bananas are much used as
coffee shade in some regions. The growing of bananas in coffee
plantations of the Pacific bocacosta promised to be a profitable
industry until the plants became diseased. In the spring of 1941
many of the bananas planted for export in the bocacosta were being
cut from the cafetaks, on a large scale. There have been noticed,
especially in the Coban region, banana plants with teratological
inflorescences that give an unusual appearance to the plants. While
not investigated carefully, it appeared that the whole inflorescence
had been transformed into a mass of foliaceous, much imbricated
bracts. In the small fincas such plants are left until the foliage
withers, when they have a fantastic appearance.

Besides the production data for "banano," the Guatemalan
Department of Agriculture reports for 1938-39 the production of
"guineo" as 3,405,896 stems. The term guinea (probably from
Guinea of Africa) in Central America embraces all forms of bananas,
but these figures evidently are intended to cover varieties other
than Gros Michel, and of these additional forms several are cultivated
upon a large scale, for food. There is, so far as we know, no good
account of the cultivated varieties of bananas, at least of those
grown in Central America, and we do not know what Latin names
should be applied to them. The local names vary from one part
of Guatemala to another. The fruit characters are not well preserved
in herbarium specimens, and we have not taken detailed notes of
the varieties found in different parts of the country. Some of the
principal varieties, however, are enumerated below. We have had

190 FIELDIANA: BOTANY, VOLUME 24

references also to some other forms, such as the "cantiado," "ha-
banero," "datil," and "criollo," whose characters are unknown to us.

Musa sapientum var. Champa Baker. Guinea manzano;
Guinea de oro; Guinea manzanita.

Plants smaller than in the common banana; fruits much smaller,
rather pale or bright yellow; pulp rather deep yellow, very soft
and fine-grained, fairly sweet. This banana is much esteemed by
some persons, and most people will agree that it is the best banana
to eat raw, after the common one. It is much used as food for
cage birds, and is usually plentiful in the markets, but it is a variety
of relatively little economic importance. The guinea de oro and
guinea manzano are sometimes considered as distinct varieties, the
former having a very yellow skin and deep yellow, sweet flesh, the
guineo manzano being slightly larger and having a pale yellow skin
like the common banana. The two are not easy to recognize when
so separated unless specimens of both are at hand for comparison.
This type of banana seems to flourish at high elevations and is
planted, for instance, about Huehuetenango, where the leaves
sometimes are severely frosted.

Musa sapientum var. rubra (Firm.) Baker. Guineo morado;
banano morado, red banana.

Easily recognized by its reddish petioles and the reddish or
purplish tinting of the leaves. The fruits are large, the skin con-
spicuously colored dull red or purplish. The flesh is rather coarse
but of fair quality for eating. The ripe fruits are eaten raw, the
green ones cooked in various ways. This form is planted abundantly
in all the lowland regions, and is especially abundant about San
Pedro Carcha and beyond, where it is planted to shade coffee in
many of the fincas. The red banana is exported in small quantities
to the United States, where it commands a higher price than the
common banana, presumably because of its scarcity. In Central
America it is considered inferior to the common banana, as it must
be if the flavor and texture of the two varieties are compared.

Guineo bianco; Sakitul (Quecchi); Guineo de pajaro. A variety
not very common except about Coban, where it is highly esteemed.
The fruit is of a very pale yellow, and a little larger and thicker
than the "Guineo de oro." The pulp is pale orange and rather tart.

Guineo de coche (Huehuetenango, Coban; "pig banana," Guineo
de San Antonio, central region). This is the commonest banana of

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 191

the Coban market, and abundant in that of Huehuetenango. It
is said to be the variety most planted in all the Huehuetenango region.
The fruit is short and thick, of a bright but pale green when ripe.
It is eaten raw, but in quality is decidedly inferior.

Guinea majonche; Majonche; Majoncho. This is one of the most
important fruits of the Guatemalan lowlands, especially of the
Oriente, and equally or more important in Salvador and some other
parts of Central America, being the banana most often cooked for
the table. After maize, rice, and frijoles negros it is possibly the
most important article of food among the poorer people. The fruit
is relatively thicker than the common banana, pale buff or pinkish
buff, the pulp coarse, fibrous, and of poor flavor. When cooked,
however, the fruit is fairly palatable, with at least as much flavor
as ordinary potatoes, and it serves at least to diversify the sadly
monotonous diet of the working classes of Central America.

Ravenala madagascariensis Sonn. Palma de viajero (fide
Aguilar). This, the famous traveler's tree, native of Madagascar,
is planted occasionally for ornament in gardens of Guatemala City,
in the bocacosta of Quezaltenango and San Marcos, and doubtless
in other places. It is easily recognized by the fan-like arrangement
of the huge banana-like long-stalked leaves. The larger plant has
a well-developed trunk. The genus Ravenala consists of two species,
one of which, strangely enough, is native in the Guianas.

Strelitzia reginae Banks. Native of the Cape region of Africa,
occasionally cultivated for ornament in Guatemala City and probably
elsewhere, but rare. In general appearance it somewhat resembles
the smaller Heliconias, and has a similar, brightly colored inflores-
cence of but few narrow spreading bracts.

ZINGIBERACEAE. Ginger Family
Reference: K. Schumann, Pflanzenreich IV. 46. 1904.

Perennial herbs, usually tall and often very large and stout; true stems usually
very short, but stemlike organs often formed by the tightly rolled leaf sheaths;
leaves commonly long-sheathing, the blades sessile or petiolate, broad or narrow,
frequently with a large ligule; flowers bracteate, often large and showy, the bracts
subtending 1 or several flowers, the inflorescence racemose, spicate, capitate, or
paniculate, terminating a leafy stem or arising from the base of the plant on a
short, bracteate peduncle; flowers irregular, perfect; calyx tubular, usually trilo-

192 FIELDIANA: BOTANY, VOLUME 24

bate, commonly cleft on one side, often marcescent after anthesis; corolla funnel-
form, trilobate, the posterior lobe usually larger than the others and enclosing
them in bud; stamens 6, but only 1 fertile, the others consisting of petal-like or
dentiform staminodia; anther introrse, 2-celled, dehiscent by longitudinal slits;
ovary normally 3-celled; fruit capsular, loculicidally trivalvate or dehiscent by
loculicidal slits, the exocarp often fleshy; seeds few or numerous, variable in form,
arillate.

The family is a tropical one, abundantly represented in both
hemispheres, but with relatively few genera and species in Central
America. Besides the genera reported here, one other, Dimerocostus,
is represented in Central America, in Panama and Costa Rica.

Leaves arranged spirally. Large and tall, coarse plants with elongate, leafy
stems; flowers arranged in a terminal and conelike, headlike or spikelike

inflorescence Costus.

Leaves 2-ranked.

Lip of the corolla trilobate; cultivated plant with the odor and taste of ginger.

Zingiber.

Lip of the corolla not trilobate; plants not with the odor and taste of ginger.
Flowers in dense spikes or racemes terminating leafy stems; introduced plants.

Flowers racemose, white, pink, and yellow Alpinia.

Flowers in dense spikes, the perianth pure white Hedychium.

Flowers in panicles arising on short leafless scapes from the base of the tall
leafy sterile stems, or in one species the inflorescence cone-like and
terminating a leafy stem Renealmia.

ALPINIA Roxburgh

Robust herbs of medium size with leafy stems, usually growing in dense clumps;
leaves large, distichous, petiolate or sessile; inflorescence racemose or subspicate,
sometimes paniculate, terminating the stem, the bracts large and imbricate, the
flowers large and showy; calyx turbinate to tubular, with 3 teeth or small lobes,
often cleft above on one side; corolla tube equaling or longer than the calyx, the
lobes commonly unequal, the posterior one much larger; lip usually broad and
much larger than the corolla lobes, the lateral staminodia small and inconspicuous;
ovules numerous; fruit capsular, dry, the seeds angulate.

The species are all natives of the Old World, chiefly of the East
Indies.

.Alpinia speciosa (Wendl.) Schum. Fl. Kaiser- Wilhelmsl. 29.
1887. Zerumbet speciosum Wendl. Sert. Hann. pi. 19. 1798. A.
nutans Roscoe in Sm. Exot. Bot. 2: pi. 106. 1805. Languas speciosa
Merrill, Enum. Philip. PI. 234. 1922. Perlas; Perlas del Oriente;
Collar de la reina.

Planted commonly in gardens for ornament, principally in the
lowlands but also at middle elevations, doubtless naturalized in
some places, as elsewhere in Central America. Native of China and
Japan.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 193

Plants usually about a meter high and forming dense clumps; leaves short-
petiolate, acuminate, narrowed at the base, 70 cm. long and 10 cm. wide or smaller,
glabrous except on the puberulent margin, the ligule 1 cm. long or less; panicles
up to 30 cm. long, the bracts 2-3-flowered, broadly elliptic, whitish, about 3 cm.
long; calyx almost 2 cm. long, white; corolla white, the tube 1 cm. long, the lobes
broadly elliptic, obtuse, 3 cm. long, red at the apex; lip broadly ovate, 4 cm. long,
subtrilobate, spotted and streaked with red and yellow; capsule globose, 2 cm. in
diameter.

Known elsewhere in Central America as "lirio de Colon,"
"grano de oro," "lagrimas de San Juan," "lagrimas de Nazareno,"
etc. The plant is much planted about the humbler dwellings of
Central America, and often persists where they have disappeared.
The large and showy inflorescences are brought to the markets,
especially for use as decorations during religious and other fiestas.

COSTUS L.

Reference: Rowlee, The genus Costus in Central America, Bull.
Torrey Club 49: 283. 1922.

Tall or low herbs, often very large, rarely acaulescent, the stout stems con-
sisting of the tightly sheathing leaf bases; leaves spirally arranged, sessile or short-
petiolate, glabrous or pilose; flowers yellow, red, or white, disposed in a dense,
cone-like spike, this terminal or borne on a short, leafless scape at the base of the
leafy stem, the bracts broad, 1-2-flowered, coriaceous, closely imbricate, sometimes
with leafy, terminal appendages; calyx tubular, coriaceous or membranaceous,
more or less trilobate at the apex; corolla tube slender, equaling or longer than the
calyx, the lobes imbricate, subequal; outer stamens or staminodia none, the lip
large, obovate, more or less bilobate at the apex; stamen with a broad, petaloid
filament, the anther 2-celled, usually shorter than the free portion of the filament;
ovary with numerous ovules; capsule trigonous, trivalvate, usually dehiscent by
loculicidal slits; seeds angulate.

About fifteen species are known from Central America. The
genus is a large one, of approximately one hundred species, widely
dispersed in the tropics of the various continents. The plants often
constitute a conspicuous element of the undergrowth of the wet
lowland forests of Guatemala, being prominent because of their
unusual appearance and showy inflorescences, that persist for a
long time. In Central America generally the succulent stems are
variously used in domestic medicine, particularly for treating
gonorrhea and related diseases, for which the common people
believe them to be highly effective.

Inflorescences borne on short leafless scapes arising at the base of the leafy stem.

C. sepacuitensis.
Inflorescences terminating the leafy stems.

Bracts of the spikes bearing spreading leafy appendages.

194 FIELDIANA: BOTANY, VOLUME 24

Plants densely hirsute with long spreading hairs C. villosissimus.

Plants with sparse pubescence of short appressed hairs C. Bakeri.

Bracts of the spikes not appendaged, closely appressed, at least during anthesis.

Spikes ovoid C. pulverulentus.

Spikes fusiform, cylindric, or globose.

Spikes fusiform, the bracts closely appressed in age C. sanguineus.

Spikes cylindric or globose, the bracts in age lax and often spreading.
Spikes about 7 cm. thick; plants tall, usually much more than 2 meters

high C. laevis.

Spikes about 4 cm. thick; plants smaller, commonly 1-2 meters high.

Spikes red, cylindric C. ruber.

Spikes green, often globose C. congestus.

Costus Bakeri Schum. Pflanzenreich IV. 46: 387. 1904. Cana
de Cristo; Cana de la Virgen.

Wet thickets, especially in ravines along small streams, mostly
at 400-1,400 meters; Santa Rosa; Escuintla; Guatemala (type
from Barranco Eminencia, J. D. Smith 2802); Suchitepequez;
Retalhuleu; Quezaltenango; San Marcos. Chiapas; reported from
Costa Rica.

Plants stout, 1.5-2.5 meters high; leaves short-petiolate or sessile, lanceolate,
oblong, or oblanceolate, 30 cm. long or less, caudate-acuminate, narrowed at the
base, usually glabrous above, sparsely sericeous beneath, the ligule very short,
villous; spikes subglobose, about 5 cm. broad, the bracts dull red, rarely green or
yellow, the foliaceous appendages lanceolate, spreading, acuminate, the inner
sterile bracts erect and comose, the bracts all rather densely appressed-pilose;
calyx 1 cm. long, puberulent; corolla tube 1 cm. long, the lobes oblong, about twice
as long.

The plant has been reported from Guatemala as C. comosus
Roscoe. It is abundant and conspicuous in many parts of the
bocacosta, where it sometimes forms large colonies on open hillsides,
especially in the Department of Quezaltenango. While the normal
color of the bracts is dull red, scattered plants have pale yellow or
even green bracts.

Costus congestus Rowlee, Bull. Torrey Club 49: 291. pi. 14.
1922. Cana de Cristo.

Moist or wet thickets, 1,350 meters or lower; Alta Verapaz;
Jutiapa; Escuintla (type from Escuintla, J. D. Smith 2036); Solola;
Suchitepequez; Quezaltenango; Huehuetenango. Reported from
Oaxaca.

Plants low, usually about a meter high, rarely somewhat taller; leaves sessile,
oblanceolate, 24 cm. long and 6 cm. wide or smaller, acuminate, narrowed to the
base, glabrous; spikes green, subglobose or ellipsoid, about 4 cm. long; bracts

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 195

in flower closely imbricate, obtuse, glabrous; ovary glabrous; petals large, pale
yellow, hyaline, obtuse; lip about 4 cm. long.

This is very close to C. ruber, from which it may not be specifically
distinct.

Costus laevis Ruiz & Pavon, Fl. Peruv. 1: 3. 1798. C. splendens
Bonn. Smith & Turckh. Bot. Gaz. 33: 260. 1902 (type from Cubil-
giiitz, Alta Verapaz, Turckheim 8015). Cana de Cristo.

Wet forest or thickets, 1,300 meters or lower; Alta Verapaz;
Izabal; Santa Rosa; Escuintla; Sacatepe"quez; Quezaltenango; San
Marcos. Probably in Chiapas; Costa Rica; Panama; southward to
Peru and Brazil.

A coarse stout plant, commonly 2-3 meters high, often forming large colonies;
leaves mostly oblanceolate, on petioles as much as 2 cm. long, caudate-acuminate,
narrowed to the base, glabrous or nearly so, mostly about 25 cm. long and 6-7 cm.
wide, or larger, the ligule 5 mm. long; spikes cylindric or oblong, as much as 22 cm.
long and 7 cm. broad, the bracts red and green, broadly elliptic, very obtuse,
coriaceous, somewhat sericeous or glabrate; ovary sericeous-tomentose; calyx 1 cm.
long, glabrous; capsule obpyramidal, 2.5 cm. long, 1.5 cm. broad, the seeds 3 mm.
long.

Costus pulverulentus Presl, Rel. Haenk. 1: 41. 1830.

At 390 meters; Suchitepequez (Rio Sis, J. D. Smith 2800).
Originally described from southern Mexico.

Leaves sessile, oblong-lanceolate, abruptly acuminate, attenuate to the base,
13-16 cm. long and 4.5-5.5 cm. wide, glabrous above, squamulose-puberulent
beneath, the ligule short, truncate; spikes terminal, sessile, 6.5 cm. long, sub-
tended at the base by 3 or more leaves; bracts closely imbricate, ovate-rounded,
retuse or emarginate or obtuse and mucronate, short-scabrous, 2 cm. long and wide.

A little known species, its systematic position doubtful. The
Guatemalan collection was referred by Baker to C. Malortieanus
Wendl., a Costa Rican species.

Costus ruber Griseb. Cat. PI. Cub. 256. 1866. Cana de Cristo;
Cana agria; Cu, Pacuite (Quecchi); Cooltze (Pete"n, Maya, fide
Lundell).

Wet forest or thickets, 1,400 meters or lower, most plentiful at
low elevations; Pete"n; Alta Verapaz; Izabal; Zacapa; Escuintla;
Suchitepequez; Retalhuleu. Southern Mexico; British Honduras to
Panama; Colombia; West Indies.

Plants stout, commonly 1-1.5 meters high; leaves subsessile, elliptic to ob-
lanceolate, mostly 10-20 cm. long and 5-7 cm. wide, abruptly short-acuminate,
tapering to the base, glabrous or nearly so, the ligule 3-5 mm. long; spikes ellipsoid,

196

FIELDIANA: BOTANY, VOLUME 24

about 5-7 cm. long and 4 cm. thick, the bracts coriaceous, broadly ovate, acute
or usually obtuse, dull red or orange-red, 1-flowered; ovary puberulent; calyx
shallowly trilobate, glabrous; corolla yellow or orange-red, the tube 1 cm. long,
the lobes oblong, acute, the broader posterior one 2 cm. long; capsule subtrigonous,
puberulent, the seeds numerous.

FIG. 35. Costus villosissimus. Habit of uppermost portion of plant; X

This is a very common plant in the wet forests of the North
Coast region, as it is also in many other parts of Central America.
It has been reported frequently from Guatemala and other parts
of Central America as C. spicatus Jacq.

Costus sanguineus Bonn. Smith, Bot. Gaz. 31: 122. 1901.

Wet thickets and mixed forest, 450 meters or lower; Alta Verapaz
(type from Cubilgiiitz, Turckheim 7686); Izabal; Huehuetenango.
Honduras; Nicaragua; Costa Rica.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 197

Plants stout, 1-2 meters high; leaves subsessile, oblanceolate, about 20 cm.
long and 6 cm. wide, short-acuminate, narrowed to the base, pilose-sericeous,
especially beneath, the short ligule villous; spikes fusiform, or elongate-cylindric,
up to 10 cm. long and 3-4 cm. thick, or often much more elongate; bracts dull
red, orange, or green, thick-coriaceous, rather closely imbricate and not at all
spreading in age, rounded or very obtuse at the apex, often tomentose on the
margins; ovary sericeous-tomentose; calyx campanulate, 8-9 mm. long, pilose;
corolla purplish red, 5.5-6 cm. long, the lobes lanceolate, acute, equaling the tube;
lip truncate, 5-lobate.

Called "cana de Cristo" in Honduras. A showy plant because
of the unusually long and often brightly colored spikes.

Costus sepacuitensis Rowlee, Bull. Torrey Club 49: 286. pi. 13.
1922.

Type from Finca Sepacuite", Alta Verapaz, 0. F. Cook & R. F.
Griggs 596. Known only from the original collection.

Leaves sessile, narrowly elliptic-oblong, about 30 cm. long and 9 cm. wide,
acute or short-acuminate, soft-pilose on both surfaces, ciliate, the ligule 1.7 cm.
long; flowering scapes arising at the base of the plant, 40 cm. long or more, leafless
but bearing numerous puberulent sheaths; spikes somewhat elongate, the bracts
broadly ovate, 4 cm. long and 3-4 cm. wide, obtuse or rounded, glabrous; fruit
pubescent, obovoid, 15 mm. long, 8 mm. wide; calyx deeply lobate, the unequal
lobes about 13 mm. long, ovate, acute.

Costus villosissimus Jacq. Fragm. Bot. 55. pi. 80. 1800-1809.

Wet mixed lowland forest, 350 meters or lower; Alta Verapaz;
Izabal. Southern Mexico; British Honduras along the Atlantic
coast to Panama; Lesser Antilles; South America.

Plants stout, 1-2.5 meters high, the sheaths rufous-hirsute; leaves sessile or
nearly so, lanceolate, acuminate, acute at the base, about 30 cm. long and 10 cm.
wide, rufous-hirsute or pilose on both surfaces, more densely so beneath; spikes
ovoid, dark red, the outer bracts rufous-villous, leafy-appendaged, as much as
9 cm. long and 4 cm. wide, acute; calyx 1.5 cm. long, the lobes only 4 mm. long;
corolla tube 1.5-2 cm. long, the lobes oblong, acute, 6 cm. long, yellow; lip as
much as 9 cm. long. (Fig. 35.)

Called "choschogo" in Veracruz.

It is probable that Curcuma longa L., turmeric, native of India,
is cultivated occasionally in Guatemala, as it is in some other parts
of Central America. In general appearance it is much like ginger.
The thick roots are the source of turmeric, employed as a dye and
condiment. In Guatemala the plant is called "camotillo" and
"tintura de yodo."

198 FIELDIANA: BOTANY, VOLUME 24

Elettaria cardamomum Maton, cardamom, native of the East
Indies, has been planted on a rather large scale in the lowlands of
Alta Verapaz, about Cubilgiiitz and elsewhere, and it is reported
as in cultivation in Escuintla. Its aromatic seeds have long been
an article of commerce, for use as a spice. The plant is a coarse,
tall perennial with large, aromatic, lanceolate leaves. The recumbent
flowering branches (concinni) arise from the fleshy base of the plant
and are distinct from the upright leafy stems. The flowers of the
concinnus consist of small lanceolate greenish sepals and a prominent
enlarged white labellum, veined reddish-purple along the middle. It
is stated that several "varieties" are grown in Alta Verapaz. The
plants there are said to be much subject to disease, but quantities
of the seeds have been harvested for export.

HEDYCHIUM Koenig

Herbs of medium size, usually cespitose or forming dense colonies, with leafy
stems, the leaves distichous, sessile or petiolate; flowers large and showy, arranged
in a dense, terminal spike, subtended by broad, imbricate bracts, the flowers
solitary or several from each bract; calyx tubular, more or less trilobate or entire,
usually cleft on one side; corolla tube long and slender, much exceeding the calyx,
the lobes linear, imbricate; outer 2 staminodia petaloid and resembling corolla
lobes, the lip large or small, broad, usually unguiculate; filament commonly
elongate, the anther linear; capsule loculicidal.

The species are all or chiefly native in the East Indies.

Hedychium coronarium Koenig in Retz. Obs. Bot. 3: 73. 1783.
Perlas de Oriente; Tzi (Quecchi).

Often planted in gardens for ornament; abundantly naturalized
in marshes and wet thickets along the Pacific bocacosta, in Alta
Verapaz, and doubtless in other places; native, according to Schu-
mann, of the Himalayan region, but now naturalized widely in
tropical America and other regions; Alta Verapaz; Izabal; Escuintla;
Suchitepequez; Retalhuleu; San Marcos.

A coarse herb about a meter high with leafy stems; leaves sessile, lanceolate
or oblong-lanceolate, 10-50 cm. long, 3-10 cm. wide, acuminate, acute at the base,
glabrous above, sparsely pilose beneath, the ligule 2-3 cm. long; spikes ellipsoid,
dense, with the flowers 10-20 cm. long and 4-8 cm. broad, the bracts green, 4-5.5
cm. long, ovate, obtuse, subcoriaceous, 2-3-flowered; calyx 4 cm. long; corolla
tube very slender, 8 cm. long, the lobes 3-3.5 cm. long.

Called "mariposa" in Salvador. A handsome and showy plant
because of the large, pure white flowers which, however, are delicate
and soon wither. In many places along the Pacific slope at low and

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 199

middle elevations the plant forms vast colonies, sometimes of several
acres, through the marshes. It grows in association with Calathea
and other native plants in areas far from dwellings where no other
imported plants are to be seen. With the marsh plants it makes a
typical association of perfectly natural appearance. Under such
conditions it is difficult to believe that it is not a native plant, and
it is not easy to judge how it has been dispersed so widely, if intro-
duced. There is no good reason, however, for supposing that it is
not an escape from cultivation.

Kaempferia rotunda L., native of the East Indies, is grown rarely
for ornament in Guatemalan gardens. The plant is acaulescent,
and usually leafless at time of flowering. The few large orchid-like
flowers arise from the rootstock and appear just above the ground,
one at a time. The petals are white and lavender, the sepals whitish
and reflexed. The leaves are purple beneath.

RENEALMIA L. f.

More or less aromatic, often very tall herbs, the stems leafy; leaves petiolate
or sessile, lanceolate or oblong, glabrous, long-sheathing, the ligule usually short;
inflorescence borne on a short, leafless scape arising at or near the base of the
leafy stem, or conelike and terminating a leafy stem; bracts relatively small,
withering after anthesis, the flowers in loose or dense racemes or panicles, the
peduncles and pedicels commonly dark red, the flowers white, yellow, or red,
small; calyx tubular or turbinate, closed in bud, later tridentate or trilobate,
often cleft on one side; corolla tube usually longer than the calyx, the lobes equaling
or longer than the tube, obtuse; lip short, more or less trilobate, the lateral
staminodia short and dentiform; anther sessile or on a very short filament; ovary
commonly glabrous, 3-celled, the ovules numerous; capsule 3-celled, crowned by
the persistent calyx, 3-valvate; seeds angulate, enclosed in a large, fleshy, usually
red or orange aril.

A genus of about sixty species, in tropical America and Africa.
Several species besides those listed here are native in southern
Central America.

Inflorescences terminal on leafy stems R. cernua.

Inflorescences borne on leafless scapes arising at the base of the leafy stems.

Plants very large, sometimes 4 meters high; capsule 2-3 cm. long; calyx almost
2 cm. long R. exaltata.

Plants smaller, 2 meters high or less; calyx about 5 mm. long. . . .R. aromalica.

Renealmia aromatica (Aubl.) Griseb. PI. Brit. W. Ind. 601.
1864. Alpinia aromatica Aubl. PI. Guian. 3. 1775. A. occidentalis
Swartz, Prodr. Veg. Ind. Occ. 9. 1788. R. occidentalis Sweet, Hort.

FIG. 36. Renealmia aromatica. A. Habit of plant; X 1 /a. B. Leaf detached
from stem; X J^. C. Inflorescence in position, and base of stem; X H-
D. Ovary in longitudinal section; X 2. E. Ovary in transverse section; X 2.
F. Fruit; X 2.

200

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 201

Brit. ed. 2. 493. 1830. Tzi (Alta Verapaz); Chucho (Alta Verapaz);
Nabay (Pete"n, Maya).

Wet mixed forest, 1,500 meters or lower; Pete"n; Alta Verapaz;
Izabal; Jutiapa; Santa Rosa; Escuintla; Suchitepequez; Solola;
Huehuetenango. Southern Mexico; British Honduras to Panama;
West Indies; northern South America.

Sterile stems sometimes 2 meters high but commonly lower; leaves sessile or
subsessile, lanceolate or oblong, acuminate, acute at the base, 15-50 cm. long,
5-12 cm. wide, glabrous; panicles arising at the base of the plant, 15-30 cm.
long, long-pedunculate, the rachis puberulent, the lower bracts as much as 8 cm.
long, lanceolate, acute, glabrous, usually dark red, pedicels 1 cm. long or less;
calyx 5-6 mm. long, red; ovary pilosulous or glabrate; corolla yellow, the elliptic
lobes equaling the tube; capsule subglobose, red, diameter about 1 cm. (Fig. 36.)

A characteristic species of the wet lowland forests, abundant in
many places. Here probably are to be referred collections reported
by Captain Smith as R. mexicana Klotzsch and R. breviscapa Poepp.
& Endl. The latter is a South American species, unknown in North
America. It is questionable whether R. mexicana, separated by
Schumann from R. aromatica because of its smaller size, is really
different from that species. At least, all the Guatemalan specimens
of this alliance appear to be conspecific, and properly referable to
R. aromatica. The Indians of Alta Verapaz use the pulp of the fruit,
it is said, for flavoring meat soup. The leaves are covered with tallow
and applied to fractures, sprains, and surface inflammation to reduce
them.

Renealmia cernua (Swartz) Macbride, Field Mus. Bot. 11: 14.
1931. Costus cernuus Swartz ex Roem. & Schult. Syst. Veg. 1: 25.
1817. R. strobilifera Poepp. & Endl. Nov. Gen. & Sp. 2: 26. pi. 136.
1838.

Wet mixed forest, at or near sea level; Izabal (Puerto Barrios).
Ranging to Panama and southward to Peru.

Stems very leafy, commonly about a meter high but sometimes 2 meters;
leaves sessile, lanceolate or linear-lanceolate, acuminate, acute at the base, glabrous,
about 30 cm. long and 6 cm. wide or smaller; inflorescence spicate, terminating
the stem, ovoid or ellipsoid, 5-10 cm. long, at first erect, later nutant, the bracts
coriaceous, orange, closely imbricate, striate, lanceolate; flowers solitary, the
ovary glabrous; calyx tubular, 1.5 cm. long; corolla yellow, coriaceous, the tube
shorter than the calyx, the lobes ovate, equaling the tube; lip 1 cm. long, linear-
oblong; capsule ellipsoid, 10-12-seeded.

Very common in lowland forests of Costa Rica and Panama, and
known also from Nicaragua, but strangely enough, no specimens
have been seen from Honduras.

202 FIELD IANA: BOTANY, VOLUME 24

Renealmia exaltata L. f. Suppl. PI. 79. 1781. Huehuechucha;
Jazmin de monte (Huehuetenango).

T-

Wet thickets or dense mixed forest, 1,200 meters or lower;
reported from Alta Verapaz (Coban); Chiquimula; Santa Rosa;
Solola; Huehuetenango; Quezaltenango; Santa Rosa. Southern
Mexico; southward to Panama and the Guianas; West Indies.

Plants very large and coarse, usually 2 meters high or more, often forming
large colonies; leaves sessile or short-petiolate, elongate-lanceolate or oblanceolate,
up to a meter long and 6-12 cm. wide, short-acuminate, long-attenuate at the
base, glabrous; panicles basal, 20-30 cm. long, red, subtomentose, long-peduncu-
late; flowers in small, pedunculate inflorescences, the bracts lanceolate, 8 cm. long
or less, the pedicels about 5 mm. long; calyx almost 2 cm. long, bilobate, sparsely
sericeous; corolla pink, the tube equaling the calyx, the lobes obtuse, 1 cm. long;
capsule ellipsoid, striate, 2-3 cm. long, striate; seeds large, yellowish, lustrous,
3-4 mm. in diameter, surrounded by a white aril.

ZINGIBER Adans. Ginger

Perennial herbs with aromatic rhizomes, the floriferous stems usually distinct
from the leafy ones, or the inflorescence sometimes terminating a leafy stem;
flowers commonly solitary in the bracts, subtended by a spathaceous bractlet,
the bracts dense or lax; calyx tridenticulate, usually cleft on one side; corolla tube
slender, dilated toward the apex, the lobes imbricate, often very unequal, the
posterior one concave and erect, the anterior ones narrower; lateral staminodia
small or large, colored, the lip deflexed, entire, emarginate, or shallowly bilobate;
ovules numerous; capsule subglobose or ellipsoid, the pericarp thin, trivalvate or
irregularly ruptured; seeds large, surrounded by a lacerate aril.

The species are all natives of the Old World, and chiefly of the
East Indies.

Zingiber officinale Roscoe, Trans. Linn. Soc. 7: 348. 1807.
Amomum Zingiber L. Sp. PL 1. 1753. Gengibre; Xenxibre (Quecchi).

Planted frequently in gardens and perhaps naturalized locally.
Probably native of the East Indies, but cultivated in all tropical
regions.

Stems a meter high or more, from a tuberous rhizome; leaves numerous,
sessile, lanceolate or linear-lanceolate, long-acuminate, narrowed at the base, as
much as 20 cm. long and 2 cm. wide; flower spikes long-pedunculate, ellipsoid,
dense, about 5 cm. long and 2-2.5 cm. thick, the bracts ovate, cuspidate, pale
green, 2.5 cm. long; calyx crenate, almost 1 cm. long; corolla yellowish green, the
tube 2 cm. long or less, the lobes subequal, lanceolate, acute; lip oblong-obovate,
purplish, yellow-dotted.

The rhizomes of ginger are widely used as a condiment and drug.
The plant usually is propagated by pieces of the rhizome, and in

[ STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 203

Central America it is said to flower rarely or not at all. Schumann
states that the capsules are unknown. The dried rhizomes are
offered for sale commonly in the markets. It is said that the Quecchi
Indians of Lanquin grow the plants in some quantity and carry the
roots for sale to the capital and other remote markets.

j CANNACEAE. C anna Family

Reference: Fr. Kraenzlin, Cannaceae, Pflanzenreich IV. 47. 1912.

Large erect leafy herbs, the thick stems arising from slender rhizomes or
usually from thick tuberous rhizomes; inflorescences terminal, usually branched,
sometimes simple, the flowers spicate, asymmetric, mostly red or yellow; sepals 3,
imbricate, subequal or one of them smaller, generally herbaceous; petals 3, alter-
nate with the sepals, one of them always smaller than the others, more or less
connate at the base into a tube, 1 or 2 reflexed, or all sometimes reflexed; an-
droecium more or less altered into petal-like staminodia, or the outer staminodia
sometimes absent, usually 2-3, the inner staminodia 2, one (the lip) revolute,
the other petaloid, bearing a dimidiate 1-celled anther on one side, connate at the
base or to the middle with a usually linear, petaloid style; style inserted obliquely
on the ovary, the stigma terminal, obliquely inserted on the style, deeply sulcate;
ovary inferior, subglobose, papillose; capsule subglobose or ellipsoid, 3-celled,
loculicidally dehiscent, membranaceous at maturity; seeds several in each cell,
extremely hard; endosperm white, very hard.

A single genus, with about fifty species, mostly natives of tropical
America, a few in tropical Africa and Asia, some of them naturalized
in the Old World tropics.

CANNAL. Canna

Staminodia besides the lip always 2; flowers yellow.

Leaves glaucous beneath C. discolor.

Leaves green beneath.

Flowers spotted; bracts about 1 cm. long C. lutea.

Flowers not spotted; bracts about 2 cm. long C. sanctaerosae.

Staminodia always 3, variously connate; flowers red or orange.

Flowers about 15 cm. long C. Tuerckheimii.

Flowers mostly 5-7 cm. long.

Plants large and coarse, generally 2-3 meters high, sometimes lower; flowers

orange or orange-red C. edulis.

Plants low and relatively slender, usually about a meter high; flowers bright
or pale red, with no orange tint C. indica.

Canna discolor Lindl. Bot. Reg. pi. 1231. 1829.

Reported as collected in Guatemala, without specific locality,
by Bernoulli; Suchitepequez (Rio Sis, J. D. Smith 2799); also from
Retalhuleu. West Indies; Colombia.

204 FIELDIANA: BOTANY, VOLUME 24

Plants large, 2-3 meters high, purplish, glabrous; leaves long-sheathing, the
blades as much as 90 cm. long and 40 cm. wide, filiform-apiculate, deep green
above, purple-marginate, vinous-purple beneath; racemes many-flowered, the
branches purplish-pruinose; bracts 2-flowered, oblong, short-acute, the lowest
2.5-3 cm. long, 10-12 mm. wide; sepals lanceolate, obtuse, 1.5 cm. long; petals
lanceolate, connate only at the base, acuminate, 4 cm. long; staminodia 2, cuneate-
oblong, purple above or within, yellow outside, about 6 cm. long; lip of about the
same length, shallowly 2-lobate; whole flower 7 cm. long or more; capsule 3-4 cm.
long, more than 2 cm. broad.

Canna edulis Ker, Bot. Reg. 9: pi. 775. 1823. Platanitto;

Cucuyus; Lirio.

Moist or wet thickets, sometimes in rather open forest, often in
second growth, 2,500 meters or less, most common at lower eleva-
tions; Alta Verapaz; Zacapa; Santa Rosa; Escuintla; Guatemala;
Quiche"; Quezaltenango; San Marcos. Southern Mexico; British
Honduras to Panama; West Indies; South America.

Plants large and coarse, usually 2-3 meters high, the rhizomes very large,
thick, and tuber-like; leaves large, oblong or ovate-oblong, acute, green, 60 cm.
long and 18 cm. wide or sometimes larger; inflorescence simple or branched;
sepals oblong-lanceolate, subacute, 12 mm. long; petals oblong-lanceolate, 4 cm.
long, 7-8 mm. wide; 3 upper staminodia oblong, much longer than the petals,
more than 6 cm. long, 12 mm. wide; lip slightly shorter, recurved into almost a
circle, orange-red or orange; capsule obovoid; whole flower connivent and little
spreading, 7.5 cm. long.

Called "bijao" in Honduras; "chancala" (Yucatan, Maya). This
plant is common and widely distributed in the lowlands of Guate-
mala, and often is more or less a weed. Like most of the native
species, it has small flowers, as compared with cultivated cannas.
In the field it is easy of recognition by its large size and orange-red
flowers. The starchy rhizomes are cooked and eaten in some parts
of tropical America, but so far as we know, no use is made of them
in Central America.

Canna generalis L. H. Bailey, Hortus 118. 1930. Bander a es-
panola; Bijagua.

This is the common large-flowered canna of gardens, of which
very many varieties are known, distinguished by size, coloring of
the foliage, and size, form, and color of the flowers. Most of the
cultivated cannas are believed to be of hybrid origin and of probably
very mixed ancestry. One of the principal elements in the ancestry
of the large-flowered plants is C. flaccida Salisb., native in the south-
eastern United States, which has as handsome flowers as most
cultivated forms. The native species, especially C. indica, often

FIG. 37. Canna indica. A. Inflorescence; X z /a- B. Fruit in position; X 1.
C. Leaf; X H.

205

206 FIELDIANA: BOTANY, VOLUME 24

are planted in Guatemalan gardens, but the majority of the culti-
vated plants are of "improved" varieties. They are planted in all
inhabited regions, but thrive best at middle or low elevations. They
often are seen half wild in hedges and in thickets near dwellings,
and normally spread so rapidly that the plants must be dug out in
order to protect less aggressive vegetation.

Canna indica L. Sp. PI. 1. 1753. Bijagua; Bijao; Cucuyus.

Mostly in moist or wet thickets, or along streams, frequent in
second growth, often invading cultivated ground, especially coffee
plantations, 1,900 meters or less, rarely naturalized at higher eleva-
tions; Alta Verapaz; Izabal; Zacapa; Jutiapa; Santa Rosa; Escuintla;
Sacatepe"quez; Quiche"; Huehuetenango; Suchitepequez; Retalhuleu;
Quezaltenango; San Marcos. Mexico; British Honduras to Salvador
and Panama; West Indies; South America.

Plants mostly 1-1.5 meters high; leaves oblong or ovate-oblong, the larger
about 50 cm. long and 20-25 cm. wide, green, acute, more or less unequal at the
base; racemes simple or sparsely branched, the bracts pruinose, the upper ones
broadly oblong, very obtuse, 1.5 cm. long; sepals 1 cm. long, much shorter than
the bracts, obtuse; petals lanceolate, acuminate, 3-4 cm. long, bright or deep red,
united only at the base; 3 outer staminodia obovate-spatulate or oblanceolate,
the lip narrower. (Fig. 37.)

Called "platanillo" in Honduras and elsewhere. This species
often is planted for ornament in Central America, and even in the
United States, but the flowers are too small to make the plant a
desirable one for cultivation. It was noted as growing abundantly
in a wet barranco near Chichicastenango, where it doubtless is
introduced.

Ganna lutea Mill. Gard. Diet. ed. 8. no. 4. 1768.

Moist or wet thickets, 350 meters or less; Retalhuleu; reported
from Escuintla. Mexico; Costa Rica and Panama; West Indies;
South America.

Plants usually about a meter high; leaves green, oblong or broadly lanceolate,
the larger about 40 cm. long and 13-15 cm. wide, acute; racemes mostly simple;
bracts pruinose, about 1 cm. long and wide, or the uppermost much smaller;
sepals oblong, acute; petals slightly connate at the base, lanceolate, acuminate,
pale or bright yellow; upper 2 staminodia connate at the base, ligulate, obtuse,
longer than the petals; whole flower 6 cm. long; capsule ellipsoid or obovoid, about
2.5 cm. long and 2 cm. broad.

This species seems to be rare in Central America, and few col-
lections of it have been made there.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 207

Canna sanctaerosae Kraenzlin, Pflanzenreich IV. 47: 40. 1912.

Known only from the type, Santa Rosa, Dept. Santa Rosa,
1,000 meters, Heyde & Lux 4290.

Plants large, glabrous; leaves oblong, 50 cm. long or more, about 23 cm. wide;
racemes simple or sparsely branched, the bracts oblong, obtuse or subacute, as
much as 2 cm. long; sepals oblong, subobtuse, 9-12 mm. long; petals lanceolate,
acuminate, short-connate at the base, 3.5-4 cm. long; staminodia 2, lanceolate,
acute, 5.5 cm. long, 7 mm. wide; lip linear, coiled into a circle; parts of the flower
connivent in anthesis except for the lip.

Perhaps only a form of C. lutea.

Canna Tuerckheimii Kraenzlin, Pflanzenreich IV. 47: 70. 1912.

Known only from the type, Cubilgiiitz, Alta Verapaz, 350 meters,
Turckheim 11.513.

Plants very large, 4-5 meters high; leaves very large, more than 70 cm. long
and 36 cm. wide, slightly paler beneath; inflorescence simple, elongate; bracts
early deciduous, the pedicels 13 mm. long; sepals subequal, ovate, obtuse, 13 mm.
long, accrescent in age; petals ovate-lanceolate, acuminate, connate only at the
base, the largest 6 cm. long and 1 cm. wide; staminodia 3, connate to the middle,
obtuse, reflexed at almost a right angle, 8 cm. long, toward the apex 8-9 mm.
wide; whole flower about 6 cm. broad, deep orange.

MARANTACEAE

Reference: K. Schumann, Marantaceae, Pflanzenreich IV. 48.
1902.

Perennial herbs, small or often very large, acaulescent or often with elongate
stems; leaves distichous, penninerved, usually vaginate, the sheath rarely ligulate
at the apex, the petiole commonly callose at the apex; flowers small or large,
perfect, dichlamydeous, very irregular, generally geminate, arranged in heads,
spikes, racemes, or panicles, each flower or group of flowers subtended by a con-
spicuous bract and usually by several bractlets; sepals free, equal or slightly
unequal; petals united below into a usually long tube, contorted, the outermost
one usually largest and more or less cucullate at the apex, white or colored; fertile
stamen 1, dimidiate, the anther 1-celled, dehiscent by a longitudinal slit; staminodia
2, another one usually present, this cucullate, all the staminodia free or connate
with the stamen, outer petaloid staminodia often also present; ovary inferior,
3-celled or 1-celled; ovules solitary in each cell, erect, anatropous; style simple,
recurved at the apex and concealed by the cucullate staminodium, elastically
exserted in anthesis; fruit capsular, loculicidal, sometimes nut-like or baccate,
1-3-seeded; seeds hard and osseous, provided with a lobulate or lamellate aril;
embryo hippocrepiform, the endosperm abundant.

About 23 genera, in the tropics of both hemispheres. The genus
Myrosma also is represented in Central America.

208 FIELDIANA: BOTANY, VOLUME 24

Ovary 3-celled, the fruit usually 3-seeded. Inflorescence usually of one or more
dense simple spikes, rarely (in one species) branched but then dense and

head-like Calathea.

Ovary 1-celled, the fruits 1-seeded. Inflorescence often much branched and open.

Perianth purple; rachis of the inflorescence zigzag Thalia.

Perianth usually white or yellowish, rarely purple; rachis of the inflorescence not
zigzag.

Bracts of the inflorescence early deciduous Maranta.

Bracts of the inflorescence persistent.

Staminodia 1 ; bracts of the inflorescence about 4 cm. long . . . Ischnosiphon.
Staminodia 2; bracts of the inflorescence 1.5 cm. long or less. .Stromanthe.

CALATHEA G. F. W. Meyer

Perennial herbs, the stems leafy at the base, simple or rarely branched; leaves
vaginate, on long or short petioles, the sheath not produced at the apex; flowers
spicate, the spikes sessile or pedunculate, the peduncle naked or bearing a single
leaf or rarely 2 leaves; bracts usually spirally arranged and closely imbricate,
rarely distichous; pairs of flowers usually more than 3 in each bract, the bractlets
membranaceous or indurate, rarely none; sepals equal or nearly so, small or large;
corolla tube equaling or much longer than the sepals; outer staminode solitary,
usually large; ovary 3-celled; fruit capsular, small, cartilaginous-thickened at the
apex, sometimes 3-corniculate, crustaceous or harder, 3-celled; seeds trigonous,
convex dorsally, arillate.

Species about one hundred, all in tropical America. A few
besides those listed here are found in other parts of Central America.

Bracts of the flower spike few, usually 5 or fewer, the whole spike 2 cm. long or

less. Plants low, the peduncle leafless, very slender C. microcephala.

Bracts of the flower spike usually numerous, at least more than 5, the whole spike

usually much more than 2 cm. long.
Bracts distichous, usually coriaceous or pergamentaceous, not green. Plants

usually very tall and coarse.
Bracts hard, coriaceous, as much as 4 cm. long, the spikes subterete or at least

not strongly compressed C. lutea.

Bracts soft and scarious or parchment-like, 3 cm. long or shorter, the spikes

strongly compressed C. insignis.

Bracts spirally arranged, not at all distichous.
Scape leafless.

Flower heads about 8 cm. broad, about as broad as long C. altissima.

Flower heads or spikes 3.5 cm. broad or less, longer than broad.

Bracts mostly 6 cm. long or longer, long-attenuate C. verapax.

Bracts less than 1 cm. long, obtuse or acute C. coccinea.

Scape bearing one or more large leaves.

Scape bearing several leaves; leaves small, 6 cm. wide or less.

C. Warscewiczii.
Scape bearing a single leaf or, if 2-3, the leaves much larger.

Inflorescence sessile; lowest bract much longer than the upper ones,

usually exceeding the inflorescence C. macrochlamys.

Inflorescence long-pedunculate; lowest bract similar to the upper ones
and always much shorter than the inflorescence C. Allouia.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 209

Calathea Allouia (Aubl.) Lindl. Bot. Reg. 14: sub pi. 1210.
1828. Maranta Allouia Aubl. PL Guian. 3. 1775. C. macrosepala
Schum. Pflanzenreich IV. 48: 83. 1902. Phrynium violaceum Roscoe,
Monandr. PL pi. 37. 1828. C. violacea Lindl. Bot. Reg. 11: sub
pi. 932. 1825. C. Allouia var. violacea Woodson, Ann. Mo. Bot.
Gard. 29 : 332. 1942. Mox, Max (Coban, Quecchi) ; Chufle; Platanillo.

Moist or wet, mixed forest, 1,400 meters or less, most frequent
at low elevations; Izabal; Zacapa; Santa Rosa; Escuintla; Sacate-
pe"quez; Suchitepequez; Huehuetenango. Southern Mexico; British
Honduras to Panama; West Indies; northern South America.

Plants large and coarse, 1-1.5 meters high, glabrous, the slender roots terminat-
ing in small tubers; radical leaves long-petiolate, the petioles slender, the callus
2 cm. long, puberulent above or glabrous; leaf blades oblong or ovate-oblong,
somewhat inequilateral, short-acuminate, as much as 60 cm. long and 20 cm. wide,
rounded at the base, thin, slightly paler beneath; scape bearing a single leaf similar
to the basal ones and long-petiolate; flower spike ellipsoid, 5-10 cm. long, long-
pedunculate, the peduncles short-pilose; bracts green, membranaceous, numerous,
spirally arranged, 2 cm. long, rounded at the apex, glabrous or sparsely appressed-
hirtellous; bractlets 12 mm. long, lanceolate; ovary villous at the apex; sepals
lanceolate, obtuse, 8 mm. long; corolla pale yellow or whitish, the tube 2.5 cm.
long, hirsute, the lobes oblong-lanceolate, 1 cm. long; outer staminode white,
12 mm. long, obovate, 2-lobate; capsule turbinate, 8 mm. long, stramineous,
the exocarp crustaceous; seeds trigonous, sulcate, grayish yellow.

The young tender flower spikes are cooked and eaten commonly
in Guatemala as well as in other parts of Central America. They
are seen frequently in the lowland markets. It is reported that in
the West Indies the tubers borne on the roots are cooked and eaten,
but we have no data to the effect that they are utilized in Central
America. About Coban and elsewhere the leaves are much used
for wrapping tamales and other foods. The nerves of the leaves
leave their impression upon the tamalitos, and this pattern is much
admired. The leaves remain soft and pliable, unlike those of some
other species, and have no unpleasant flavor. In some localities
of the Pacific bocacosta the cultivation of this and other Calatheas
for their leaves is an agricultural operation second only to the grow-
ing of maize. In Salvador the plant is sometimes called "macus."
Var. violacea is a form differing only in its purplish corollas. It is
common in Panama but probably does not extend so far north as
Guatemala.

Calathea altissima (Poepp. & Endl.) Koernicke, Bull. Soc.
Nat. Moscou 35, pt. 1: 141. 1862. Phrynium altissimum Poepp. &
Endl. Nov. Gen. & Sp. 2: 20. 1838. Hoja de cuero.

210 FIELDIANA: BOTANY, VOLUME 24

Moist or wet, mixed forest, 350 meters or less; Pete"n; Alta Vera-
paz; Suchitepequez. Honduras; Costa Rica; Panama; southward
to Peru.

Plants large and coarse, 2 meters high or less; leaves long-petiolate, the
callus 13 cm. long or less, complanate or subterete, glabrous; leaf blades oval-
oblong or oval, 60 cm. long and 30 cm. wide, rounded and abruptly short-pointed
at the apex, rounded at the base and abruptly short-decurrent, glabrous, almost
concolorous, thin but stiff; flower spike head-like, about as broad as long, 8 cm.
broad, long-pedunculate, the scape naked, about as long as the leaves; bracts
spirally arranged, green, the outer ones coriaceous, oblong-ovate or broader, as
much as 10 cm. long, acuminate, appressed-hispidulous, often cleft in age, the
inner ones shorter and narrower, densely pubescent; ovary sericeous; sepals 18
mm. long, lanceolate, pubescent at the apex; corolla cream-colored, pubescent,
the tube 3 cm. long, the lobes lanceolate, 13 mm. long; outer staminode yellowish,
broadly obovate, 18 mm. long.

Called "bijagiiillo" in Honduras.

Calathea coccinea Standl. & Steyerm. Field Mus. Bot. 23: 39.
1944.

Known only from the type, Dept. San Marcos, wooded rocky
slopes above Finca El Porvenir, along Rio Cabus, southern slopes
of Volcan de Tajumulco, 1,300-1,500 meters, Steyermark 37945.

A low perennial, the roots numerous, long and slender, each ending in an
ellipsoid tuber 2.5 cm. long; leaves absent at anthesis, unknown, the young leaves
sparsely and minutely puberulent beneath; scapes naked, very slender, glabrous,
about 30 cm. long; spikes 3.5-5 cm. long, 1.5-2 cm. broad, the bracts 7-8, red,
subdistichous or spirally inserted, imbricate, broadly ovate, abruptly acuminate,
glabrous or nearly so, 1 cm. long; bractlets elliptic-ovate, subobtuse, 8-9 mm.
long; ovary glabrous; sepals elliptic-oblong or oblong-oblanceolate, subobtuse,
10 mm. long, 5-nerved, glabrous; corolla red, glabrous, 3 cm. long, the lobes half
as long as the tube, acute; outer staminode 10-12 mm. long, obovate.

Calathea eximia (Mathieu) Koernicke, Gartenflora 7: 87. 1858.
Maranta eximia Mathieu, Cat. 1853. This was grown in Europe from
plants collected by Warscewicz, probably in Central America, and
has been ascribed to Guatemala. There is no evidence that it is
native here, and both its habitat and specific status are uncertain.

Calathea insignis Peters, in Mart. Fl. Bras. 3, pt. 3: 124. 1890.
P. crotalifera Wats. Proc. Amer. Acad. 24: 86. 1889 (type collected
near Izabal, Izabal, Sereno Watson). C. sclerobractea Schum. Pflan-
zenreich IV. 48: 73. 1902 (type collected between San Andre's Osuna
and Rochela, Escuintla, Preuss 1446). Machan; Mohan (Izabal,
fide Blake).

FIG. 38. Calathea insignis. A. Inflorescence, X M and detached leaf, X '/
B. Habit of plant; X 1 /.

212 FIELDIANA: BOTANY, VOLUME 24

Dense wet forest or wet thickets, 1,500 meters or less, most com-
mon at low elevations; Izabal; Alta Verapaz; Escuintla; Suchite-
pequez; Retalhuleu; Solola; San Marcos. Southern Mexico; British
Honduras to Panama; southward to Ecuador.

Plants large and coarse, often 2 meters high and probably even larger; leaves
long-petiolate, the callus about 10 cm. long, terete; leaf blades oblong or oval,
about 60 cm. long and 25 cm. wide or even larger, rounded and abruptly short-
tipped at the apex, rounded at the base, glabrous, concolorous or somewhat paler
beneath, stiff and paper-like; stems bearing 1 or more leaves, the spikes several
or numerous, long-pedunculate, 25 cm. long or shorter, 5-6 cm. wide, much com-
pressed, the bracts numerous, bronze-colored, about 3 cm. long, as broad as long,
rounded or emarginate at the apex, distichous, densely imbricate, coriaceous,
glabrous or pubescent only at the apex; ovary glabrous; sepals 1 cm. long, linear-
oblong, obtuse, glabrous; corolla pale yellow; capsule ellipsoid, 9 mm. long,
stramineous; seeds trigonous, convex dorsally, castaneous, aril white. (Fig. 38.)

Called "bijagiiillo" in Honduras; "hoja blanca" (Veracruz). The
leaves are used like those of C. lutea. In describing C. crotalifera
Watson stated that the specific name was given because the inflores-
cences suggested the rattles of a rattlesnake, and the resemblance
is rather suggestive. C. sclerobractea was separated by Schumann
on tenuous characters that seem to have little importance.

Calathea lutea (Aubl.) G. F. W. Meyer, Prim. Fl. Esseq. 10.
1818. Maranta lutea Aubl. PI. Guian. 3. 1775. C. discolor G. F. W.
Meyer, op. cit. 7. Hoja de sal; Maxdn, Moxan (Quecchi and Maya) ;
Hoja de cuero; Platanillo.

Wet forest or open or forested swamps and marshes, 300 meters
or less, but often planted at higher elevations; Pete"n; Alta Verapaz;
Izabal; Escuintla; Suchitepequez ; Retalhuleu; Quiche"; probably in
all the Pacific coast departments. Southern Mexico; British Hon-
duras to Panama; southward to Peru; West Indies.

Plants large and coarse, often 3 meters high or more; leaves large, on very long
petioles, the callus often 13 cm. long, glabrous; leaf blades often more than a
meter long and 60 cm. wide but mostly smaller, elliptic or oval, usually rounded
and apiculate at the apex, rounded at the base, thick and stiff, glabrous, pruinose
beneath but the pale coat often deciduous in large sheets; scapes bearing one or
more leaves, the spikes slightly compressed, numerous, often 20 cm. long; bracts
numerous, mostly dark reddish brown, broad, 4 cm. Jong, coriaceous and very
hard, distichous, glabrous or nearly so; ovary villous at the base; sepals linear,
obtuse, 1 cm. long; corolla pale yellow, the tube almost 3 cm. long, the lobes
elliptic, 1.5 cm. long, sericeous; outer staminode pale yellow, obovate, 2 cm. long;
capsule usually 1-seeded; seed 8 mm. long, grayish castaneous, the aril white.

Called "bijao" in Honduras. Although growing most often in
open swamps, this plant grows perfectly well in well drained soil,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 213

and sometimes grows naturally on open hillsides in regions of heavy
and continuous rainfall. It is of great economic importance in
Guatemala, especially along the Pacific foothills, where it often is
planted on a large scale. In some places from Retalhuleu to Escuin-
tla there are good-sized fields, with the plants set in rows. Great
quantities of the leaves are gathered, piled flat, and tied in large
bunches, which are transported to all parts of the country, and are
to be seen in almost all the markets. They serve in place of paper
for wrapping all kinds of articles, especially salt and soap. Banana
leaves when green are much used as temporary wrappers but when
dry they are altogether useless. Calathea leaves are tough, and serve
very well for this purpose. They are much used, too, as thatch on
small temporary structures.

Calathea macrochlamys Woodson & Standl. Field Mus. Bot.
22: 134. 1940. Hoja de cuero.

Wet forest, 150-200 meters; Huehuetenango (between Ixcan
and Rio Ixcan, Steyermark 49250). Oaxaca; Tabasco.

Plants large and coarse, a meter high or more, the scape bearing a large leaf,
sparsely pilose with subappressed hairs; leaves long-petiolate, stiff and paper-
like, the callus 8.5-11 cm. long; leaf blades oval or broadly oblong, about 50 cm.
long and 18-20 cm. wide, rounded and apiculate at the apex, rounded and abruptly
contracted at the base, almost concolorous; inflorescence arising from the axil
of the cauline leaf, sessile, at first 5-6 cm. long and 3-4 cm. broad, after anthesis
recurved and somewhat elongate, subpaniculate, with few short branches; basal
bract lance-oblong, 5.5-11 cm. long, long-attenuate, appressed, the inner bracts
numerous, oblong or ovate-oblong, 12-20 mm. long, pale brown, sparsely sericeous
or almost glabrous; sepals ovate-oblong, obtuse or subacute, 11 mm. long; ovary
glabrous, 3-celled; petals 3.

Called "pozolillo" in Oaxaca. The species is anomalous in
Calathea because of its branched inflorescence, but the 3-celled
ovary necessitates its reference here unless a new genus is erected
for it.

Calathea microcephala (Poepp. & Endl.) Koernicke, Bull.
Soc. Nat. Moscou 35, pt. 1: 125. 1862. Phrynium microcephalum
Poepp. & Endl. Nov. Gen. & Sp. 3: 20. pi. 128, figs, a, b. 1838.
Maranta micans Mathieu, Cat. 1853. C. micans Koernicke, op.
cit. 126. C. albicans Brongn. ex Schum. Pflanzenreich IV. 48: 112.
1902. Yuquilla silvestre.

Dense wet forest, 200 meters or less; Alta Verapaz; Izabal;
Huehuetenango (near Ixcan). Tabasco; British Honduras along
the Atlantic coast to Panama; southward to Peru and the Guianas.

214 FIELDIANA: BOTANY, VOLUME 24

Plants small, about 30 cm. high, from a thick rootstock; radical leaves usually
numerous, on very long and slender petioles, thin, the petioles densely puberulent;
leaf blades asymmetric, elliptic to lance-oblong, mostly 15-19 cm. long and 3.5-6
cm. wide, acute or acuminate at each end; scape naked, very slender, equaling or
shorter than the leaves; bracts 3 or sometimes as many as 5, spirally inserted,
imbricate, ovate, short-acuminate, green, glabrous, 1 cm. long or less; ovary
glabrous; sepals linear-lanceolate, 7 mm. long; corolla white, the tube 8 mm.
long, the lobes oblong-lanceolate, 5 mm. long; outer staminode obovate, 7 mm.
long, unguiculate.

Calathea verapax Bonn. Smith, Bot. Gaz. 31: 124. 1901.

Wet mixed forest, 1,000 meters; Alta Verapaz (type from Rubel-
cruz, Turckheim 1269; collected also at Sacolal). Honduras.

Plants tall, slender, from a thick rootstock, the scape naked, the callus 4 cm.
long, puberulent or glabrous; leaf blades elliptic to oblong, about 40 cm. long and
16 cm. wide, rounded and short-acuminate at the apex, rounded at the base,
glabrous; spike turbinate, 10-12 cm. long, the bracts numerous, spirally arranged,
ovate-lanceolate, scarlet, glabrous, long-acuminate, as much as 6.5 cm. long;
ovary glabrous; sepals lanceolate, acuminate, 2 cm. long; corolla tube 3 cm. long,
the lobes lanceolate, acuminate, 1.5 cm. long.

Calathea Warscewiczii (Mathieu) Koernicke, Gartenflora 7:
87. 1858. Maranta Warszewiczii Mathieu, Cat. 1853. Culebrin.

Wet mixed forest, 500-800 meters; Suchitepequez; often grown
for ornament. Costa Rica.

Plants slender, 90 cm. high or less; cauline leaves usually numerous, on petioles
12 cm. long or shorter, often crowded on short lateral shoots, the callus 3 cm. long
or shorter, puberulent or glabrate; leaf blades lanceolate to elliptic-oblong, as
much as 30 cm. long and 13 cm. wide but mostly 15 cm. long or shorter; spikes
3-6 cm. long, generally short-pedunculate, the bracts broadly ovate, spirally
arranged, acute or rounded at the apex, few, about 1.5 cm. long, with conspicuous
scarious margins; ovary glabrous; sepals lanceolate; corolla tube 3 cm. long, or
shorter, the lobes oblong-lanceolate, 1.5 cm. long.

This plant is much grown in Guatemala and other parts of
Central America because of its handsome leaves. These are of
a beautiful velvety appearance on the upper surface, with broad
alternating stripes of light and dark green, while beneath they are
deep purple or brownish purple.

ISCHNOSIPHON Koernicke

Perennial herbs, often very large, the stems sometimes much branched;
leaves often very large, coriaceous or herbaceous; inflorescence consisting of few
or very numerous, terete or more or less compressed spikes, these solitary or clus-
tered; bracts coriaceous, strongly imbricate, appressed; sepals 3, free, equal, linear;

FIG. 39. lachnosiphon pruinosus. Inflorescence and detached leaf; X */*

215

216 FIELDIANA: BOTANY, VOLUME 24

corolla tube elongate, the lobes lanceolate or oblong-lanceolate; stamen tube
almost obsolete; outer staminode solitary, petaloid, obovate; anther with an
often very short, adnate appendage; ovary 1-celled, fruit capsular, crustaceous,
elongate, unequally 3-valvate; seed 1, elongate, acute, arillate at the base.

About fifteen species, in tropical America. Two or three others
are found in southern Central America.

Ischnosiphon pruinosus (Regel) Peters. Bot. Tidskr. 18: 264.
1892. Pleiostachya pruinosa Schum. Pflanzenreich IV. 48: 165.
1902. Platanillo.

Wet mixed forest, at or little above sea level; Pete"n; Izabal.
Chiapas; British Honduras, along the Atlantic coast to Panama.

A tall coarse herb 1.5-2.5 meters high; leaves long-petiolate, the callus at the
apex of the petiole terete, glabrous, about 6.5 cm. long; leaf blades mostly 40 cm.
long or less, as much as 22 cm. wide, acute or cuspidate, glabrous or essentially
so, green above, often purplish beneath; spikes 17 cm. long or less, 2-3 cm. wide,
usually several or numerous and forming a flabellate panicle; bracts 4 cm. long or
larger, glabrous, often pruinose; sepals 3 cm. long; corolla purple, the tube almost
3 cm. long, the lobes lanceolate, 1.5 cm. long; outer staminode elliptic, emarginate,
purple, 17 mm. long. (Fig. 39.)

Called "bijagiiillo" in Honduras.

MARANTA L. Arrowroot

Perennial or perhaps sometimes annual herbs, often with large starchy roots,
usually erect and branched; leaves small or large, thin, petiolate; flowers white,
racemose, the racemes sometimes paniculate, the bracts few, usually appressed,
green, finally deciduous; pairs of flowers several in each bract, pedunculate, the
terminal flower usually longer-pedicellate, the other one subsessile; bractlets none;
sepals equal, lanceolate or lance-oblong, striate; corolla tube longer or shorter
than the sepals, usually ampliate and gibbous at the base, often curved, the 3
lobes subequal, short-cucullate at the apex; stamen tube usually short; outer
2 staminodia petaloid, usually large and obovate, the other slightly shorter;
stamen free; style stout, the stigma obliquely 2-lobate; ovary glabrous or sericeous,
1-celled; fruit nut-like, indehiscent or sometimes 3-valvate, the epicarp coriaceous;
seed trigonous or short-pyramidal, often sulcate, arillate.

About twelve species, all native in tropical America. One other
species is known from southern Central America.

Ovary glabrous or pubescent only on the angles M. arundinacea.

Ovary densely white-sericeous or puberulent M. divaricata.

Maranta arundinacea L. Sp. PL 2. 1753. Yuquilla; Chuchute
(Huehuetenango) ; Tamalera (fide Aguilar).

Moist or wet thickets, sometimes on brushy rocky slopes, 1,300
meters or less, chiefly at lower elevations; Pete"n; Alta Verapaz;

FIG. 40. Maranta arundinacea. A. Uppermost flowering portion of plant;
X 1 A. B. Flower; X 2. C. Tuberous rhizome; X 2 /s. D and E. Outer petaloid
staminodia; X 1 1 A- F. Staminodium; X 1 1 A- G. Anther; X 1 1 A- H. Stami-
nodium; X 1 1 A- I. Style; X 1 1 A>

217

218 FIELDIANA: BOTANY, VOLUME 24

Izabal; Zacapa; Chiquimula; Jutiapa; Santa Rosa; Escuintla; Suchi-
tepequez; Huehuetenango. Mexico; British Honduras to Salvador
and Panama; South America.

Plants slender, erect, often much branched, usually a meter high or less, often
with rather large, tuberous rhizomes, glabrous or sparsely pilose; sheaths long,
broadly winged, the callus about 7 mm. long, terete, pilose at the apex; ligule
usually conspicuous, short, rounded; leaf blades ovate-lanceolate or lanceolate,
thin, mostly 20 cm. long and 8 cm. wide or smaller, attenuate-acuminate, rounded
at the base; inflorescence often large and much branched; bracts linear-lanceolate,
5 cm. long or shorter; pairs of flowers on peduncles 5 cm. long or shorter, the
terminal flowers on a pedicel 1 cm. long or shorter; ovary glabrous or pubescent
only on the angles; sepals about 15 mm. long, lanceolate; corolla pure white, the
tube ampliate at the base, about 13 mm. long, the lobes 8-10 mm. long; outer
staminodia obovate, emarginate, 10 mm. long, the inner half as long; fruit ellipsoid
or obovoid, 7 mm. long; seed pale red, rugulose, the aril yellowish. (Fig. 40.)

The Maya name of Yucatan is recorded as "chaac." This is a
common wild plant in the lowlands of Guatemala, where it is pre-
sumably native, and also is cultivated for its starchy roots. From
them is extracted arrowroot starch, which is used locally for starch-
ing cloth, and also as food. In some parts of the earth the plant is
cultivated extensively for the purpose. Many of the wild plants
have the appearance of annuals, and apparently have no starchy
rhizomes. During the dry season the stems and leaves wither and
die. The starch is much used in Guatemala in preparing atol and
other foods given to invalids, since it is easy of digestion.

Maranta divaricata Roscoe, Monandr. PI. pi. 27. 1828. Pla-
tanillo; doubtless called also Yuquillo.

Moist or wet thickets, 1,300 meters or less; Alta Verapaz; Izabal;
Chiquimula; Suchitepequez; Huehuetenango. Southern Mexico;
British Honduras to Costa Rica; South America.

Plants perennial, erect, in most respects exactly like the preceding species,
often much branched; bracts of the inflorescence linear, 2-4 cm. long; sepals
lanceolate, 13 mm. long; corolla pure white, the tube slightly longer than the
sepals; capsule 8 mm. long.

It is doubtful whether this is at best more than a poor variety
of M. arundinacea. Schumann separates it on the basis of smaller
leaves and slender, not thickened rhizomes. The leaves are of the
same size in both plants, and since the roots are seldom found with
herbarium specimens, it is not possible to use the root character
for practical purposes of identification.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 219

Maranta Friedrichsthaliana Koern. has been published and listed
from Guatemala but the plant is actually from Cerro de Aguacate,
Costa Rica.

STROMANTHE Sender

Mostly tall and coarse, perennial herbs arising from thick rhizomes, often
branched; leaves large, thin but stiff and paper-like, the cauline ones 2 or more;
inflorescences simply racemose or paniculate, open and lax or dense and congested,
the bracts rather remote or sometimes crowded, complicate, finally deciduous;
pairs of flowers usually numerous; bractlets none; sepals 3, free, lanceolate or
broader; corolla tube short, the lobes oblong; stamen tube very short; outer
staminodia shorter or longer than the inner ones; anther usually apiculate, the
filament appendaged; ovary 1-celled, smooth or tuberculate, glabrous or villous;
fruit capsular, 3-valvate; seed rugulose, arillate.

Species 15-20, in tropical America. A very few additional ones
are found in southern Central America.

Stromanthe lutea (Jacq.) Eichl. Abh. Akad. Berlin 1882: 81.
1883. Maranta lutea Jacq. Coll. Bot. 4: 117. 1794. Myrosma
guapilense Bonn. Smith, Bot. Gaz. 23: 251. 1897. Mojdn.

Dense wet mixed forest, at or little above sea level; Izabal.
British Honduras along the Atlantic coast to Panama; Colombia,
Venezuela, and northern Brazil.

A coarse herb 1-3 meters high, arising from a thick rhizome; basal leaves
long-petiolate, the cauline ones somewhat smaller and short-petiolate, the callus
about 2 cm. long, pubescent or glabrate; leaf blades elliptic or lance-oblong,
30-60 cm. long, 11-18 cm. wide, abruptly short-acuminate, rounded and abruptly
short-contracted at the base, green on both surfaces, puberulent or glabrous;
inflorescences long-pedunculate, lax and open, many-flowered, the racemes gemi-
nate, forming large panicles, the rachises glabrous; bracts yellowish, oblong, obtuse,
2.5 cm. long or shorter, pilose or glabrate, finally deciduous; pairs of flowers on
peduncles 6 mm. long or less, the terminal flower on a pedicel as much as 3 mm.
long; ovary glabrous or pubescent; sepals oblong-lanceolate, yellowish, 9 mm.
long; corolla yellowish, the tube 2 mm. long, the lobes oblong, 6.5 mm. long;
capsule trigonous, 8 mm. long, yellowish; seeds dark castaneous, with a yellow
aril. (Fig. 41.)

THALIA L.

Tall coarse aquatic herbs, often waxy-pruinose; leaves few, petiolate, thin but
stiff; inflorescence paniculate, usually lax and much interrupted, generally sub-
tended by a solitary leaf or a sheathing bract, the bracts deciduous or persistent;
pairs of flowers subtended by a bract, no bractlets present; sepals small, equal,
hyaline; corolla tube very short, the lobes hyaline; outer staminode obovate,
colored, usually unguiculate; stamen bearing a lateral appendage adnate to the
filament and equaling or longer than the anther; ovary 1-celled, 1-ovulate; style

FIG. 41. Stromanthe lutea. A. Inflorescence; X 2 /V B. Leaf; X
Fruit; X 4.

220

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 221

with a sometimes 2-lobate appendage at the apex; fruit nut-like, indehiscent,
the exocarp membranaceous; seed globose or ellipsoid, with a small aril at the base.

Species about seven, five in tropical America, one extending
northward into the southern United States, and one extending to
Africa. Only one is found in Central America.

Thalia geniculata L. Sp. PL 3. 1753.

In shallow water of open swamps, at or near sea level; Pete"n;
Izabal. Florida; Mexico; British Honduras to Salvador and Panama;
West Indies; South America; tropical Africa.

A tall, nearly glabrous perennial, usually 2-3 meters high; basal leaves long-
petiolate; callus of the petiole 1.5 cm. long, terete, glabrous or puberulent; leaf
blades rather stiff and paper-like, ovate-lanceolate or oblong-lanceolate, as much
as 60 cm. long and 20 cm. wide but mostly smaller; sheath of the petiole broad,
membranaceous, glabrous, not auriculate; panicles usually large, subtended by
a single large leaf, the racemes lax, with internodes 1 cm. long or less, the rachis
usually zigzag; bracts about 2 cm. long, oblong-lanceolate; ovary glabrous; sepals
oblong-ovate, 2 mm. long; corolla purple, the tube very short, the lobes 7 mm. long;
outer staminode purple, 14 mm. long, the inner staminodia half as long or shorter;
fruit ellipsoid, 1 cm. long, the exocarp membranaceous; seed grayish, minutely
tuberculate or smooth, subtended by a small whitish aril. (Fig. 42.)

Called "quento" in Tabasco; "platanillo" (Salvador). This is
a common and characteristic plant of the large open swamps along
the North Coast of Guatemala, and along the whole Atlantic coast
of Central America.

BURMANNIACEAE

Reference: F. P. Jonker, A monograph of the Burmanniaceae,
Utrecht, 1938.

Small, annual or perennial, often saprophytic herbs, usually without chloro-
phyll, the stems commonly very slender, the leaves alternate, generally reduced
to minute scales; flowers perfect, sometimes zygomorphic, usually regular; stem
bearing at its apex a single flower or a capitate or branched, usually bifid, inflores-
cence; perianth corolla-like, the limb consisting of 2 whorls of 3 segments, the tube
cylindric or trigonous, often winged or costate; anthers usually 3, sessile or nearly
so in the perianth throat, the anthers laterally dehiscent; style 3-branched; ovary
inferior, 1-celled, with axial placentae; ovules numerous; fruit commonly capsular,
irregularly dehiscent, or with transverse slits at the top, the seeds numerous, very
small.

The family is an essentially tropical one, with few representatives
in Central America. The plants are little collected, chiefly because
they are rather tenuous, and grow among grasses and other herbs
where it is difficult to discover them. One other genus is known from
Central America, Thismia, with a single species in Panama.

FIG. 42. Thalia geniculata. A. Inflorescence; X M. B. Leaf; X

222

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 223

Flowers subcapitate, clustered at the top of the stem Burmannia.

Flowers not capitate, in simple or bifid racemes.

Perianth limb deciduous; seeds subglobose to ellipsoid Gymnosiphon.

Perianth limb persistent; seeds linear.

Flowers in simple, lax racemes Apteria.

Flowers chiefly in bifid inflorescences Dictyostega.

APTERIA Nuttall

Slender annuals with almost filiform stems, the roots short, threadlike, the
stems usually simple, bearing few racemosely disposed flowers; leaves reduced to
small scales; flowers often nutant, the perianth campanulate or salverform, the
lobes 6, the outer ones ovate, the inner ones narrower but of the same length,
linear-lanceolate; tube of the corolla more than 3 times as long as the lobes;
stamens inserted in sacs in the perianth tube below the inner lobes, the filaments
short and thick, with a wing on the external side, the anther cells transversely
dehiscent; style filiform, elongate, trifid at the apex; capsule dehiscent between
the placentae, crowned by the persistent corolla; seeds minute, oblong to ovoid
or ellipsoid.

The genus consists of three species, the others South American.

Apteria aphylla (Nutt.) Barnhart ex Small, Fl. Southeast.
U. S. 309. 1903. Lobelia aphylla Nutt. Amer. Journ. Sci. 5: 297.
1822.

In swamps, at sea level; Izabal. Southern United States; Mexico;
British Honduras; Costa Rica; West Indies; Brazil; Bolivia.

Plants glabrous, usually simple, 5-20 cm. high, whitish or purplish; leaves
1.5-3 mm. long, scale-like; flowers 1 to several, long-pedicellate; perianth purple
or blue, sometimes white, 8-13 mm. long, the outer lobes one-third as long as the
tube, ovate, acute, the inner lobes lanceolate or linear-lanceolate, obtuse; capsule
ovoid or obovoid, 6 mm. long and 4 mm. broad or smaller; seeds brown, reticulate.

Perhaps the typical habitat for this delicate plant is grassland,
but the few plants found at Puerto Barrios were growing in a dark
Manicaria swamp.

BURMANNIA L.

Annual or perennial, usually saprophytic plants with or without chlorophyll;
stems simple or branched; leaves in the saprophytic species reduced to scales;
flowers solitary or clustered at the top of the stem in cyme-like or headlike inflores-
cences; perianth tube cylindric-trigonous, the 3 outer lobes much larger than the
inner, the latter sometimes minute or absent; anthers 3, sessile in the perianth
throat below the inner lobes; style filiform, shortly trifid; ovary trigonous; capsule
crowned by the persistent perianth, irregularly dehiscent; seeds numerous, oblong
or ellipsoid.

224 FIELD IANA: BOTANY, VOLUME 24

The genus is a large one, in tropical and subtropical regions of
both hemispheres. In Central America has been found one other
species, B. Wercklei Schlechter, in Costa Rica, with green leaves and
blue flowers.

Flowers about 3.5 mm. long, 2-many on each stem B. capitata.

Flowers 9-11 mm. long, usually solitary on the stems B. flava.

Burmannia capitata (Walt.) Mart. Nov. Gen. & Sp. 1: 12.
1824. Anonymos capitatus Walt. Fl. Carol. 69. 1788.

Wet grassland, usually in pine forest, at or little above sea level;
British Honduras (Honey Camp; All Pines). Southern United
States; Chiapas; Panama; West Indies; South America.

Plants slender, 3-30 cm. high, the wiry stems usually simple, bearing at the
apex a dense, headlike inflorescence of few or numerous small flowers; a few
basal, linear or linear-lanceolate leaves usually present, the cauline leaves scale-
like, 2 mm. long; flowers wingless, usually white or yellowish, erect, subsessile,
3.5 mm. long; perianth tube 1.5 mm. long, the outer lobes triangular, with involute
margins, 0.5 mm. long, the inner lobes linear or narrowly oblanceolate, obtuse,
not much shorter than the outer ones; seeds brownish yellow.

Burmannia flava Mart. Nov. Gen. & Sp. 1: 11. pi. 5. 1824.

Open pine ridges, at or little above sea level; British Honduras
(Toledo District, Swasey Branch, Monkey River, P. H. Gentle
3788). Southern Florida; Cuba; Costa Rica; Colombia to Paraguay.

Plants erect, very slender, annual, 6-25 cm. high; stems usually simple, some-
times branched, mostly 1-flowered but sometimes 2-9-flowered; basal leaves linear,
7 mm. long and 1.5 mm. wide or smaller, obscurely 3-nerved; stem leaves few,
scale-like, acute or acuminate, 2-4 mm. long; bracts lanceolate, acute or sub-
obtuse, 4-5 mm. long; flowers pale yellow, erect, sessile or short-pedicellate, 9-11
mm. long; outer perianth lobes erect, triangular, obtuse, 1.5-2 mm. long, the inner
lobes 1.5-2 mm. long, oblong or subspatulate; perianth tube trigonous-cylindric,
3.5-4 mm. long; wings of the perianth narrow, extending from the base of the
limb to the base of the ovary; capsule obovoid, 4-5.5 mm. long, irregularly dehis-
cent; seeds numerous, minute, oblong, yellowish brown, sometimes slightly curved.

DICTYOSTEGA Miers

Slender saprophytes, without chlorophyll, usually simple, glabrous, the root-
stock bearing ovate, acute or acuminate scales; leaves reduced to minute, attenuate
scales; inflorescence usually bifid, the flowers more or less secund along the
branches; perianth tube often constricted, the limb 6-lobate, the outer lobes erect,
ovate, acute, the inner ones smaller and shorter, ovate, obtuse to rounded at the
apex, erect or reflexed; anthers 3, sessile in the upper part of the perianth tube
below the inner lobes, not in sacs, the cells bursting transversely in the constriction;
style elongate, filiform in the lower part, dilated at the top and trifid; ovary 3-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 225

sulcate, with 3 parietal placentae; perianth persistent upon the fruit, the limb
not deciduous; capsule dehiscent by 3 irregular valves; seeds numerous, linear-
lanceolate, minute, the testa reticulate.

The genus consists of only two species, the other one South
American. The generic name has often been written Dictyostegia.

Dictyostega orobanchoides (Hook.) Miers, Proc. Linn. Soc.
1: 61. 1840. Apteria orobanchoides Hook. Icon. PI. 3: pi 254. 1840.
D. campanulata Karst. Linnaea 28: 422. 1856.

Dense, moist or wet, mixed forest, 1,500 meters or lower, Chi-
quimula (Cerro Tixixi, Steyermark 31589). Southern Mexico;
British Honduras; Costa Rica; South America.

Stems stouter than in most Central American plants of the family, yellowish
white, 7-30 cm. high; leaves reduced to ovate acute scales 1-4 mm. long, appressed;
pedicels 1-4 mm. long, curved or spreading; flowers white or pinkish, 2.5-5 mm.
long; perianth tube often constricted, the outer lobes 0.5 mm. long, the inner ones
smaller, at first erect, later reflexed; capsule obovoid or subglobose, 2.5 mm.
long or shorter; seeds yellowish.

GYMNOSIPHON Blume

Annual, erect saprophytes with very slender stems, usually simple in the
Central American species; inflorescence terminal, with few or numerous flowers,
bifid, the flowers somewhat secund, sessile or short-pedicellate, leaves minute,
scale-like; perianth limb 6-lobate, the outer lobes much larger than the inner ones;
stamens inserted below the inner perianth lobes, the anthers dehiscent by a
median, horizontal cleft; ovary ovoid or subglobose, the ovules numerous; style
elongate, shortly trifid at the apex; perianth limb circumscissile after anthesis
below the insertion of the stamens, the tube persistent upon the capsule; capsule
dehiscent at the apex or irregularly and longitudinally; seeds ovoid or subglobose,
reticulate.

About thirty species are known, widely distributed in the tropics
of America, Africa, Asia, and the East Indies. In Central America
one other species is reported, G. panamensis Jonker, of Panama.

Flowers 8-12 mm. long; stems stout G. suaveolens.

Flowers 9 mm. long or less; stems almost filiform G. Tuerckheimii.

Gymnosiphon suaveolens (Karst.) Urban, Symb. Antill. 3:
438. 1903. Benitzia suaveolens Karst. Linnaea 28: 420. 1928.

Dense wet forest, 1,800 meters or lower; Alta Verapaz; Huehue-
tenango. Southern Mexico; British Honduras; Costa Rica; Panama;
Colombia to Brazil.

Plants relatively stout, 8-30 cm. high, whitish, simple; inflorescence bifid,
3-17-flowered; leaves 1-3 mm. long, ovate or lanceolate, obtuse; pedicels 2-6 mm.

226

FIELDI ANA: BOTANY, VOLUME 24

long, the flowers 8-12 mm. long, white or blue; perianth tube 2-3.5 mm. long, the
limb 4-5 mm. long, the outer lobes ovate, obtuse, with lanceolate lateral lobes
equaling or surpassing the midlobe, the inner perianth lobes clavate, half as long
as the outer ones or shorter; capsule ellipsoid or obovoid, 3.5-5 mm. long.

FIG. 43. Gymnosiphon Tuerckheimii. A. Habit of plant; X 1. B. Flower
partly dissected; X 5. C. Fruit with attached perianth; X 5. D. Ovary, opened;
X 9. E. Upper part of style with stigmas; X 7. F. Stamen; X 7.

Gymnosiphon Tuerckheimii Jonker, Monogr. Burmann, 197.
1938.

Wet mixed forest, 350-1,700 meters; Alta Verapaz (type from
Cubilgliitz, Turckheim 11.475); Solola, British Honduras; Atlantic
lowlands of Honduras.

Plants white or purplish, 7-9 cm. high, usually simple, the inflorescence
bifid, 3-16-flowered, the branches 2 cm. long or less, the pedicels 1 mm. long;
leaves ovate, 1-1.5 mm. long; flowers up to 9 mm. long, the tube 4 mm. long, the
limb to 4 mm. long, the outer lobes ovate, obtuse, the lateral lobes lanceolate, about
equaling the midlobe, the inner lobes small, ovate, acute; capsule subglobose or
obovoid, 1.5 mm. long. (Fig. 43.)

Guatemalan material has been reported under the name G.
tenellus (Benth.) Urban, which pertains to a South American
plant.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 227

CASUARINACEAE. Beefwood Family

Trees, the leaves reduced to scales, the branchlets rigid, erect or pendulous,
often deciduous, frequently verticillate, usually jointed at the nodes; leaf scales
verticillate, small, appressed, often short-connate into a sheath; flowers unisexual,
the staminate in cylindric or 4-angulate, simple or compound spikes, terminal,
the pistillate forming globose or ovoid spikes or cones, these terminal or lateral;
flowers of both kinds solitary in the verticillate bracts or scales, sessile, bibracteo-
late; perianth segments 1-2 in the staminate flower, concave or cucullate, cir-
cumscissile at the base and pushed off by the expanding stamen; stamen 1, the
filament inflexed in bud, the anther large, the cells distinct, longitudinally dehis-
cent; perianth none in the pistillate flower, the ovary 1-celled; style short, the
branches linear, stigmatose from base to apex; ovules 2, affixed collaterally above
the base of the cell, ascending; fruit an ovoid or cylindric cone, the bracts and
bractlets in age indurate and accrescent, crowded, the whole cone somewhat
woody; nut laterally compressed, smooth, produced into a wing at the apex,
indehiscent; seed solitary, the testa membranaceous; endosperm none; embryo
straight, the cotyledons equal, complanate, the short radicle superior.

The family consists of a single genus, with about twenty-five
species, chiefly in Australia.

CASUARINA Forster. Beefwood

Casuarina equisetifolia Forst. Char. Gen. PI. 104. 1776. Pino
de Australia; Pinabete (Jutiapa).

Planted commonly in Guatemala, in parks, plantations, along
roadsides, and elsewhere, in the lowlands and in the mountains up
to at least 2,400 meters. Native of tropical Asia and Australia.

A large pine-like tree, sometimes 20 meters tall or more with a trunk a meter
in diameter, the branches pale green, slender, verticillate, often drooping, the
bark dark gray or blackish; branchlets very slender, resembling the stems of
Equiselum, somewhat angulate, the scales in whorls of 6-8, 1-3 mm. long, acute,
appressed, ciliate; staminate spikes 1-4 cm. long; fruit heads subglobose, about
2 cm. in diameter.

Called "pino" and "cipreY' in Yucatan. One of the common
park trees of Guatemala and all Central America. The tree is most
distinctive in appearance because of its unusual foliage. It gives but
little shade, because of the absence of normal leaves. It is one of
the trees that thrives in the vicinity of salt water. Probably it
grows without cultivation in some parts of Guatemala. The wood
is creamy yellow or pale pink, hard and strong. It is valued in
regions where the trees are native, but is not utilized in America,
even when available in some quantity, as in Florida. Some other
species besides C. equisetifolia are planted in Guatemala City, but
their determination is doubtful. It is not known how long C. equiseti-

228 FIELDIANA: BOTANY, VOLUME 24

folia has been planted in Central America, but specimens were col-
lected in Mexico more than a century ago. The bark is said to be
rich in tannin, and to give reddish and blue-black dyes in the regions
where the trees grow naturally.

PIPERAGEAE. Pepper Family

References: C. De Candolle, Piperaceae (of the West Indies), in
Urban, Symb. Bot. 3: 159-274. 1902; Piperacearum clavis analytica,
Candollea 1: 65-415. 1923. William Trelease, The Piperaceae of
Panama, Contr. U. S. Nat. Herb. 26: 15-50. 1927; The Piperaceae
of Costa Rica, op. cit. 115-226. 1929.

Herbs or shrubs, terrestrial or often epiphytic, rarely (in America) scandent,
pubescent or glabrous; leaves simple, alternate, opposite, or verticillate, entire,
palmate-nerved or penninerved; stipules present or none; flowers minute, usually
green, often whitish or pale yellowish, rarely dark red, bracteolate, without a
perianth, arranged in usually very dense, ament-like spikes, these pedunculate,
terminal or opposite the leaves, or sometimes axillary, rarely several together and
borne on a common peduncle; stamens commonly 2-6, hypogynous, the anthers
erect, the 2 cells distinct or confluent into one, longitudinally dehiscent, the
filaments free or rarely adnate at the base to the ovary; ovary superior, sessile
or rarely stipitate, 1-celled, 1-ovulate; style 1, the stigmas 2-5, or the stigma often
simple and penicillate; ovule erect, orthotropous; fruit small, berry-like, inde-
hiscent; seed small, with a membranaceous testa, the endosperm copious, the
embryo minute.

Genera 5 or fewer, only two of them reaching continental North
America.

The following account of the Guatemalan Piperaceae has been
prepared wholly by the senior author, and is based almost exclusively
upon the very ample material in the Herbarium of Chicago Natural
History Museum. The work has been greatly facilitated by the loan
of a substantial number of type specimens from the University of
Michigan Herbarium, and by about 200 sheets lent by the United
States National Herbarium. The latter group included a large
number of collections not represented in the Herbarium of Chicago
Natural History Museum, all or most of which had been marked
as new species by Trelease.

This family of plants, so far as it is represented in North America,
has had a rather unusual and decidedly unfortunate history which
can be appreciated fully only by one who has attempted recently to
work with the group, which, contrary to what some might think
after superficial study of the literature, is not one of the most difficult
families of tropical American plants. Its apparent difficulty results

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 229

from the actually very large number of specific entities involved, a
number that has been exaggerated by the somewhat eccentric
methods used in study of the Piperaceae by the two men who have
done the most work with the family.

Practically all the American Piperaceae were described during
the years 1869-1944, and during these years the group received
attention from only two men, Casimir De Candolle and William
Trelease. Most of the papers by the former were published in the
period from 1860 to 1923; most of those by the latter from 1927
to the 1940's. The only good account of North American Piperaceae,
excluding an account of Peperomia mentioned under that genus, is
that by De Candolle describing the West Indian species. De Can-
dolle's earliest work upon the family, in the Prodromus, also is good.
In later years, however, the same author, with more abundant new
material for study, became too enthusiastic in proposing new con-
tinental species, and named a very large number without apparent
reference to West Indian species, presumably assuming they must
be new because of their remote occurrence. Trelease believed
that the species of Piperaceae were extremely local, as many of
them are. However, he went still further, and assumed that those
of one country were distinct from all those (except a very few wide-
spread and more or less "weedy" species) of the adjoining countries.
Such narrow limits of distribution are not found in any other group
of Central American plants, nor, in the opinion of the writers, does
it exist in the Piperaceae. In his recognition of limiting geographic
units, Trelease was scarcely consistent. He treated the whole of
Mexico as one Piperaceous unit, and each of the Central American
countries as a unit of equal standing. Now each of these Central
American countries is about equal in area to one of the states of
Mexico, obviously making such an alignment improbable.

We are inclined to believe that instead of being local in their
occurrence the species of Piper and Peperomia, at least large numbers
of them, range very widely. Certainly many are common to the
West Indies and the mainland, and many of them extend from
Mexico to Costa Rica and Panama, and probably far southward
into South America. The treatment of the Guatemalan Piperaceae
here presented is a rather liberal one as regards specific lines, but it
is believed to afford an idea of the number of species of the family
occurring in the region. Ultimately some of the species here recog-
nized probably will have to be reduced, but this reduction in numbers
will be compensated by the discovery of species at present unknown

230 FIELDIANA: BOTANY, VOLUME 24

from the country, and by the division in some cases of groups of
plants here treated as a single species. So far as concerns the names
here used for the species of both Piper and Peperomia, the matter
is different. Aside from the few names whose original authors were
Linnaeus and Swartz, there are few that may not be supplanted by
some earlier name published from the West Indies, South America,
or Panama and Costa Rica.

In preparation of this account we have had available authentic
material in good quantity from Mexico, Salvador, Honduras, and
the West Indies, and the names from those regions have been used
when applicable. There have been available for study also photo-
graphs of most of the species described from Costa Rica and Panama,
but not usually actual specimens. Because of the slight characters
on which many of the species are based, names from those regions
have not been adopted here unless their application was obvious.
The nomenclature of the Central American species can not be
stabilized until all the West Indian and South American species
are considered, and such a task, involving monographic study of the
American Piperaceae, was out of the question.

Guatemala has been fortunate in that few Piperaceae have been
described from the country. Those that have been published seem
to be for the most part valid species, and it seems necessary to
publish here a number, fortunately not very large, of new Guate-
malan species. If, however, Trelease had lived to complete and
publish his studies upon the Guatemalan Piperaceae, the case
would have been very different. We submitted to him a large part
of the Piperaceae collected on the first three of the four Guatemalan
expeditions of this Museum, and he had completed their study at
the time of his death. He did not receive any of the material from
the fourth expedition.

The Piperaceae are a shining example of certain groups of tropical
plants that possess little or no practical or even esthetic importance,
yet occupy much space in a regional flora and require the expenditure
of a large amount of time for their treatment. The one important
member of this large family of plants is black pepper, Piper nigrum L.
(called in Guatemala "pimienta de Castilla"), whose dried fruits are
used throughout the earth as a condiment. Black pepper has been
grown at various places in the lowlands of Alta Verapaz, but
apparently it has not proved successful there. In the Quecchi
language black pepper is said to be called "caxlane"n." The fruits
of some of the native species of Piper have a faint taste of pepper,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 231

but they are not used locally for flavoring, so far as we know. The
dried spikes of various native species of Piper often are on sale in
the Guatemalan markets, for use in household medicine. The plants
find a good deal of use in country medicine for treating various ail-
ments. Dieseldorff reports the Quecchi name of one such medicinal
Piper as "ticrac ha," but the usual name for the species throughout
Guatemala and other parts of Central America is "cordoncillo," an
allusion to the cord-like spikes. Usually the country people have
no names for any of the species of Peperomia.

Plants herbaceous, usually epiphytic; leaves generally very thick and fleshy;
stigma 1 Peperomia.

Plants woody, or rarely herbaceous but then usually very large plants, almost
always terrestrial; leaves not fleshy; stigmas 2-5 (usually 3-4) Piper.

PEPEROMIA Ruiz & Pavon

References: Hugo Dahlstedt, Studien iiber Slid- und Central-
Amerikanische Peperomien, Svensk. Vet. Akad. Handl. 33, no. 2:
11-218. pis. 1-18. 1900; Arthur W. Hill, A revision of the geophilous
species of Peperomia, with some additional notes on their morphology
and seedling structure, Ann. Bot. 21: 139-160. pi. 15. 1907. William
Trelease, The peltate Peperomias of North America, Bot. Gaz. 73:
133-147. pis. 1-4. 1922.

Mostly small, perennial or rarely annual herbs, erect to prostrate, generally
epiphytic, rarely terrestrial; leaves alternate, opposite, or often verticillate, usually
thick and succulent; flowers very small, green, spicate, the spikes slender, solitary
or clustered, subtended by orbicular or elliptic bracts; stamens 2, the filaments
short or rarely longer than the bracts; anthers transverse-oblong or subglobose,
the two cells confluent to form one 2-valvate one; ovary sessile or contracted at
the base and substipitate, obtuse to rostrate at the apex; stigma undivided,
generally penicillate, terminal or lateral near the apex; ovule one, erect from the
base of the cell; fruit usually almost minute, with a thin pericarp, almost dry;
endosperm farinose.

Species probably almost 1,000, widely distributed in the tropics
of both hemispheres, but the great majority of them American.
Many others besides those listed here are found in other parts of
Central America. The plants of this genus are of little or no economic
importance, but some of them are much used in Guatemala in
domestic medicine, especially in the Coban region. Some of the
plants are of rather handsome appearance and suitable for cultiva-
tion. It may be remarked that of all Central American plants the
Peperomias are probably the most difficult to make into herbarium
specimens, since they yield up their moisture only under extreme

232 FIELDIANA: BOTANY, VOLUME 24

desiccation. Without artificial heat it is almost impossible to dry
them. Even if the leaves are scored or punctured at close intervals,
the openings quickly are filled with a sticky sap that hardens and
retains the internal sap indefinitely.

Leaves verticillate or opposite.

Flower spikes racemose; leaf blades cordate at the base P. Treleasei.

Flower spikes solitary; leaf blades not cordate at the base.

Leaves opposite (those at the apex of the branches sometimes verticillate).
Stems and leaves glabrous.

Leaves 3-nerved r P. pseudopereskiifolia.

Leaves 5-nerved (the nerves sometimes obscured in drying) . P. glutinosa.
Stems and leaves densely or sparsely pubescent, at least over part of their

surfaces.
Leaf blades orbicular, as broad as long or broader.

Leaves less than 1 cm. wide P. lenticularis.

Leaves about 1 cm. wide P. pecuniifolia.

Leaf blades various in shape but not orbicular.

Leaves very obtuse or rounded at the apex, mostly broadest slightly
above the middle P. humilis.

Leaves mostly acute or subacute or at least somewhat pointed at the
apex, broadest at or usually below the middle ... P. guatemalensis.
Leaves all or mostly verticillate.

Stems pubescent, usually densely so but the hairs often minute and incon-
spicuous.

Leaves small, mostly 7 mm. wide or narrower, chiefly cuneate-oblong or

oblanceolate-oblong, rounded or emarginate at the apex. P. galioides.

Leaves larger, mostly 10-20 mm. wide or widen, mostly broadest at or

near the middle.

Leaf blades orbicular or nearly so, broadly rounded at the apex, coria-
ceous when dried P. pecuniifolia.

Leaf blades various but not orbicular, mostly obtuse or subacute, at
least somewhat pointed at the apex, drying thin and not at all
coriaceous.

Stems very densely pilose with long spreading hairs ... P. hondoana.

Stems merely puberulent or sparsely pilose with short, mostly sub-
appressed hairs, never densely long-pilose.

Lower leaves very unlike the upper ones, rounded-obovate and
about 1 cm. long or less, the upper ones elliptic or ovate-
elliptic and 2.5 cm. long P. heterodoxa.

Lower leaves little if at all different from the upper ones.

Stems pubescent with very short, incurved hairs . P. Liebmannii.
Stems pilose with slender, spreading or subappressed hairs.

P. blanda.
Stems glabrous or essentially so.

Leaves mostly cuneate-obovate, narrowed to the base, broadest above
the middle, broadly rounded or emarginate at the apex.

Spikes glabrous P. quadrifolia.

Spikes puberulent P. Deppeana.

Leaves variable in shape but not cuneate-obovate, usually broadest at
the middle and about equally narrowed to each end, usually narrowed
at the apex and never broadly rounded or emarginate.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 233

Leaves thin when dried and usually blackish, narrowly acuminate.

P. tacticana.
Leaves coriaceous when dried, not blackening, obtuse.

Leaf blades rhombic-ovate or rhombic-elliptic, acute at the base,
somewhat narrowed to the narrow obtuse apex. P. crassiuscula.
Leaf blades oval or elliptic, mostly very obtuse at the base, scarcely
narrowed to the very obtuse or narrowly rounded apex.

Spikes glabrous P. staminea.

Spikes densely and finely puberulent P. reflexa.

Leaves alternate, or some of the uppermost opposite or at least appearing so be-
cause of shortening of the branches, or the leaves sometimes all basal.
Leaf blades peltate, at least most of them; plants often acaulescent.
Plants with elongate stems, these usually rooting at the nodes; leaves acute

or acuminate, or at least subacute.

Leaves very large, the principal ones 12-20 cm. long or larger. .P. maculosa.
Leaves much smaller, mostly 4-10 cm. long.
Leaves conspicuously pilose or villous beneath.

Leaves pilose on the upper surface P. vegana.

Leaves glabrous on the upper surface.

Leaf blades broadly rounded at the base, villous beneath with long

fc' spreading multicellular hairs P. agitato,.

Leaf blades merely obtuse at the base, pilosulous beneath with
I very short, often curved, not conspicuously multicellular hairs.

P. Griggsn.

Leaves glabrous beneath or obscurely puberulent.
Leaf blades small, mostly 3.5-4.5 cm. long.

Leaves very obtuse, peltate far above the base; spikes fusiform to

[ subglobose, obtuse or rounded at the apex P. clavigera.

Leaves acute, peltate slightly above the base; spikes obconic, flat

or depressed at the broad apex P. dorstenioides.

Leaf blades larger, mostly 6-8 cm. long P. peltilimba.

Plants acaulescent, or with short erect stems.

Leaf blades orbicular, not pointed, peltate at or very close to the middle,

glabrous P. campylotropa.

Leaf blades mostly broadly ovate and acute or at least pointed, often

peltate far below the middle, rarely suborbicular but then pilose.
Leaves conspicuously pilose or pilosulous on one or both surfaces.
Leaf blades small, 2 cm. long or less, rounded at the apex.

P. Tuerckheimii.

Leaf blades large, mostly 3.5-8 cm. long, acute P. molithrix.

Leaves glabrous, or rarely obscurely and finely puberulent.

Inflorescences branched; fruits sessile P. claytonioides.

Inflorescences simple; fruits short-pedicellate.

Leaf blades small, mostly less than 2.5 cm. long; fruits with a long

slender style P. mexicana.

Leaf blades large, mostly 3.5-7 cm. long; fruits with a very short

style P. peltata.

Leaves not peltate; plants not acaulescent.
Leaf blades cordate or subcordate at the base.
Leaf blades small, 3 cm. long or usually shorter.

Stems usually conspicuously hispidulous; leaf blades mostly reniform
and slightly broader than long, broadly rounded at the apex.

P. hispidula.

234 FIELDIANA: BOTANY, VOLUME 24

Stems usually quite glabrous; leaf blades mostly as long as broad or

usually somewhat longer, acute to rounded at the apex.
Leaf blades mostly acute or subacute at the apex and subcordate at

the base P. pellucida.

Leaf blades mostly rounded or very obtuse at the apex and rounded

at the base P. Skutchii.

Leaf blades larger, mostly 4-7 cm. long.

Leaves somewhat penninerved or plinerved, the innermost nerves arising
well above the base of the blade.

Spikes solitary P. aurorana.

Spikes geminate P. gymnophylla.

Leaves palmate-nerved, the nerves all arising at the very base of the
blade or close to it.

Leaf blades narrowly and deeply cordate at the base P. asarifolia.

Leaf blades shallowly and broadly cordate at the base or only sub-
cordate.
Plants prostrate, with elongate, creeping and rooting stems.

P. praetenuis.
Plants erect or suberect, with very short stems.

Leaf blades 2.5-4.5 cm. long; spikes 5-9 cm. long. . .P. Bernoullii.

Leaf blades 1 cm. long or less; spikes 1.5 cm. long or shorter.

P. major.

Leaf blades rounded to long-attenuate at the base, not at all cordate.
Leaves small, mostly 1 cm. long or shorter, orbicular or broadly oval.

Stems very slender, almost filiform, creeping.
Leaf blades narrowed to an obtuse, apiculate, often subemarginate apex,

the petioles very short P. suchitanensis.

Leaf blades broadly rounded at the apex, neither apiculate nor emargi-
nate; petioles long and slender P. rotundifolia.

Leaves larger, usually much more than 1 cm. long.

Leaves conspicuously pubescent on one or both surfaces.
Leaves penninerved.

Petioles mostly 3.5-6.5 cm. long.

Leaf blades mostly 5.5-7 cm. long P. Griggsii.

Leaf blades mostly 12-18 cm. long P. frigidula.

Petioles mostly 2 cm. long or shorter.

Leaves conspicuously long-ciliate, oval or broadly ovate.

P. minarum.
Leaves not evidently ciliate, lance-oblong or ovate-oblong.

P. floresensis.
Leaves palmately nerved.

Nerves evident beneath in the dried leaves P. santa-helenae.

Nerves obsolete or nearly so beneath in the dried leaves.

Leaves narrow, mostly oblanceolate to cuneate-oblong or cuneate-
obovate, acute to attenuate-cuneate at the base, mostly
broadest toward the apex P. pililimba.

Leaves broad, broadly ovate to suborbicular or broadly elliptic,
rarely broadly pbovate, mostly rounded or very obtuse at
the base, sometimes acute.

Leaves pilose on both surfaces with straight spreading hairs.

P. flagitans.

Leaves merely puberulent on both surfaces, often obscurely so.

P. lacanana.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 235

Leaves glabrous on both surfaces or practically so, rarely obscurely

puberulent, sometimes ciliate.

Spikes paniculate or racemose or geminate or solitary but then borne
on a long erect nodose peduncle bearing much reduced, foliaceous
bracts.

Spikes paniculate; leaf blades rounded or very obtuse at the base.

P. huitzensis.
Spikes not paniculate or, if so, the leaves acute to long-attenuate

at the base.
Leaves acute to long-acuminate, with a very acute apex.

Spikes solitary on a stout terminal leafy-bracteate peduncle,

3-4 mm. thick P. limana.

Spikes paniculate, 2 mm. thick or less.

Leaf blades mostly 5-6 times as long as wide ... P. floribunda.
Leaf blades 3 times as long as wide or broader .... P. cabana.
Leaves broadly rounded or emarginate to obtuse or subacute at
the apex, if acute the tip of the leaf obtuse. . . .P. obtusifolia.
Spikes solitary on the peduncles, the peduncles mostly solitary, axillary
or terminal, if terminal and clustered the peduncles not with
foliaceous bracts.
Leaves densely and conspicuously black-punctate beneath.

Plants normally terrestrial, erect, with a stout thick, trunk-like
main stem, the internodes very short, the leaves crowded.

Leaf blades acute or acuminate at the base P. petrophila.

Leaf blades rounded or very obtuse at the base . P. jilotepequeana.

Plants normally epiphytic, never with a trunk-like main stem,

usually procumbent or repent, the internodes much elongate.

Petioles and stems glabrous P. nigropunctata.

Petioles and stems sparsely short-pilose or hispidulous.

P. glabella.

Leaves not black-punctate beneath or very sparsely and incon-
spicuously so.
Leaves somewhat penninerved, with conspicuous strong branches

diverging from the costa above the basal nerves.
Leaf blades rounded or very obtuse at the apex, or merely
subacute, often emarginate, broadly ovate or rounded-
ovate, mostly 2.5-4.5 cm. long.

Leaves conspicuously emarginate at the apex . P. condormiens.

Leaves acute or subacute, not emarginate . P. praeteruentifolia.

Leaf blades acute to long-acuminate, mostly lance-oblong to

oblong-ovate, generally 6-12 cm. long.
Leaves not ciliate P. granulosa.

Leaves, at least most of them, conspicuously ciliate near the
base or throughout.

Leaf blades lance-oblong, on petioles 1 cm. long or shorter.

P. sisiana.
Leaf blades elliptic or ovate-elliptic, the petioles mostly

1.5 cm. long or longer P. luisana.

Leaves palmately nerved, the nerves all arising at or very close
to the base.

Stems conspicuously winged p. alata.

Stems not winged.

Plants procumbent or repent, with often much elongated
stems rooting at the nodes; leaves very unequal on the

236 FIELDIANA: BOTANY, VOLUME 24

same plant, part of them suborbicular, others lanceolate
or ovate P. aggravescens.

Plants mostly erect or ascending, not rooting at the nodes;
leaves mostly almost uniform in shape.

Leaves lanceolate or oblong-lanceolate P. coarctata.

Leaves variable in shape but mostly narrowly to broadly
ovate or elliptic P. collocata.

Peperomia aggravescens Trelease in Yuncker, Field Mus.
Bot. 17: 330. 1938.

Dense, moist or wet, mixed, mountain forest, 1,400-2,300 meters;
Alta Verapaz; El Progreso. Honduras (type from El Achote, near
Siguatepeque).

Plants slender, growing on tree trunks or on logs, glabrous throughout or nearly
so, the stems simple or branched, mostly 30 cm. long or shorter, usually repent
and rooting at the nodes, the tips sometimes erect, the internodes short or elongate;
leaves alternate, on short slender petioles, often fuscescent when dried, not pellucid-
punctate, very variable upon the same plant; blades of the lower leaves orbicular
or rounded-obovate, mostly 1-1.5 cm. long, rounded at the apex, rounded to
broadly cuneate at the base; blades of the upper leaves lanceolate to ovate,
generally 2-5 cm. long, acute to acuminate or sometimes obtuse, obtuse or acute
at the base, slightly paler beneath, obscurely 3-5-nerved; peduncles slender,
terminal, generally solitary, simple, naked; spikes very slender, mostly 6 cm. long
or shorter, 1 mm. thick, glabrous; bracts rounded, centrally peltate; ovary some-
what obliquely apiculate, the stigma subapical.

Peperomia agitata Trelease & Standley, sp. nov.

Moist or wet, mixed, mountain forest, terrestrial or sometimes
epiphytic, 1,500-2,400 meters; endemic; Quezaltenango; San Marcos
(type from Barranco Eminencia, above San Rafael Pie de la Cuesta,
Standley 68629; in Herbarium of Chicago Natural History Museum).

Plants slender, simple or sparsely branched, the stems succulent, prostrate
or repent, rooting at most of the nodes, the internodes elongate, glabrous, often
finely and rather conspicuously granular; leaves alternate, rather small, the
petioles slender, equaling or shorter than the blades; leaf blades thick and fleshy
when fresh, thin when dried and often fuscescent, not pellucid-punctate, broadly
ovate or rounded-ovate, mostly 5-8 cm. long and 3.5-4.5 cm. wide, rather abruptly
acuminate, rounded at the base, peltate well above the base, deep green and
glabrous above, somewhat paler beneath, pilose or villous-hirsute with rather
sparse but conspicuous, long, spreading, mostly yellowish hairs, about 7-nerved;
spikes unknown.

Planta gracilis, caulibus simplicibus vel sparse ramosis prostratis vel repen-
tibus, ad nodos radicantibus, internodiis elongatis glabris; folia inter minora alterna
longipetiolata in sicco saepe fuscescentia non pellucido-punctata, late ovata vel
rotundo-ovata, subabrupte acuminata, basi rotundata, 7-nervia, supra glabra,
subtus sparse pilosa vel villoso-hirsuta; spicae non visae.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 237

Peperomia alata Ruiz & Pavon, Fl. Peruv. 1: 31. pi. 48, f. b.
1798.

On wet shaded limestone rocks, 300-900 meters; Alta Verapaz.
British Honduras (Valentin); West Indies; South America.

Plants glabrous, the stems erect, solitary or clustered, simple or sparsely
branched, narrowly but evidently winged by the decurrent bases of the petioles,
succulent, with short or elongate internodes, sometimes decumbent at the base
and rooting at the lowest internodes; leaves alternate, on petioles 1 cm. long or
shorter, usually fuscescent when dry, pellucid-punctate, slightly paler beneath,
lanceolate to lance-oblong or oblong-elliptic, commonly 6-10 cm. long and 3-4 cm.
wide, abruptly acuminate or narrowly long-acuminate, attenuate to the base,
ciliolate, 3-5-nerved; peduncles terminal, longer than the petioles; spikes very
slender, densely flowered, sometimes 12 cm. long or more but usually shorter,
1-1.5 mm. thick, glabrous; bracts orbicular, centrally peltate, subsessile; ovary
half immersed in the rachis; fruits sparsely glandular, globose, obliquely and
obscurely mucronate at the apex.

Peperomia asarifolia Schlecht. & Cham. Linnaea 5: 75. 1830.
P. Heydei C. DC. in Bonn. Smith, Bot. Gaz. 19: 9. 1894 (type
from Laguna de Ayarza, Jalapa, Heyde & Lux 3834). P. Heydei
var. minor C. DC. op. cit. 10 (type from Estanzuela, Santa Rosa,
Heyde & Lux 3835).

Terrestrial in moist or rather dry forest, often growing on rocks,
1,000-1,800 meters; Zacapa; Jalapa; Santa Rosa; Huehuetenango.
Southern Mexico.

Plants succulent, erect or suberect, about 30 cm. high or lower, glabrous
throughout, the stems solitary or several, simple or sparsely branched, very
thin, inconspicuously pellucid-punctate or epunctate, somewhat paler beneath,
the petioles very slender, those of the lowest leaves as much as 15 cm. long, those
of the upper leaves mostly much shorter; leaf blades ovate-orbicular or almost
reniform, mostly 6-11 cm. long and 4.5-11 cm. wide, broadly rounded to some-
what pointed at the apex, rather deeply and narrowly cordate at the base, 9-11-
nerved, the innermost nerves sometimes arising slightly above the base of the
blade; peduncles terminal and axillary, about equaling the petioles; spikes very
slender, equaling or exceeding the leaves, as much as 19 cm. long but often much
shorter, 1.5 mm. thick, glabrous; flowers mostly arranged in rather remote whorls,
the bracts orbicular; ovary obovoid, obliquely complanate at the apex and obli-
quely stigmatiferous, pale-glandular; fruits emersed and very shortly stipitate,
subglobose.

Peperomia aurorana Trelease & Standley, sp. nov.

Known only from the type, Dept. Guatemala, perhaps in the
barranco near La Aurora, alt. about 1,400 meters, Ignacio Aguilar
517 in 1940; in Herbarium of Chicago Natural History Museum.

238 FIELDIANA: BOTANY, VOLUME 24

Plants small, erect, about 15 cm. high, the stems rather stout, simple or with
a few thick succulent branches, glabrous; petioles very slender, 7.5 cm. long or
shorter, glabrous; leaf blades thin, green when dried, paler beneath, not pellucid-
punctate, ovate or broadly ovate, 4.5-6.5 cm. long, 3.5-4.5 cm. wide, acute,
broadly rounded at the base and shallowly cordate, 7-9-plinerved, the inner nerves
arising far above the base of the blade but below the middle; peduncles axillary,
short, slender, glabrous, simple, naked; spikes very slender, glabrous, pale green,
about 3 cm. long, little more than 1 mm. thick, laxly flowered, the bracts broadly
ovate, peltate, short-pedicellate, subacute.

Planta nana erecta, caule simplici vel pauciramoso crasso glabro; folia tenuia
longipetiolata epunctata, subtus paullo pallidiora, , ovata vel late ovata, acuta,
basi late rotundata et breviter cordata, 7-9-plinervia, nervis interioribus bene
supra basin laminae nascentibus; pedunculi axillares breves simplices nudi, spicis
gracillimis glabris ca. 3 cm. longis vix ultra 1 mm. crassis laxifloris; bracteae late
ovatae peltatae breviter pedicellatae subacutae.

The type was presumably collected in the Department of Guate-
mala but possibly at some distance from Guatemala City.

Peperomia Bernoulli! ("Bernouillii") C. DC. Linnaea 37: 367.
1871-73. P. violaefolia C. DC. in Donn. Smith, Bot. Gaz. 19: 8.
1894 (type from Palin, Guatemala, J. D. Smith 2580). P. izakoana
Trelease in Standl. Journ. Wash. Acad. Sci. 13: 366. 1923 (type
from Izalco, Salvador). P. Uphofii Trelease in Standl. Field Mus.
Bot. 22: 10. 1940 (type from Salvador).

Terrestrial, on moist or wet, shaded banks in forest, 1,000 meters
or lower; Escuintla; Guatemala; Suchitepequez (type from Maza-
tenango, Bernoulli 37). El Salvador.

Plants small, erect, mostly 12 cm. high or less, with a very short, succulent,
simple or sparsely branched stem, glabrous; leaves slender-petiolate, the petioles
4.5 cm. long or shorter; leaf blades green when dried, thin, not pellucid-punctate,
mostly rounded-ovate or rounded-deltoid, commonly 2.5-4.5 cm. long, acute or'
narrowed to an obtuse apex, rounded to subcordate at the base, usually 5-nerved;
peduncles solitary, simple, naked, terminal and axillary, the spikes very slender
and laxly flowered, much exceeding the leaves; bracts orbicular, peltate at the
middle, subsessile; ovary emersed, ovoid, the stigma apical, papillose; fruits sub-
sessile.

Peperomia blanda (Jacq.) HBK. Nov. Gen. & Sp. 1: 67. 1815.
Piper blandum Jacq. Icon. PI. Rar. 2: 2. 1786-93. ? Peperomia
blanda var. cobana C. DC. Candollea 1: 380. 1923; 3: 122. 1926
(type from Coban, Alta Verapaz, Turckheim 11.1754).

On trees or rocks in moist or wet, mountain forest, 900-2,400
meters; Alta Verapaz(?); Jalapa; Guatemala; San Marcos. Central
and southern Mexico; probably southward in Central America;
South America.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 239

Plants succulent, erect or ascending, the stems usually branched above, rather
densely pilose with long slender lax hairs, the internodes mostly elongate; leaves
generally quaternate, on short or elongate petioles, green or fuscescent when dried,
obscurely if at all pellucid-punctate, elliptic or ovate-elliptic, mostly 3-6.5 cm.
long and 1-3 cm. wide, narrowed to the obtuse or subacute apex, obtuse or acute
at the base, 3-nerved, thin when dried, thinly short-pilose on both surfaces;
peduncles mostly terminal and clustered, equaling or much longer than the petioles;
spikes slender, erect or recurved, 4-8 cm. long, 1-1.5 mm. thick, remotely flowered,
glabrous; bracts suborbicular, peltate near the middle, subsessile.

Peperomia campylotropa A. W. Hill, Ann. Bot. 21: 156. 1907.
P. bracteata A. W. Hill, Ann. Bot. 21: 155. 1907 (type from Huehue-
tenango, Sactos, C. &E. Seler 2731).

Terrestrial, in open or shaded, wet to rather dry places, often
in alpine meadows or on or about rocks, mostly at 2,400-3,600
meters; Huehuetenango. Central and southern Mexico.

Plants glabrous, acaulescent, arising from a hypogaean tuber, this subglobose,
1-2 cm. in diameter, emitting fibrous roots; leaves few or numerous, erect, the
petioles very slender, 3-8 cm, long, or sometimes as much as 20 cm.; leaf blades
orbicular or ovate-orbicular, 1.5-3.5 cm. long, peltate at the middle, not or very
obscurely pellucid-punctate, rather thin when dried, fleshy when fresh, 6-9-nerved;
peduncles slender, simple, naked, equaling or longer than the petioles; spikes
pale green, rather densely flowered, equaling or usually exceeding the leaves;
bracts ovate, acuminate, 1.5 mm. long; fruit globose-ovoid, 1.7 mm. long, ver-
ruculose.

Dahlstedt and most other authors have referred this plant to
P. umbilicata Ruiz & Pavon, which it much resembles, and it may
be preferable to treat it as a form or variety of that species. The
species is distinct from all other Central American ones. In general
appearance it is highly suggestive of Hydrocotyle umbellata (Umbelli-
ferae). In Guatemala it is noteworthy as being the only truly alpine
Peperomia. The Guatemalan specimens may be referable to P.
bracteata A. W. Hill, but the material appears to be conspecific
with that of the Mexican specimens referred by Hill to P. campylo-
tropa.

Peperomia cilibractea C. DC. Candollea 1: 360. 1923.

This name was published in a key to the species of Peperomia.
The type, according to a photograph of a specimen in the Berlin
Herbarium, is Bernoulli & Cario 2686, said to be from Guatemala.
The specimen is a poor one, consisting mostly of detached leaves,
and the key characters give so few details regarding the specimen
that we have been unable to associate it definitely with any of the

240 FIELDIANA: BOTANY, VOLUME 24

Guatemalan plants we have seen. The name is probably synonymous
with one of those listed here.

Peperomia clavigera Standl. & Steyerm., sp. nov.

Terrestrial on wet limestone cliffs, 300-500 meters; Alta Verapaz
(type collected along road between Chajmayic and Sebol, Steyer-
mark 45738, in Herbarium of Chicago Natural History Museum).

Plants glabrous, with more or less elongate stems, or the stems sometimes
short, the internodes often elongate; leaves alternate, the petioles slender, 5 cm.
long or less; leaf blades very thick and fleshy when fresh, fuscescent when dry,
sparsely pellucid-punctate, when fresh grayish green above and silvery white
beneath, rounded-ovate, mostly 3-4.5 cm. long and 3-4 cm. wide, very obtuse at
the apex, broadly rounded at the base, peltate some distance above the base,
doubtless palmate-nerved but the nerves wholly obscured in the dry leaves;
peduncles axillary, simple, bearing a small fleshy bract above or below the middle,
equaling or shorter than the petioles, slender; spikes fusiform to obovoid, sometimes
ellipsoid, usually 7-15 mm. long, 5-7 mm. thick, obtuse or rounded at the apex,
acute or subacute at the base, very succulent, very densely many-flowered; anthers
borne on short filaments; fruits emersed, densely crowded, spreading, globose-
ovoid, obtuse at the apex, contracted into the short capitellate apical stigma.

Planta glabra, caule brevi vel elongate et prostrato vel repente, internodiis
interdum elongatis; folia alterna in sicco fuscescentia, in vivo crasse carnosa,
longipetiolata, sparse pellucido-punctata, rotundato-ovata parva obtusissima, basi
late rotundata, bene supra basin laminae peltata, nervis obsoletis; pedunculi
axillares simplices supra vel infra mediam bracteatae, petiolis aequilongis vel
brevioribus; spicae fusiformes, obovoideae vel interdum subglobosae, carnosae,
dense multiflorae, 7-15 mm. longae 5-7 mm. crassae; fructus emersus globoso-
ovoideus obtusus, stigmate apicali capitellato.

A distinct species, remarkable for the very short and thick, glo-
bose to club-shaped flower spikes.

Peperomia claytonioides Kunth, Ind. Sem. Hort. Berol. 11.
1847 (described from cultivated plants said to be of Guatemalan
origin). P. ovatopeltataC. DC. Journ. Bot. 4: 133. 1866. P.pinulana
C. DC. Bot. Jahrb. 10: 289. 1888 (type collected above Pinula,
Jalapa, F. C. Lehmann 1693). P. claytonioides var. longiscapa C.
DC. ex Trelease, Bot. Gaz. 73: 138. 1922 (type from Guatemala,
the locality unknown). P. claytonioides var. pinulana Trelease,
op. cit. 139.

On shaded rocks or in rather dry soil, sometimes epiphytic, 1,200-
1,600 meters; Baja Verapaz; Jalapa; Huehuetenango. El Salvador.

Plants acaulescent, from a small globose hypogaean tuber; leaves usually few,
erect, glabrous, the petioles very long and slender; leaf blades ovate-orbicular,
3-7 cm. long, somewhat narrowed to an obtuse tip, broadly rounded or subcordate

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 241

at the base, peltate near or far below the middle, thin when dried, slightly paler
beneath; peduncles long and slender, sometimes 30 cm. long but mostly shorter,
erect, usually sparsely branched, sometimes simple; spikes very slender, sparsely
and remotely flowered, short-pedunculate, little more than 1 mm. thick, short
or usually elongate, glabrous; bracts ovate-orbicular, acute; fruits subglobose,
0.8 mm. in diameter, rugose, produced at the apex into a slender appendage, or
the appendage sometimes very short.

Peperomia coarctata Trelease & Standley, sp. nov.

On trees in moist or wet, mountain or lowland forest, rarely
terrestrial, 300-2,000 meters; endemic; Sacatepe*quez; Suchitepe-
quez; Retalhuleu; Quezaltenango (type from Quebrada San Geronimo
Finca Pirineos, lower southern slopes of Volcan de Santa Maria,
Steyermark 33365, in Herbarium of Chicago Natural History Mu-
seum).

Stems erect or pendent, usually much branched, slender or rather stout,
very sparsely and inconspicuously pubescent or glabrous, with short or elongate
internodes; leaves small, usually blackish when dried and thin, slightly paler
beneath, sparsely pellucid-punctate, the petioles slender, glabrous, 7 mm. long
or shorter; leaf blades lanceolate to elliptic-lanceolate or ovate, mostly 2.5-6 cm.
long and 1-2 cm. wide, acute to long-acuminate and with an obtuse or narrowly
rounded tip, cuneate at the base, glabrous, obscurely 3-nerved; peduncles solitary
and axillary or terminal and clustered, longer than the petioles; spikes very slender,
pale green, erect, remotely or densely flowered, mostly 3.5-6 cm. long, scarcely
1 mm. thick, glabrous; bracts very small, orbicular, centrally peltate, sessile;
ovaries semi-immersed.

Herba erecta vel pendens ramosa, sparsissime inconspicue pubescens vel
glabra; folia parva in sicco vulgo nigrescentia atque tenuia, subtus paullo pallidiora,
sparse pellucido-punctata; lamina lanceolata usque elliptico-lanceolata vel ovata
acuta vel longiacuminata basi cuneata glabra, inconspicue 3-nervia; pedunculi
singuli et axillares vel terminales et aggregati petiolis longiores; spicae gracillimae
erectae laxiflorae vel densiflorae, plerumque 3.5-6 mm. longae, vix 1 mm. crassae,
glabrae; bracteae parvae orbiculares centro peltatae sessiles; ovaria semi-immersa.

Peperomia cobana C. DC. in Bonn. Smith, Bot. Gaz. 19: 260.
1894.

On trees in moist or wet forest, 250-1,450 meters; endemic;
Alta Verapaz (type collected near Coban, Turckheim 78; known
only from the Coban region).

Plants glabrous, rather slender, the stems simple or sparsely branched, erect
or procumbent and rooting at the lower nodes; leaves alternate, the slender petioles
as much as 5 cm. long but mostly shorter; leaf blades thick and fleshy when fresh,
rather thick when dried and often fuscescent, opaque, elliptic to elliptic-lanceolate
or oblong-elliptic, mostly 10-13 cm. long, acute or acuminate, usually with an
acute tip, acute to attenuate at the base, the costa emitting 4-6 slender nerves
on each side, these often obsolete or obscure; peduncles terminal, half as long as

242 FIELDIANA: BOTANY, VOLUME 24

the leaves, bearing usually 3 slender-pedunculate erect spikes, the peduncles
naked, the spikes long and slender, densely flowered, 1 mm. thick; bracts orbicular, I
centrally peltate; ovary partly immersed, obliquely subulate at the apex, stigmati-
ferous anteriorly.

Peperomia collocata Trelease in Yuncker, Field Mus. Bot.
17: 332. 1938 (type from El Achote, above Siguatepeque, Honduras).
P. laudabilis Trelease in Yuncker, op. cit. 335 (type from El Achote,
Honduras). P. tressis Trelease in Yuncker, op. cit. 339 (type from
El Achote, Honduras). Siempreviva (Quezaltenango).

Dense, moist or wet, mixed, mountain forest, usually epiphytic
but sometimes terrestrial, 500-2,900 meters; Alta Verapaz; Zacapa;
Chiquimula; Jalapa; Guatemala; Sacatepe"quez; Chimaltenango;
Retalhuleu; Quezaltenango; San Marcos; Huehuetenango. Hon- ;
duras, and perhaps farther southward in Central America.

Plants erect or decumbent, the stems usually stout, simple or sparsely
branched, with short or elongate internodes, solitary or clustered, the plants
glabrous throughout; leaves alternate, thick and succulent when fresh, thin when
dried and often fuscescent, not pellucid-punctate, on petioles 1 cm. long or shorter;
leaf blades variable in size and shape, elliptic to rounded-elliptic, rhombic-ovate,
or rhombic-lanceolate, mostly 3.5-7 cm. long and 1.5-3 cm. wide, very obtuse to
acuminate with an obtuse tip, obtuse or broadly cuneate at the base, 3-5-nerved,
somewhat paler beneath; peduncles terminal and axillary, the terminal ones some-
times clustered, slender, longer than the petioles, simple, naked; spikes pale green,
sometimes 9 cm. long but usually shorter, 1-1.5 mm. thick, glabrous, erect or
recurved; remotely flowered; bracts orbicular, centrally peltate; ovary partly
immersed, obliquely apiculate, the stigma subapical.

The plant is a common one in many of the mountain areas,
and one of the most abundant of all the local Peperomia species.
The extensive collections that we have referred to this form
include 27 sheets annotated by Trelease as types of new species.
It is highly probable that a much older name can be found for this
plant. Two of the collections we place here were determined by
De Candolle as P. chrysocarpa C. DC., a Costa Rican species to
which all the material may be referable.

Peperomia condormiens Trelease in Yuncker, Field Mus.
Bot. 17: 332. 1938 (type from El Achote, above Siguatepeque,
Honduras). P. inaudax Trelease in Yuncker, op. cit. 334 (type
from El Achote, Honduras).

Moist or wet forest, sometimes in forest of pine and Liquidambar,
1,500-2,400 meters, usually epiphytic; endemic; Zacapa (Sierra de
las Minas); Huehuetenango. Mountains of Honduras.

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 243

Plants glabrous throughout, the stems erect, decumbent, or sometimes pendent,
slender, only slightly succulent, with mostly short internodes, often rooting at
the lower nodes; leaves small, somewhat coriaceous when dried, alternate, not
pellucid-punctate, mostly green when dried, the petioles slender, 5 mm. long or
shorter; leaf blades orbicular to rounded-ovate, 1.5-3 cm. long, 1.5-3 cm. wide,
rounded at the apex and conspicuously emarginate, rounded to broadly cuneate
at the base, paler beneath, essentially penninerved but perhaps more accurately
5-7-plinerved with the inner nerves arising far above the base; peduncles terminal,
solitary, equaling or slightly longer than the petioles; spikes slender, pale green,
glabrous, densely flowered, erect, 3-7 cm. long, little more than 1 mm. thick;
bracts orbicular, centrally peltate, short-pedicellate; ovaries ovoid, semi-immersed,
glabrous, abruptly contracted at the apex into a short but distinct style.

A closely related species is P. tenella (Swartz) A. Dietr. of the
West Indies, which has very similar but narrower leaves. The leaves
of the West Indian species are uniformly much smaller than those
of the continental material, which apparently constitutes a fully
distinct species.

Peperomia crassiuscula Millsp. Field Mus. Bot. 2: 33. 1900.
P. Lundellii Trelease in Standl. Field Mus. Bot. 12: 406. 1936 (type
from Honey Camp, British Honduras, C. L. Lundell 96a). P.
Wagneri Trelease in Yuncker, Field Mus. Bot. 9: 276. 1940 (type
collected near Coyoles, Honduras).

Epiphytic or on moist rocks, 400 meters or lower; Pete*n; Alta
Verapaz; Zacapa. Yucatan Peninsula of Mexico; British Honduras;
Atlantic lowlands of Honduras.

Plants stout and coarse, erect or sometimes pendent, very succulent, glabrous
throughout, the stems sometimes 8 mm. thick, sulcate when dried; leaves very
thick and fleshy, coriaceous when dried, in verticils of 4-6, short-petiolate, opaque,
mostly rhombic-ovate, 2.5-7 cm. long, 1.5-4 cm. wide, subacute with an obtuse
tip, acute or subacute at the base, 3-5-nerved, but the nerves usually obscure;
peduncles terminal, solitary or several, simple, stout, generally much longer than
the petioles; spikes erect, straight or somewhat curved, mostly 5-11 cm. long,
often several times exceeding the leaves, laxly or densely flowered; fruits partly
immersed, globose-ovoid, somewhat obliquely apiculate at the apex.

The Maya name of Yucatan is reported as "exlabon ac." This
species and P. glutinosa may eventually be united as a single
species. For this it is probable that an earlier name may be found
among West Indian species.

Peperomia Deppeana Schlecht. & Cham. Linnaea 5: 180. 1830.
P. chucanebana Trelease ex Standl. Field Mus. Bot. 10: 156. 1931.

On tree trunks in moist or wet, lowland forest, 900 meters or
lower; Pete"n; Alta Verapaz; Izabal; Escuintla(?); Huehuetenango.

244 FIELDIANA: BOTANY, VOLUME 24

Southern Mexico; British Honduras; Honduras and probably farther
southward in Central America; reported from Brazil.

Plants glabrous, perennial, often forming dense clumps or mats, the stems
slender, with rather short internodes, short or often elongate, sometimes pendent,
generally rooting at the nodes; leaves thick and fleshy, pale green, usually very
pale when dried, opaque, in whorls of 3-4, rounded-obovate, mostly 5-7 mm.
long, rounded at the apex, subacute to almost rounded at the base, 1-nerved or
obscurely 3-nerved from the base; spikes terminal, on long slender simple
peduncles, erect, densely flowered, much exceeding the leaves, chiefly 1-1.5 cm.
long, 1.5 mm. thick, densely and rather conspicuously puberulent; bracts orbicular,
centrally peltate; ovary immersed; fruits very small, immersed, ovoid, with a
conspicuous, somewhat elongate style.

Trelease has labeled some of the Guatemalan material as repre-
senting two unpublished species. The name Peperomia polochicana
Trelease has been reported for a plant of Pete*n, but the name has
not been published formally.

Peperomia dorstenioides Standl. & Steyerm., sp. nov. Hoja
de culebra.

Known only from the type, Huehuetenango, on rocky bluffs in
dense wet forest, between Ixcan and Finca San Rafael, 200-800
meters, Steyermark 49410 (type in Herbarium of Chicago Natural
History Museum).

Plants acaulescent or nearly so, arising from a short slender creeping rhizome,
glabrous throughout; leaves small, fuscescent when dried, when fresh dark green
above with 3 silver-gray longitudinal bands, silvery beneath, the petioles slender,
3 cm. long or less; leaf blades thick and succulent when fresh, rather thin when
dry, densely pellucid-punctate, ovate or ovate-elliptic, mostly 3-4.5 cm. long and
1.5-2.5 cm. wide, acute or subacute, rounded at the base, peltate slightly above
the base, 3-nerved; peduncles very slender, terminal or axillary, about equaling
the petioles; inflorescence (spike) obconic or funnelform, about 9 mm. high and
broad, subacute at the base, truncate or depressed at the apex, naked outside,
the upper or inner surface densely flowered; fruits at least partly immersed,
globose, scarcely 0.5 mm. in diameter, rounded at the apex, slightly granular,
the minute stigma subapical.

Plantae subacaulescentes rhizomatosae glabrae; folia parva in sicco fusces-
centia, graciliter longipetiolata in vivo carnosa; lamina dense pellucido-punctata
acuta vel subacuta, basi rotundata, paullo supra basin laminae peltata, 3-nervia;
pedunculi gracillimi petiolis subaequilongi terminates vel axillares; inflorescentia
obconica vel infundibuliformis ca. 9 mm. alta et aequilata basi acutiuscula, apice
truncata vel depressa, extus nuda, apice intusque densiflora; fructus subimmersus
parvus subglobosus apice rotundatus, stigmate minuto subapicali.

A most curious and unusual plant, its inflorescences strongly
suggestive of those of some species of Dorstenia.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 245

Peperomia flagitans Trelease in Yuncker, Field Mus. Bot. 17:
334. 1938. Xaxpec (Coban, Quecchi).

Moist or wet forest, on trees or on mossy rocks or cliffs, often
on limestone, 1,500 meters or lower; Alta Verapaz; Izabal; Chi-
quimula; Sacatepe"quez(?); Huehuetenango. British Honduras;
Honduras, the type collected near Siguatepeque.

Plants erect to procumbent and rooting at the lower nodes, the stems stout
or slender, mostly 35 cm. long or less, densely pilose with soft spreading hairs,
the internodes short or elongate; leaves alternate, thick and succulent when
fresh, rather thin and often fuscescent when dry, pellucid-punctate, on short or
elongate petioles, variable in shape, orbicular to rounded-ovate or broadly elliptic
or rhombic-elliptic, mostly broadest at or below the middle, broadly rounded to
very obtuse at the apex, cuneate to rounded at the base, densely pilose on both
surfaces with long spreading soft hairs, mostly 5-nerved; peduncles terminal,
simple, naked, mostly solitary, densely pilose; spikes erect, rather stout, much
exceeding the leaves, about 6 cm. long and 2 mm. thick, densely flowered; bracts
centrally peltate, orbicular, pedicellate, densely ciliate.

There is some confusion between Trelease's description of P.
flagitans (which he describes as glabrous) and the plant here treated
under that name, which is densely pilose. The abundant Guate-
malan material agrees with a specimen of the Yuncker Honduran
collection cited as the type of P. flagitans.

Peperomia floresensis Trelease in Yuncker, Field Mus. Bot.
9: 275. 1940.

Epiphytic or on rocks or banks, sometimes on limestone, 500-
1,000 meters or lower; Alta Verapaz; Escuintla. British Honduras;
Honduras, the type from Las Flores, Department of Yoro.

Stems often much elongate, usually pendent, sometimes rooting at the nodes,
the internodes elongate, sparsely or rather densely pilose with chiefly spreading
hairs; leaves alternate, the stout petioles 1-2 cm. long, rather sparsely pilose;
leaf blades very thick and fleshy when fresh, rather thick and fuscescent when
dried, narrowly to broadly lance-oblong, mostly 4.5-10 cm. long and 1.5-4 cm.
wide, rather abruptly acuminate or acute, obtuse or rounded at the base, sparsely
hirsute on both surfaces with slender white spreading hairs, or in age sometimes
glabrate; peduncles axillary, solitary, equaling or longer than the petioles, pilose,
simple, naked; spikes slender, densely flowered, 6-15 cm. long, often pendulous,
in fruit as much as 6 mm. thick, the rachis glabrous; bracts pale green, orbicular,
centrally peltate; fruits emersed, cylindric, about 2 mm. long, oblique at the
apex, spreading at almost a right angle, often granular.

Peperomia floribunda (Miq.) Dahlstedt, Svensk. Vet. Akad.
Handl. 33, no. 2: 154. 1900. Erasmia floribunda Miq. Inst. Versl.
Med. 1842: 81. 1843. Peperomia miradoresiana C. DC. in DC.

246 FIELDIANA: BOTANY, VOLUME 24

Prodr. 16, pt. 1: 395. 1869. Piper patulum Mart. & Gal. Bull.
Acad. Brux. 10, pt. 2: 128. 1843, non P. patulum Bertol. 1840.
Piper miradorense C. DC. in DC. Prodr. 16, pt. 1: 380. 1869. Pepe-
romia macrophylla C. DC. in Donn. Smith, Bot. Gaz. 19: 8. 1894
(type from Palin, Guatemala, J. D. Smith 2579). Peperomia flori-
bunda var. macrophylla Dahlstedt, loc. cit.

Most often terrestrial but sometimes epiphytic, in moist or wet,
mixed forest, often on cliffs, sometimes on limestone, 500-2,500
meters; Alta Verapaz; Izabal; Escuintla(?) ; Guatemala; Chimal-
tenango; Suchitepequez; Retalhuleu; Quezaltenango; San Marcos.
Southern Mexico.

Plants glabrous, erect or nearly so, or sometimes with decumbent and rooting
bases, the stems simple or very sparsely branched, with mostly short internodes;
leaves alternate, on petioles mostly 3 cm. long or shorter, thin, narrowly oblanceo-
late or oblong-oblanceolate, mostly 12-25 cm. long and 2.5-4 cm. wide, attenuately
long-acuminate, long-attenuate to the base, pellucid-punctate, usually somewhat
paler beneath, penninerved, the nerves 10 or fewer on each side, ascending at a
very narrow angle; inflorescences axillary, slender-pedunculate, mostly equaling
or even longer than the leaves, paniculate, lax, bearing mostly 3-5 spikes, these
on rather short, slender peduncles, mostly 4-8 cm. long, very slender, at anthesis
scarcely 1.5 mm. thick; bracts usually close together, orbicular, peltate, almost
sessile; fruits narrowly cylindric, obliquely ascending, subacute, 1.5 mm. long.

A common plant of the more humid forests of the Pacific slopes
of the volcanoes, abundant in many localities, especially on very
wet banks. The ample Guatemalan material is remarkably uniform,
and Trelease indicated only one new species among the specimens
we refer here. He used the name P. macrophylla for the Guatemalan
plant, but there is no obvious reason for supposing that it is different
from the Mexican plant whose name we use for the species.

Peperomia frigidula Trelease & Standley, sp. nov.

Terrestrial in dense wet mixed mountain forest, 1,400-1,600
meters; endemic; Alta Verapaz (type collected near Tactic, above
bridge across Rio Frio, Standley 89493, in Herbarium of Chicago
Natural History Museum; known only from this general region).

Plants stout, erect, 40-70 cm. high, the stems about 7 mm. thick, succulent,
generally simple, with short internodes, rather densely pilose with slender, mostly
appressed hairs; leaves alternate, thick and fleshy when fresh, rather thin and
green or fuscescent when dried, the petioles mostly 4-6 cm. long, appressed-
pilose; leaf blades elliptic or oblong-elliptic, commonly 9-16 cm. long and 4-7 cm.
wide, rather abruptly and shortly acute-acuminate, acute or subobtuse at the
base, scarcely if at all pellucid-punctate, deep green above, thinly appressed-
pilose with rather long hairs, slightly paler beneath, sparsely appressed-pilose or

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 247

glabrate, penninerved, the nerves 4-5 on each side, ascending at an acute angle;
spikes unknown.

Planta alta erecta, caule vulgo simplice crasso, internodiis brevibus subdense
pilis adpressis pilosis; folia magna alterna longipetiolata in vivo crassa carnosa,
in sicco tenuia, elliptica vel oblongo-elliptica, subabrupte breviter acuto-acuminata,
basi acuta vel subobtusa, supra sparse pilis longiusculis adpresso-pilosa, subtus
paullo pallidiora sparse adpresso-pilosa vel glabrata, penninervia, nervis utroque
latere 4-5 angulo angusto adscendentibus; spicae ignotae.

Peperomia galioides HBK. Nov. Gen. & Sp. 1: 71. pi. 17.
1815. P. galioides var. longifolia DC. ex Dorm. Smith, Enum. -PL
Guat. 4: 134. 1895, nomen.

Usually epiphytic in forest, often on oak trees, frequently in
rather dry regions, sometimes on rocks or on soil, 1,300-3,300
meters; Alta Verapaz; El Progreso; Zacapa; Chiquimula; Guate-
mala; Sacatepe"quez; Chimaltenango; Huehuetenango; Quezalte-
nango; San Marcos. Central and southern Mexico; El Salvador
and Honduras to Panama; northwestern South America.

Plants erect, perennial, succulent, mostly 30 cm. high or less, often forming
dense clumps on the branches or trunks of trees, the stems mostly simple below
but bushy-branched above, densely and finely puberulent, the internodes equaling
or mostly shorter than the leaves; leaves thick, generally pale green and often
yellowish, mostly in whorls of 4-5, spreading or often reflexed, sessile or nearly so,
the upper ones generally oblanceolate or linear-oblanceolate, the lower ones
cuneate-oblong or cuneate-obovate, chiefly 1-2 cm. long and 3-7 mm. wide, obtuse
or rounded at the apex, 1-nerved or obscurely 3-nerved, glabrous or practically
so, slightly paler beneath, pellucid-punctate and usually finely dark-punctate;
spikes terminal and axillary, usually several times as long as the leaves, solitary
at the apex of the stem or often geminate or several, on slender, short or elongate,
naked peduncles, 1-1.5 mm. thick, mostly 2-5 cm. long, remotely flowered;
bracts minute, orbicular, centrally peltate, sessile or nearly so; ovaries semi-
immersed, the stigma glandular, papillose; fruit ovoid, subrostellate at the apex.

This is a species of wide distribution, as here limited, only slightly
variable, of distinctive appearance and easy of recognition. We
find no basis for separating Mexican and Central American material
from that of South America, whence the species was described
originally. Trelease has described a number of its forms, if such
they are, from farther south in Central America, and has assigned
new names to about twenty of the Guatemalan collections. Like
many other local species, this plant sometimes sheds its leaves
during the dry season. The new leaves that develop after the next
rains fall are often longer and relatively narrower than the preceding
ones. In this and many other species the mature fruits are very
glutinous and adhere tenaciously to the fingers when the plants
are handled.

248 FIELDIANA: BOTANY, VOLUME 24

Peperomia glabella (Swartz) A. Dietr. Sp. PI. 1: 156. 1831.
Piper glabellum Swartz, Prodr. Veg. Ind. Occ. 16. 1788. Peperomia
caulibarbis Miq. Syst. Piper. 98. 1843. ?P. glabella var. microphylla
C. DC. ex Bonn. Smith, Enum. PI. Guat. 7: 30. 1905, nomen.
Peperomia longeacuminata Trelease ex Standl. Field Mus. Bot. 10:
157. 1931 (described from Lancetilla Valley near Tela, Honduras).
Peperomia pololensis Trelease in Standl. Field Mus. Bot. 12: 406.
1936 (type from Monte Polol, Pete"n, C. L. Lundell 3040). Peperomia
rubefacta Trelease in Yuncker, Field Mus. Bot. 17: 338. 1938 (type
from El Rincon, near Siguatepeque, Honduras). P. rubefacta var.
cangrejalensis Trelease in Yuncker, Field Mus. Bot. 9: 276. 1940
(type from Mount Cangrejal, near La Ceiba, Honduras). Verdolaga
(Izabal).

On trees in moist or wet, mixed, mountain or lowland forest, 2,100
meters or lower; Pete"n; Alta Verapaz; Izabal; Zacapa; San Marcos.
British Honduras; Honduras and probably farther southward in
Central America; West Indies; South America.

Stems prostrate, procumbent, or often pendent, rooting at the lower nodes,
slender but succulent, black-punctate, simple or branched, with elongate internodes,
almost glabrous but sparsely hispidulous on the angles made by the decurrent
petiole bases; leaves small, alternate, on slender petioles 1 cm. long or less, the
petioles hispidulous or at least ciliate; leaf blades obovate-elliptic to elliptic-
rhombic, mostly 4.5 cm. long and 2.5 cm. wide or smaller, short-acuminate to
very obtuse at the apex, more or less cuneate at the base, glabrous, densely black-
punctate, 5-plinerved, the inner nerves arising little above the base of the blade;
peduncles axillary, mostly solitary, simple, naked, about 1 cm. long, glabrous;
mature spikes 12 cm. long or shorter, 1 mm. thick, glabrous, rather sparsely
flowered, finely black-pun cticulate; bracts rounded, centrally peltate, subsessile;
ovary emersed, obliquely stigmatiferous below the apex; fruit ovoid, smooth,
obliquely acute at the apex, sessile.

This is very closely related to P. nigropunctata, and separated
only on trifling characters, but it is maintained as a distinct species
by both Dahlstedt and De Candolle, who, however, differ as to the
nomenclature of the two forms.

Peperomia glutinosa Millsp. Field Mus. Bot. 1: 293. pi. 12.
1896.

Usually epiphytic in moist or wet forest, sometimes on banks or
rocks, 1,400 meters or lower; Pete"n; Izabal; Zacapa; Chiquimula;
Jutiapa; Huehuetenango; San Marcos. Yucatan Peninsula of
Mexico; British Honduras.

Plants glabrous, stout, erect, as much as 40 cm. high, or sometimes procum-
bent, or with elongate pendent stems, these simple or sparsely branched, angulate

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 249

when dried, with elongate internodes; leaves all or mostly opposite, or the upper-
most ternate, very thick and fleshy when fresh, subcoriaceous when dry, opaque,
on stout petioles 1 cm. long or shorter, rhombic-ovate or elliptic-ovate, mostly
3.5-7.5 cm. long and 2-4 cm. wide, obtuse to acuminate with an obtuse tip, cuneate
at the base, 3-5-nerved, the nerves sometimes prominent beneath but often almost
wholly obscured ; peduncles terminal, mostly solitary and simple, generally several
times as long as the petioles; spikes erect or sometimes curved, as much as 13 cm.
long but mostly shorter, little more than 2 mm. thick, pale green, rather densely
or often sparsely flowered; bracts oval or suborbicular, centrally peltate; ovaries
partly immersed, globose-ovoid, somewhat obliquely apiculate.

Peperomia granulosa Trelease, Journ. Wash. Acad. Sci. 19:
328. 1929 (type from Lancetilla Valley near Tela, Honduras).
P. chicbulana Trelease in Standl. Field Mus. Bot. 17: 230. 1937
(type from La Libertad, Pete"n, C. L. Lundell 3099). P. dantoana
Trelease in Yuncker, Field Mus. Bot. 9: 274. 1940 (type from Rio
Dan to near La Ceiba, Honduras). P. riocangrejalensis Trelease in
Yuncker, op. cit. 275 (type collected near La Ceiba, Honduras).

On trees in moist or wet, lowland forest, 1,000 meters or lower;
Pete"n; Alta Verapaz; Izabal; Suchitepequez; Retalhuleu; Huehue-
tenango. Southern Mexico; British Honduras; Honduras and
probably farther southward.

Plants usually pendent, glabrous throughout or nearly so, the branches
slender, elongate, simple or sparsely branched, with often much elongate internodes;
leaves thick and fleshy when fresh, thick when dried and often blackish, usually
opaque, on petioles 2.5 cm. long or usually much shorter, lance-oblong or elliptic-
oblong, mostly 5-10 cm. long and 2-4.5 cm. wide, acuminate or long-acuminate,
acute or obtuse at the base, not ciliate or very obscurely so, paler beneath, penni-
nerved or 5-7-plinerved with the inner nerves arising far above the base, the
nerves usually not visible except against a strong light; peduncles terminal, solitary,
simple, naked, generally 1.5 cm. long; spikes slender, often pendent, densely
flowered, commonly 5-10 cm. long, scarcely 2 mm. thick at anthesis but thicker
in fruit, often curved, glabrous; bracts orbicular, centrally peltate, short-pedicel-
late; fruits emersed, widely ascending, oblong, about 1.5 mm. long or even larger,
obliquely short-rostrate, granular.

It is probable that there are several earlier names for this plant,
which is common and probably of wide range in the Central American
lowlands. We are using a name whose application is certain, instead
of referring the material with question to one of the West Indian or
South American species. The present species has been reported
from Guatemala under the name Peperomia distachya (L.) A. Dietr.

Peperomia Griggsii C. DC. Candollea 1: 364, 390. 1923; ex
Schroeder, Candollea 3: 125. 1926.

250 FIELDIANA: BOTANY, VOLUME 24

Moist or wet, lowland forest, on logs or rocks, 900 meters or
lower; endemic; Alta Verapaz (type from Finca Sepacuite", 0. F.
Cook & R. F. Griggs 253) ; Izabal.

Plants rather small and slender, the stems elongate, branched, repent, rooting
at the nodes, sparsely puberulent, the internodes elongate; leaves alternate, the
petioles slender, 3.5 cm. long or shorter, puberulent with mostly curved hairs;
leaf blades thick and succulent when fresh, thin when dried, green and only
slightly fuscescent, not pellucid-punctate, ovate or elliptic-ovate, 5-7 cm. long,
2.5-4 cm. wide, acute or short-acuminate, rounded at the base, peltate very slightly
above the base, glabrous above, paler beneath, densely puberulent with chiefly
curved hairs, about 7-plinerved, the innermost nerves arising far above the base;
peduncles terminal, short or elongate, each bearing 2 geminate secondary peduncles
and at their base a small foliaceous bract; immature spikes longer than the
secondary peduncles, very slender, erect, 2-3 cm. long, 1 mm. thick, glabrous,
very densely flowered.

Peperomia guatemalensis C. DC. in Bonn. Smith, Bot. Gaz.
19: 10. 1894.

Epiphytic or terrestrial on soil or rocks, moist or wet, usually
mixed, mountain forest, 600-3,300 meters; endemic; Alta Verapaz;
Zacapa; Sacatepe"quez (type from Acatepeque, J. D. Smith 2587);
Chimaltenango; Quezaltenango; San Marcos.

Plants rather slender but succulent, mostly 30 cm. high or less, the stems
simple or sparsely branched, erect or decumbent, sometimes rooting at the lower
nodes, the nodes elongate, sparsely or densely pilose; leaves thin when dried, chiefly
opposite, the uppermost sometimes verticillate, on short nr elongate, slender
petioles, lanceolate to broadly ovate, mostly 3-6 cm. long, acute or subacute,
with a usually obtuse tip, cuneate to obtuse at the base, pellucid-punctate, green
above, usually glabrous, somewhat paler beneath, sparsely or densely pilosulous,
generally 5-nerved; peduncles axillary or terminal, usually simple, solitary or
clustered, equaling or shorter than the petioles, naked; spikes slender, glabrous,
pale green, densely or sparsely flowered, about 1.5 mm. thick, often 10 cm. long;
bracts elliptic, peltate near the middle, subsessile; ovary partly immersed in the
rachis, obovoid, obliquely stigmatiferous at the apex; fruit minute, subglobose,
glabrous.

The material referred here is somewhat variable and includes a
dozen sheets marked by Trelease as types of new species or varieties.
It is questionable whether it would not be preferable to combine
all the material with P. humilis, from which it is by no means sharply
or satisfactorily separable.

Peperomia gymnophylla C. DC. Candollea 1: 363, 390. 1930;
ex Schroeder, Candollea 3: 126. 1926.

Known only from the type, Alta Verapaz, Finca Sepacuite",
0. F. Cook & R. F. Griggs 256.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 251

Plants glabrous, slender, the stems suberect or procumbent and rooting at the
nodes, with elongate internodes; leaves alternate, the petioles slender, sometimes
^6 cm. long but mostly shorter; leaf blades broadly ovate or rounded-ovate, 4-5.5
cm. long, 3-4.5 cm. wide, acute or rounded and subapiculate at the apex, broadly
rounded or usually subcordate at the base, green above, somewhat paler beneath,
multiplinerved, the inner nerves arising only slightly above the base of the blade;
peduncles terminal, as much as 4 cm. long, the spikes geminate, slender, about
7 cm. long and 2 mm. thick, rather closely flowered, the secondary peduncles
1-1.5 cm. long; bracts minute, rounded, peltate; ovaries ovoid, obliquely truncate
at the apex, the stigma anterior.

We know this species only by a photograph of the type, in the
United States National Herbarium. It does not seem to agree with
any recent Guatemalan collections.

Peperomia heterodoxa Standl. & Steyerm., sp. nov.

Known only from the type, Huehuetenango, Sierra de los
Cuchumatanes, between Santa Ana Huista and Rancho Lucas, 800-
900 meters, Steyermark 51340 (in Herbarium of Chicago Natural
History Museum).

Plants erect, the stems stout, simple below, sparsely branched above, as much
as 4 mm. thick below, the branches divaricate, with short or elongate internodes,
finely puberulent or in age glabrate; leaves mostly quaternate, the lower leaves
rather thick and fleshy, very shortly petiolate, oval, elliptic, or oval-obovate,
mostly 8-10 mm. long, rounded at the apex, subacute at the base, densely granular;
blades of the upper leaves thin, on slender petioles 4 mm. long or shorter, elliptic
or ovate-elliptic, mostly 2-2.5 cm. long and 7-12 mm. wide, acute with an obtuse
tip, acute at the base, green when dried, opaque or nearly so, deep green above,
pale beneath, glabrous, 3-nerved; peduncles terminal, solitary or clustered, about
as long as the petioles or elongate, the spikes very slender, longer than the leaves,
6 cm. long or more, scarcely 1 mm. thick, remotely flowered; fruits emersed,
minute, ovoid, slightly narrowed to the apex, the stigma subapical.

Planta erecta, caulibus crassis inferne simplicibus, superne sparse ramosis,
ramis gracilibus divaricatis, internodiis brevibus vel elongatis minute puberulis
vel glabratis; folia plerumque quaternata biformia, inferiora brevissime petiolata
crassa ovalia, elliptica vel ovali-obovata plerumque 8-10 mm. longa, apice rotun-
data, basi subacuta, dense granulosa; folia superiora tenuia breviter graciliter
petiolata, elliptica vel ovato-elliptica, vulgo 2-2.5 cm. longa et 7-12 mm. lata,
acuta, acumine obtuso, basi acuta, subtus pallida, glabra, 3-nervia; pedunculi
terminales singuli vel aggregati simplices nudi, spicis gracillimis vix 1 mm. crassis
laxifloris; fructus minutus ovoideus, stigmate subapicali.

The plant is noteworthy for its strongly dimorphous leaves, the
variation being much greater than we have observed in any other
species of the region. It bears some resemblance to P. L/iebmannii
C. DC.

Peperomia hispidula (Swartz) A. Dietr. Sp. PI. 1: 165. 1831.
Piper hispidulum Swartz, Prodr. Veg. Ind. Occ. 15. 1788.

252 FIELDIANA: BOTANY, VOLUME 24

Moist or wet mountain forest, 2,600 meters; Huehuetenango
(Sierra de los Cuchumatanes, between Todos Santos and San Juan
Atitan, Steyermark 51941). Southern Mexico; West Indies; South
America.

Plants terrestrial, small and very slender, usually repent, the stems succulent,
sparsely branched, hispidulous, at least near the nodes; leaves very small, alternate,
slender-petiolate, thin and translucent when dried, rounded or subreniform,
mostly 6-10 mm. long and 7-12 mm. wide, broadly rounded at the apex, emargi-
nate at the base, sparsely pilosulous on the upper surface, glabrous and minutely
punctulate beneath, obscurely 5-nerved; peduncles opposite the leaves, much
longer than the petioles, pilosulous below; spikes about three times as long as the
leaves, filiform, glabrous, sparsely flowered; bracts orbicular, centrally peltate,
subsessile; style about equaling the ovary; fruit ellipsoid, hispidulous, stipitate
at the base, mucronate at the apex; stigma small, glabrous.

Peperomia hondoana Trelease & Standley, sp. nov.

On shaded rocks or rather dry bluffs, 400-1,800 meters; endemic;
Zacapa (type from base of Sierra de las Minas, between Rio Hondo
and the waterfall, Steyermark 29401, in Herbarium of Chicago
Natural History Museum; also at Aguas Calientes); Huehuetenango
(between La Libertad and Paso del Boqueron).

Plants forming dense clumps, the stems erect or sometimes decumbent at
the base, stout, simple below, sparsely branched above, with very slender branches,
very densely villous-hirsute with long slender brownish spreading multicellular
hairs, the upper branches much less copiously villous, the internodes short or
elongate; leaves verticillate, mostly in whorls of 4-6, on short stout petioles;
lower leaves broadly obovate or rounded-obovate, mostly 1.5-2 cm. long, rounded
at the apex, broadly cuneate at the base, fuscous when dry, rather thin, not pel-
lucid-punctate, very densely pilose on both surfaces with slender spreading hairs;
upper leaves larger, on longer petioles, appearing thinner, cuneate-obovate to
obovate-elliptic, mostly 3-4 cm. long and 1.5-2 cm. wide, obtuse or rarely sub-
acute, acute at the base, sparsely short-pilose chiefly on the nerves, 3-nerved;
peduncles terminal, generally clustered, very slender, sparsely pilose, several times
as long as the petioles, the spikes longer than the leaves, sparsely flowered, 6 cm.
long or shorter, less than 1 mm. in diameter, glabrous; bracts centrally peltate,
sessile or nearly so, suborbicular.

Caules vulgo caespitosi crassiusculi erecti vel basi decumbentes, inferne
simplices, superne sparse ramosi, densissime pilis longis patentibus multilocularibus
brunneis villoso-hirsuti; folia plerumque 4-6-nata, breviter crasseque petiolata,
inferiora minora late obovata vel rotundo-obovata apice rotundata, basi late
cuneata, utrinque dense pilis tenuibus patentibus pilosa; folia superiora majora
tenuiora longius petiolata, cuneato-obovata vel obovato-elliptica, obtusa vel rare
subacuta, basi acuta, sparse praesertim ad nervos pilosa, 3-nervia; pedunculi
terminales aggregati sparse pilosi, spicis foliis longioribus gracillimis sparsifloris
glabris vix 1 mm. diam.; bracteae centre peltatae suborbiculares.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 253

Peperomia huitzensis Standl. & Steyerm., sp. nov.

Known only from the type, Huehuetenango, Cerro Huitz, between
Mimanhuitz and Yulhuitz, Sierra de los Cuchumatanes, 1,500-2,000
meters, Steyermark 48613 (type in Herbarium of Chicago Natural
History Museum).

Plants terrestrial, erect, about 30 cm. high, the stems rather slender, simple,
with short internodes, thinly pilose with weak yellowish hairs; leaves alternate,
the petioles rather slender, 2-3.5 cm. long, sparsely pilose or glabrate; leaf blades
broadly oval or elliptic, about 12 cm. long and 6-7 cm. wide, the apex not seen,
rounded at the base, rather densely pilose above with short weak hairs, slightly
paler beneath and similarly pilose, very sparsely and coarsely pellucid-punctate,
penninerved, the nerves about 6 on each side, obscure; peduncles terminal, about
equaling the petioles, sparsely and weakly pilose, bearing about 7 racemosely
disposed secondary peduncles, these slender, 2.5 cm. long or shorter; spikes very
slender, greenish white, 13-17 cm. long, little more than 1 mm. thick, rather
densely flowered, glabrous; ovules immersed in the rachis.

Planta terrestris erecta, internodiis brevibus sparse fulvescenti-pilosis; folia
modica alterna in sicco fuscescentia, graciliter longipetiolata, late ovalia vel
elliptica, basi rotundata, sparse grosse pellucido-punctata, utrinque sat dense pilis
gracillimis fulvescentibus laxis pilosa, penninervia, nervis utroque latere ca. 6;
pedunculi terminales singuli erecti, pedunculis secundariis ca. 7 racemose dis-
positis; spicae gracillimae vix ultra 1 mm. crassae glabrae sat densiflorae, bracteis
orbicularibus centro peltatis.

None of the few leaves present on the single plant of the type
sheet shows a perfect apex, and it is uncertain whether the leaves
are acute or obtuse. The species represented seems to be a very
distinct one.

Peperomia humilis (Vahl) A. Dietr. Sp. PI. 1: 168. 1831.
Piper humile Vahl, Enum. PL 1: 349. 1804. Peperomia Langsdorffii
Miq. Syst. Piper. 116. 1843. Peperomia Johnstonii Trelease in Standl.
Field Mus. Bot. 22: 9. 1940 (type from Volcan de Agua, Sacate-
pe"quez, John R. Johnston 220). Verdolaga (Jalapa).

Epiphytic or terrestrial in moist or wet, mountain forest, 1,600-
2,850 meters; Chiquimula; Jalapa; Guatemala; Sacatepe"quez ;
Chimaltenango; Solola; Suchitepequez; Quezaltenango; San Marcos;
Huehuetenango. Probably extending farther southward in Central
America and northward into southern Mexico; West Indies; South
America.

Plants usually erect and 40 cm; high or less, the stems slender or stout, simple
or sparsely branched, succulent, with mostly elongate internodes, rather sparsely
short-pilose with weak hairs or glabrate; leaves rather small, opposite or the
uppermost ternate, on slender petioles 1.5 cm. long or shorter; leaf blades thin,
at least when dried, obovate to elliptic or rounded-elliptic, generally broadest

254 FIELDIANA: BOTANY, VOLUME 24

above the middle, mostly 2-6 cm. long and 1-2.5 cm. wide, rounded or very obtuse
at the apex, cuneate at the base, sparsely or densely short-pilose on both surfaces
with slender, often yellowish hairs, green above, paler beneath, 3-nerved; peduncles
terminal, solitary or often clustered, mostly simple, short, the spikes very slender,
4-12 cm. long, 1-1.5 mm. thick, glabrous, laxly flowered; bracts suborbicular,
centrally peltate, short-pedicellate; ovary emersed, ovoid, obliquely stigmatiferous
slightly below the apex, the stigma globose, papillose.

As here treated this is a rather variable species, but not unreason-
ably so. Ten of the sheets were annotated by Trelease as types of
new species or varieties, but the material seems to be referable satis-
factorily to the common West Indian species, which heretofore has
not been recorded definitely from the mainland of North America.

Peperomia jilotepequeana Trelease & Standley, sp. nov.
Cigarrillo (Chiquimula).

Moist or wet banks, usually in dense forest, 1,500-2,000 meters;
Chiquimula (Cerro Brujo); Sacatepe"quez; Chimaltenango (type
collected along the road between Chimaltenango and San Martin
Jilotepeque, Standley 80901; also on Volcan de Fuego; type in
Herbarium of Chicago Natural History Museum). Southern Mexico.

Plants terrestrial, erect, 50 cm. high or less, the stems very thick, long-enduring,
with very short internodes, simple and naked below, sometimes with a few branches
above, leafy near the apex, the internodes when dry ochraceous and closely and
conspicuously transverse-rugose; leaves thin, not pellucid-punctate, densely and
finely black-punctate beneath, on long slender petioles 7 cm. long or shorter; leaf
blades ovate to oblong-ovate or elliptic-ovate, mostly 8.5-14 cm. long and 4-5.5
cm. wide, acuminate, rounded or very obtuse at the base, penninerved or plinerved,
with 3-4 nerves on each side, these all arising well below the middle of the blade,
ascending at a very narrow angle; peduncles terminal, simple, naked, solitary or
clustered, very slender, about equaling the petioles, 14 cm. long or shorter, 2 mm.
thick, rather densely flowered, the rachis glabrous; bracts small, orbicular, peltate
at the center, sessile or nearly so, black-punctate; ovaries immersed.

Planta terrestris erecta, caule crasso inferne simplici atque aphyllo, superne
interdum sparse ramoso, internodiis brevissimis transverse rugosis; folia tenuia
longe graciliter petiolata, opaca, utrinque dense nigro-puncticulata, ovata, oblongo-
ovata vel elliptico-ovata, acuminata, basi rotundata vel obtusissima, 7-9-plinervia,
supra glabra, subtus paullo pallidiora ad nervos breviter patenti-pilosa; pedunculi
terminales petiolis subaequilongi simplices nudi, solitarii vel pauci, spicis usque
ad 14 cm. longis, 2 mm. crassis densifloris; bracteae orbiculari-peltatae nigro-
puncticulatae.

Part of the material we refer here was marked by Trelease as
representing four other species and varieties. The plants apparently
lose all their leaves during the dry season, developing new ones each
wet season at the apex of the rather hard and almost woody
stem.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 255

Peperomia lenticularis Dahlstedt, Svensk. Vet. Akad. Handl.
3, no. 2: 154. pi. 5, /. 2. 1900.

On trees in moist or rather dry forest, 300-600 meters; Santa
Rosa; Retalhuleu (type from Retalhuleu, Bernoulli & Cario 2703).
El Salvador; Honduras; Brazil.

Plants very succulent, glabrous almost throughout, the stems much elongate
and slender, creeping and rooting at the nodes, bearing a few curved hairs; leaves
opposite, or the uppermost rarely ternate, lenticular and very thick, almost sessile,
suborbicular, broadly rounded at the apex, broadly rounded or somewhat sub-
cordate at the base, 5-10 mm. long and wide, sparsely ciliate; spikes mostly
terminal, the peduncles simple and solitary, short, hirtellous, the spikes dense,
4 cm. long or shorter, 1-1.5 mm. thick, the rachis glabrous; bracts small, rounded;
fruits 0.7 mm. long, obliquely appendaged at the apex.

Peperomia Liebmannii C. DC. Linnaea 37: 383. 1871-73.

On rocks in shaded moist places, 1,000-1,600 meters; Alta
Verapaz; Huehuetenango (Sierra de los Cuchumatanes). Southern
Mexico, the type from Oaxaca.

Plants erect, 35 cm. high or less, the stems slender but succulent, simple
below, sparsely branched above, puberulent with short curved hairs, the internodes
elongate; leaves small, in verticils of 3-5, the petioles slender, 12 mm. long or
less, puberulent; leaf blades thin, often brownish when dried, obovate-rhombic
or broadly elliptic, mostly 2-2.5 cm. long and 10-13 mm. wide, very obtuse or
subacute at the apex, acute or subacute at the base, puberulent or pilosulous on
both surfaces, somewhat paler beneath, ciliate, 3-nerved; peduncles axillary and
terminal, simple, the terminal ones often clustered, about equaling the petioles,
puberulent, naked; spikes longer than the leaves, slender, about 1 mm. thick,
densely flowered, the rachis glabrous; bracts suborbicular, centrally peltate,
sessile; ovary partly immersed, obliquely stigmatiferous at the apex, puberulent.

Peperomia limana Trelease & Standley, sp. nov.

Terrestrial or epiphytic in moist or wet, mixed or pine, mountain
forest, 1,800-2,800 meters; endemic; El Progreso (Sierra de las
Minas); Zacapa (type from Sierra de las Minas, in canyon along
Rio Lima, below Finca Alejandria, Steyermark 30048; in Herbarium
of Chicago Natural History Museum); Quiche"; Huehuetenango
(Sierra de los Cuchumatanes); San Marcos (volcanoes of Tacana
and Tajumulco).

Plants glabrous, erect, 40 cm. high or less, the stems simple, sometimes
decumbent at the base, very stout, with short internodes; leaves alternate, green
or fuscescent when dry, thick and fleshy when fresh, finely and densely pellucid-
punctate, the petioles stout or slender, 3 cm. long or shorter; leaf blades elliptic
to lance-elliptic or oblong-elliptic, mostly 10-14 cm. long and 4-5 cm. wide,
acuminate, with an acute tip, acute to attenuate at the base, deep green above,
paler beneath, penninerved, the nerves about 5 on each side, very slender, ascend-

256 FIELDIANA: BOTANY, VOLUME 24

ing at a very narrow angle, some of them arising above the middle of the blade;
peduncles terminal, solitary, very stout and thick, mostly 1 cm. long or shorter,
bearing near the apex a small or large leaf; spikes stout, erect or in age recurved,
commonly 8-20 cm. long, 3.5-4 mm. thick, very densely flowered; bracts orbicular,
centrally peltate, conspicuously granular.

Planta terrestris vel epiphytica glabra, caulibus simplicibus erectis vel basi
decumbentibus, internodiis brevibus; folia alterna longiuscule crasseque petiolata
crassiuscula, dense minuteque pellucido-punctata, in sicco viridia vel fuscescentia,
elliptica usque ad lanceolato-elliptica vel oblongo-elliptica, majuscula, acute
acuminata, basi acuta vel attenuata, penninervia, nervis utroque latere ca. 5;
pedunculi terminales singuli simplices crassi prope apicem folio parvo vel magno
onusti, spicis crassis erectis vel recurvis 3.5-4 mm. crassis densifloris; bracteae
orbiculares centro peltatae granulosae.

The plant has been reported from Quiche* (on the basis of Heyde
& Lux 3456) as P. adscendens C. DC.

Peperomia luisana Trelease & Standley, sp. nov.

On trees or cliffs in lowland forest, 500 meters or lower; endemic;
cultivated in Chiquimula; Escuintla (type from San Luis, north of
Escuintla, Standley 60134; in Herbarium of Chicago Natural History
Museum); Retalhuleu.

Stems often much elongate, pendent from cliffs or from the branches of trees,
stout, with mostly elongate internodes, often rooting at the lower nodes, glabrous
or when young sparsely pilose; leaves alternate, thick and fleshy when fresh,
rather thick and blackish when dried, inconspicuously or not at all pellucid-punc-
tate, the stout petioles mostly 1-2.5 cm. long, white-hirsute; leaf blades oval
to broadly ovate or oblong-ovate, mostly 5-8 cm. long and 2-5 cm. wide, acute or
short-acuminate, rounded or very obtuse at the base, glabrous, conspicuously
ciliate, penninerved or 5-7-plinerved, with the inner nerves arising far above the
base of the blade; peduncles slender, terminal, glabrous, simple, naked, solitary,
about 2 cm. long; spikes slender, as much as 8 cm. long, glabrous, densely flowered;
bracts orbicular, centrally peltate; fruits emersed, obliquely ascending, oblong,
about 2 mm. long, obliquely subrostrate, glabrous.

Caules elongati vulgo penduli crassiusculi, internodiis elongatis, glabris vel
juvenilibus sparse pilosis; folia alterna modica longiuscule petiolata crasse
carnosa in sicco fuscescentia, ovalia usque late ovata vel oblongo-ovata, acuta
vel breviter acuminata, basi late obtusa vel rotundata, glabra, ciliata, 5-7-plinervia,
nervis interioribus bene supra basin laminae nascentibus; pedunculi terminales
singuli simplices nudi glabri, spicis longis gracilibus densifloris; bracteae rotundatae
centro peltatae; fructus anguste oblongus oblique adscendens glaber oblique
subrostratus.

Each of the four sheets of this species was marked by Trelease
as the type of a new species.

Peperomia maculosa (L.) Hook. Exot. Fl. pi. 92. 1833. Piper
maculosum L. Sp. PI. 30. 1753. Peperomia variegata Ruiz & Pavon,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 257

Fl. Peruv. 1: 33. 1798. P. suavis var. hondurensis Trelease in
Yuncker, Field Mus. Bot. 17: 338. 1938 (type from El Achote, above
Siguatepeque, Honduras). Culantro (Quezaltenango).

Moist or wet forest, epiphytic or sometimes terrestrial, often on
limestone, 1,100-1,500 meters; Alta Verapaz; Chiquimula; Quezal-
tenango; Huehuetenango. Honduras; West Indies; South America.

Plants usually large, coarse, and very succulent, the stems simple or sparsely
branched, short or somewhat elongate, usually very thick, glabrous or pilose,
procumbent or repent and rooting at the lower nodes; petioles stout, mostly long,
often exceeding the blades, sparsely pilose or glabrous; leaf blades extremely
thick and fleshy when fresh, deep green above, pale beneath, opaque, subcoriaceous
when dry, oblong-ovate or elliptic-ovate, mostly 12-23 cm. long or even larger,
acuminate or long-acuminate, rounded at the base, peltate a short distance above
the base, glabrous above, glabrous or sparsely puberulent beneath, about 7-pli-
nerved, the costa conspicuous but the lateral nerves often obsolete; peduncles
rather short and stout, several times shorter than the spikes, terminal; spikes
slender, erect, often much longer than the leaves, often 20 cm. long or more, 3 mm.
thick, rather densely flowered; bracts orbicular, centrally peltate; fruits ovoid-
globose, verrucose, protracted at the apex into a somewhat curved, beak-like
appendage.

Among the Guatemalan collections that we refer here Trelease
indicated three or four unpublished species, but all the material
appears conspecific and we find no definite character to justify our
separating it from the common West Indian P. maculosa. The plants
when fresh have a strong and distinctive spicy odor, suggestive of
ginger. About Coban this plant is well known, and the leaves often
are peddled on the streets by the Indians. The very thick leaves
are bound on the head to relieve headache. The plant is one of the
most celebrated medicinal plants of the Coban region, and is used
in treating a variety of other ailments besides headache. The species
P. suavis Trelease, of which a variety is cited above in synonymy,
has not been published formally, although a Latin diagnosis of the
variety was published, as cited.

Peperomia major (Miq.) C. DC. in DC. Prodr. 16, pt. 1:
432. 1869. Acrocarpidium majus Miq. Syst. Piper. 60. 1843. P.
tremulaeformis Trelease in Yuncker, Field Mus. Bot. 9: 276. 1940
(type from Mount Cangrejal near La Ceiba, Honduras). Yepasetas
(Alta Verapaz).

On tree trunks or rocks in moist or wet, mixed, lowland or moun-
tain forest, 2,400 meters or lower; Alta Verapaz; Izabal; Quezal-
tenango; Huehuetenango. Southern Mexico; British Honduras;
Atlantic lowlands of Honduras; West Indies; South America.

258 FIELDIANA: BOTANY, VOLUME 24

Plants repent or subscandent, the stems slender, often much elongate, with
elongate internodes, rather densely pubescent with fine appressed hairs; leaves on
long slender petioles as much as 4 cm. long, broadly ovate or usually reniform-
ovate, commonly 3-4.5 cm. long and 3-4 cm. wide, pointed or subacute at the
apex, shallowly cordate at the base or sometimes only rounded, rather thick and
succulent when fresh, obscurely if at all pellucid-punctate, deep green above,
paler beneath, often yellowish green, usually 7-nerved, the nerves evident beneath,
sparsely or rather densely pubescent on both surfaces with short appressed hairs;
peduncles bracteate, axillary, solitary, equaling or only half as long as the spikes;
spikes short, densely flowered, the bracts small, rounded, centrally peltate, sub-
sessile; ovary subimmersed; fruits cylindric-ellipsoid, papillose below, spreading,
rather long-rostrate, the beak often equaling the body of the fruit.

This species has been reported from Guatemala as P. urocarpa
(Miq.) Fisch. & Mey. The Central American material is somewhat
variable but probably all of it is conspecific. Some of the collections
approach rather closely P. serpens Swartz, a plant widely distributed
in the West Indies and South America, with somewhat smaller leaves
than P. major.

Peperomia mexicana Miquel, Syst. Piper. 75. 1843.

Terrestrial in moist or wet, shaded places, often on cliffs, 300-
2,600 meters; Izabal; Zacapa; Santa Rosa; Quezaltenango. Southern
Mexico.

Plants glabrous, arising from a small globose hypogaean tuber, acaulescent
or with a very short, tuberous stem; leaves few, green, erect on long slender petioles,
the petioles mostly 2.5 cm. long or shorter; leaf blades very thin when dried, slightly
paler beneath, broadly ovate or rounded-ovate to oblong-ovate, mostly 2.5 cm.
long or shorter, acute, broadly rounded at the base, peltate slightly above the
base, obscurely 3-nerved; peduncles very slender, simple, usually 4-5 cm. long;
spikes very slender, pale green, mostly 4-5 cm. long; bracts ovate or ovate-lanceo-
late, acute, 1 mm. long; stamens with distinct filaments; fruits cylindric, 1 mm.
long, obscurely reticulate, short-stipitate, the style slender, cylindric, half as long
as the fruit, the stigma apical, capitellate.

Peperomia minarum Standl. & Steyerm., sp. nov.

Known only from the type, El Progreso, Sierra de las Minas,
along trail between Finca Piamonte and Cimiento, Steyermark 43747
(in Herbarium of Chicago Natural History Museum).

Plants epiphytic, the stems elongate and pendent, about 5 mm. thick, with
short or elongate internodes, sparsely pilose with long slender spreading whitish
hairs; leaves alternate, thick and fleshy when fresh, rather thick and blackish when
dried, finely punctate on both surfaces but not pellucid-punctate, the petioles
stout, about 1 cm. long, thinly pilose; leaf blades broadly ovate or oval-ovate,
mostly 5-7 cm. long and 3-4 cm. wide, rather abruptly acute, rounded at the base,
deep green on the upper surface and sparsely hirsute with long spreading hairs,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 259

conspicuously ciliate, somewhat paler beneath, glabrous or nearly so, 5-7-plinerved
or almost penninerved, the inner nerves arising far above the base of the blade,
visible only against a strong light; spikes unknown.

Planta epiphytica, caulibus elongatis pendulis crassis, internodiis brevibus vel
elongatis, sparse pilis longis paten tibus albidis pilosis; folia alterna crasse petiolata
in sicco fuscescentia non pellucido-punctata, petiolis pilosis; lamina late ovata
vel ovali-ovata subabrupte acuta, basi rotundata, ciliata, supra sparse pilis
patentibus hirsuta, subtus paullo pallidior glabra vel glabrata, 5-7-plinervia,
nervis obscuris; spicae ignotae.

Peperomia molithrix Trelease & Standley, sp. nov.

Epiphytic on tree trunks or growing among or on rocks in dense,
moist or rather dry forest, 200-1,700 meters; endemic; Zacapa (type
collected near Zacapa, Standley 74685; in Herbarium of Chicago
Natural History Museum); Chiquimula (near Conception de las
Minas) ; Jalapa (Chahuite, northwest of Jalapa) ; Santa Rosa.

Plants erect or ascending, 25 cm. high or less, the stems very thick and stout,
usually simple, naked below, leafy near the apex, with very short internodes,
sparsely pilose or glabrate; leaves apparently deciduous during the dry season,
very thin, alternate, translucent but not pellucid-punctate, the petioles very
slender, 7.5 cm. long or shorter, glabrous or short-pilose; leaf blades somewhat
paler beneath, ovate or broadly ovate, mostly 5-8 cm. long and 3.5-4.5 cm. wide,
acute, rounded or subcordate at the base, peltate a short distance above the base,
sparsely and inconspicuously short-pilose above, glabrous beneath or sparsely
short-pilose on the nerves, about 7-nerved, the nerves arising at the point of
attachment of the petiole; peduncles terminal, simple, naked, very slender, about
as long as the petioles, glabrous; spikes very slender, much exceeding the leaves,
14 cm. long or shorter, 1.5 mm. thick, densely flowered, the rachis glabrous or
sparsely and minutely puberulent; fruits emersed, subglobose or ovoid-globose,
about 0.7 mm. long, the stigma apical or nearly so.

Planta erecta vel adscendens, caule crasso vulgo simplici prope apicem foliato,
internodiis brevissimis sparse pilosulis vel glabratis; folia tenuia longissime graciliter
petiolata epunctata alterna, ovata vel late ovata, acuta, basi rotundata vel sub-
cordata, supra sparse breviter pilosula, subtus glabra vel ad nervos sparse breviter
pilosula, ca. 7-nervia; pedunculi terminales longi gracillimi nudi petiolis sub-
aequilongi, spicis gracilibus foliis multo longioribus 1.5 mm. crassis densifloris,
rhachi glabra vel minute puberula; fructus emersus subglobosus, apice stigmati-
ferus.

Of the four sheets of this plant seen by Trelease, each was in-
scribed as a new species, but all of them, with one later collection,
appear to us to represent a single species. The plant has been
reported from Guatemala as P. Muelleri C. DC.

Peperomia nigropunctata Miquel, Syst. Piper. 188. 1843.
Piper acuminatum L. Sp. PI. 30. 1753. Peperomia acuminata Dahlst.
Stud. 123. 1900, not Ruiz & Pavon, 1794. P. sanfelipensis C. DC.

260 FIELDIANA: BOTANY, VOLUME 24

in Bonn. Smith, Bot. Gaz. 19: 9. 1894 (type from San Felipe, Retal-
huleu, J. D. Smith 2583). Hierba de aguila (Suchitepequez, fide
Morales).

Moist or wet, mixed forest, chiefly in the mountains, on trees
or rocks, 1,800 meters or lower; Alta Verapaz; Izabal; Zacapa;
Escuintla; Guatemala; Suchitepequez; Retalhuleu; Quezaltenango;
San Marcos; Huehuetenango. Southern Mexico; British Honduras;
doubtless extending farther southward in Central America; West
Indies; South America.

Plants glabrous throughout, the stems branched or simple, short or elongate,
often pendent, generally rooting at the lower nodes, black-punctate, the internodes
short or elongate; leaves small, on petioles 1.5 cm. long or shorter, mostly lanceolate
or rhombic-lanceolate, generally 4-8 cm. long and 1-3 cm. wide, acute or acuminate,
acute or attenuate at the base, densely black-punctate on both surfaces, 5-pli-
nerved, the nerves usually obscure, the leaves mostly blackening when dried;
peduncles axillary and terminal, simple, naked, equaling or longer than the
petioles; spikes slender, black-punctate, sometimes 14 cm. long but usually shorter,
1 mm. thick, pale green; bracts rounded, centrally peltate, subsessile; ovary sub-
impressed in the rachis, ovoid, acutish, bearing below the apex the small puberulent
stigma; fruit ovoid-globose, smooth, obliquely rostellate at the apex.

This plant has been reported from Guatemala under the erroneous
name "Peperomia nigropunctula Miq."

Peperomia obtusifolia (L.) A. Dietr. Sp. PI. 1: 154. 1831.
Piper obtusifolium L. Sp. PL 30. 1753. P. magnoliaefolium Jacq.
Coll. Bot. 3: 210. 1789. Peperomia magnoliaefolia A. Dietr. op. cit.
153. P. Gottii Trelease ex Standl. Field Mus. Bot. 10: 156. pi. 31.
1931 (description based on Honduran material, but no type cited).
P. petenensis Trelease in Standl. Field Mus. Bot. 17: 230. 1937
(type collected near La Libertad, Pete"n, C. L. Lundell 2129). P.
lancetillana Trelease in Yuncker, Field Mus. Bot. 17: 334. 1938
(type from Lancetilla Valley near Tela, Honduras). P. lancetillana
var. spathifolia Trelease, op. cit. 335. P. Yousei Trelease in Yuncker,
Field Mus. Bot. 17: 339. 1938 (type collected at El Achote, above
Siguatepeque, Honduras). P. Yunckeri Trelease, op. cit. 340. 1938
(type collected near Siguatepeque, Honduras). P. romaensis Tre-
lease in Yuncker, Field Mus. Bot. 9: 275. 1940 (type from Roma
siding, east of La Ceiba, Honduras). Culantro de montana; Sakix-
acpec (Coban, Quecchi) ; Chunupcaac (Pete"n, Maya).

Moist or wet forest, usually epiphytic but sometimes on banks
or rocks, 2,600 meters or lower; Pete"n; Alta Verapaz; Izabal; Zacapa;
Chiquimula; Guatemala; Solola; Suchitepequez; Retalhuleu; Quezal-
tenango; San Marcos; Huehuetenango. Southern Mexico; British

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 261

Honduras to Honduras and Salvador, and probably southward
throughout Central America; West Indies; South America.

Plants stout, glabrous, usually erect, sometimes decumbent, the stems simple
or sparsely branched, with short or somewhat elongate internodes; leaves alternate,
very thick and fleshy when fresh, subcoriaceous when dry, not pellucid-punctate
or very inconspicuously so, usually not darkening when dried, on stout, short
or elongate petioles; leaf blades rounded-obovate to spatulate-obovate, mostly
5-12 cm. long and 3-5 cm. wide, variable in shape and size, broadly rounded to
very obtuse at the apex, sometimes emarginate, cuneate or attenuate at the base,
the costa emitting 3-5 inconspicuous nerves on each side; peduncles 2-5 cm. long
or longer, bearing 1-2 spikes or rarely more, terminal, solitary or clustered; spikes
densely flowered, much exceeding the leaves, mostly 6-14 cm. long and 3-4 mm.
thick; bracts suborbicular, centrally peltate, very shortly pedicellate; ovary im-
mersed; fruits ovoid-cylindric or ovoid-globose, more or less rostrate at the apex.

The material referred here is decidedly variable in foliage and
also in the size and disposition of the spikes, but none of the more
obvious characters by which species might be segregated seem to
be constant, and it seems better to place all the very numerous
Guatemalan collections in a single, not unusually variable species.
No fewer than 21 of the Guatemalan specimens have been marked
by Trelease as types of new species and varieties.

Peperomia pecuniifolia Trelease & Standley, sp. nov.

Epiphytic or on rather dry rocks, 1,500-2,800 meters; endemic;
Quezaltenango (type from Volcan de Santa Maria, above Santa
Maria de Jesus, Steyermark 34235, in Herbarium of Chicago Natural
History Museum); San Marcos (Volcan de Tajumulco, near Taju-
mulco).

Plants stout, the stems elongate, apparently pendent or procumbent, sparsely
branched, pubescent, at least around the nodes, the nodes elongate; leaves mostly
ternate or quaternate, or some of the lower ones opposite, on stout petioles 8 mm.
long or shorter, very thick and fleshy when fresh, coriaceous when dried and green-
ish or yellowish, opaque, orbicular to rounded-oval, mostly 2.5-4 cm. long and
1.5-4 cm. wide, broadly rounded at the apex, sometimes slightly emarginate,
rounded at the base, usually lustrous above, glabrous or nearly so, ciliate, 5-nerved;
spikes unknown.

Planta epiphytica vel clivicola, caulibus crassis elongatis ut videtur pendulis,
sparse ramosis, saltern ad nodos pubescentibus, internodiis elongatis; folia plerum-
que 3-4-nata, inferiora interdum opposita, crassissima, non pellucido-punctata,
crasse petiolata, petiolis pubescentibus, orbicularia vel rotundo-ovalia, apice late
rotundata, interdum breviter emarginata, basi rotundata, 5-nervia, glabra vel
glabrata, ciliata; spicae ignotae.

The relationship of this Guatemalan plant is clearly with P.
trifolia (L.) A. Dietr. of the West Indies. A different species may
not be represented, but the plant of the Antilles has consistently

262 FIELDIANA: BOTANY, VOLUME 24

smaller leaves than the continental one, and the latter probably
constitutes a distinct species.

Peperomia pellucida (L.) HBK. Nov. Gen. & Sp. 1: 64. 1815.
Piper pellucidum L. Sp. PI. 30. 1753. Choec (Huehuetenango) ;
Mazamorra (Pete"n, fide Lundell).

Moist or wet thickets or mixed forest, often a weed in waste
ground about dwellings, most common below 1,000 meters but
ascending to 2,500 meters or more; Pete*n; Alta Verapaz; Izabal;
Jutiapa; Santa Rosa; Escuintla; Suchitepequez; Retalhuleu; Quezal-
tenango; San Marcos; Huehuetenango. Mexico; British Honduras
to El Salvador and Panama; West Indies; South America; intro-
duced in tropical Africa.

Plants slender and low, mostly 25 cm. high or less, erect or ascending, some-
times procumbent, appearing glabrous throughout, the stems very succulent and
brittle, the plants annual or sometimes probably more enduring and with
decumbent and rooting bases, sparsely branched; leaves thin, on long slender
petioles, deltoid-ovate or rounded-ovate, mostly 1.5-3 cm. long, acute or obtuse,
shallowly cordate to broadly rounded at the base, sparsely pellucid-punctate or
epunctate, often pale green, especially beneath, glabrous or sometimes incon-
spicuously ciliate and with a few scattered hairs on the upper surface, palmately
5-nerved; flower spikes simple, pale green, very slender and usually remotely
flowered, mostly 1.5-5 cm. long, scarcely more than 1 mm. thick, slender-peduncu-
late; bracts minute, orbicular, peltate at the center, almost sessile; fruits sub-
globose, about 0.5 mm. in diameter.

Sometimes called "lombricilla" and "hierba de sapo" in El Sal-
vador. The plant is a common weed in many parts of the Central
American lowlands.

Peperomia peltata C. DC. Ann. Cons. Jard. Bot. Geneve 2:
277. 1898. P. pedicellata Dahlstedt, Svensk. Vet. Akad. Handl. 33,
no. 2: 35. 1900 (based on the same collection as P. peltata). Agria
(Chiquimula).

On moist or wet cliffs or rocks, 900-2,400 meters; endemic;
Chiquimula; Jalapa; Santa Rosa (type from Santa Rosa, Heyde &
IMX 3829); San Marcos.

Plants acaulescent, arising from a hypogaean tuber, glabrous; leaves few,
basal, erect, the petioles very slender, twice as long as the blades or often much
longer; leaf blades broadly ovate, mostly 4-8 cm. long, acute or short-acuminate,
broadly rounded or subcordate at the base, peltate a short distance above the base,
very thin, translucent, sparsely pellucid-punctate, usually 7-nerved; peduncles
slender, arising from the tuber, erect, usually longer than the leaves; spikes very
laxly flowered, with a rather thick, glabrous rachis; bracts small, ovate, pointed

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 263

at the apex, peltate; fruit oblong-cylindric, slightly verruculose, long-stipitate,
bearing at the apex a slender suboblique conic appendage.

The type collection was once listed from Guatemala as P. ovato-
peltata C. DC.

Peperomia peltilimba C. DC. ex Trelease, Bot. Gaz. 73: 145.
1922. P. hernandifolia var. cryptocarpa Trelease, Bot. Gaz. 73: 145.
1922 (type from Finca Sepacuite", Alta Verapaz, 0. F. Cook & R. F.
Griggs 533). Culantro (Chimaltenango) ; Culantro de monte (Quezal-
tenango) ; Cordoncillo.

Generally in dense wet mixed mountain forest, terrestrial or
epiphytic, often on rocks, 1,200-2,000 meters; Alta Verapaz; Chimal-
tenango; Huehuetenango; San Marcos. Costa Rica.

Plants very succulent, glabrous, the stems usually much elongate and with
elongate internodes, stout, repent and rooting at most of the lower nodes, the tips
of the branches erect or ascending; leaves alternate, the petioles slender, mostly
somewhat shorter than the blades; leaf blades thick and succulent when fresh but
when dried rather thin, blackish, opaque or nearly so, broadly ovate or rounded-
ovate, mostly 5-9.5 cm. long and 3.5-5.5 cm. wide, rather abruptly acuminate or
long-acuminate, rounded at the base, peltate near or somewhat above the base,
mostly 7-nerved, somewhat paler beneath; peduncles terminal, solitary or geminate,
equaling or shorter than the petioles, glabrous or puberulent, bearing a rather
conspicuous, foliaceous bract; spikes mostly 2.5-4 cm. long, erect, 2.5-3 mm.
thick, very densely flowered; ovary ovoid, obliquely short-rostrate, the stigma
anterior on the beak.

An earlier name may eventually be found for this plant among
some of the species of West Indies or other regions. We place here
specimens referred by Trelease to five unpublished Guatemalan
species.

Peperomia petrophila C. DC. Linnaea 37: 369. 1866.

Moist or wet, mixed, mountain forest, sometimes along streams
or in pine forest, 1,200-2,100 meters; Alta Verapaz; Zacapa (Sierra
de las Minas); Guatemala (Volcan de Pacaya). Veracruz.

Plants erect, 30 cm. high or less, the stems thick, with very short internodes,
naked below, densely leafy above, the lower leaves deciduous, the stems simple or
with a few short branches above, the plants glabrous throughout; leaves thin
and soft, on very slender petioles 1.5-3.5 cm. long, lanceolate or linear-lanceolate,
mostly 6-14 cm. long and 1-3 cm. wide, attenuate-acuminate, acute or attenuate
at the base, not pellucid-punctate, green above when dried, somewhat paler beneath
and black-puncticulate, 5-plinerved; peduncles terminal, solitary or clustered,
about equaling the petioles or often longer, slender, glabrous; spikes rather stout,
equaling or exceeding the leaves, 9.5 cm. long and 3 mm. thick or smaller, densely
flowered; bracts orbicular, centrally peltate, subsessile; ovary emersed, obovoid,
somewhat obliquely stigmatiferous at the apex, the stigma punctiform.

264 FIELDIANA: BOTANY, VOLUME 24

In Veracruz this plant grows on wet rocks in the margins of
streams. Probably it has the same habitat in Guatemala, although
the scant notes accompanying the three Guatemalan specimens are
not conclusive regarding this point.

Peperomia pililimba C. DC. Candollea 1: 335, 403. 1923.
P. Matapalo Trelease in Standl. Journ. Wash. Acad. Sci. 13: 366.
1923 (type from San Salvador, El Salvador).

Moist, wet, or rather dry forest, epiphytic or on rocks or rocky
banks, 200-3,000 meters; Alta Verapaz (type from Coban, Turck-
heim 11.1383); Baja Verapaz; Zacapa; Chiquimula; Santa Rosa;
Escuintla; Quezaltenango; Huehuetenango. Honduras; El Salvador.

Plants erect or sometimes decumbent and rooting at the lower nodes, the stems
stout, often clustered, simple below, often sparsely branched above, sparsely or
densely pilose with spreading or subappressed hairs, the internodes short or
elongate; leaves alternate, on stout or rather slender pilose petioles 1 cm. long or
shorter, very variable in shape, oblanceolate to cuneate-oblong or cuneate-obovate,
mostly 1.5-4 cm. long and 1-2 cm. wide, generally broadest toward the apex,
rounded or obtuse at the apex, acute to attenuate at the base, rather thick and
fleshy when fresh, finely pellucid-punctate, paler beneath and often tinged with
red or pink, sparsely pilose above, mostly along the costa, more densely pilose
beneath, obscurely 3-nerved, the nerves obscure or obsolete beneath; peduncles
terminal, solitary or geminate, naked, pilose or glabrate, 1.5 cm. long or shorter;
spikes erect, densely flowered, slender, 10 cm. long or shorter, about 2 mm. thick,
the rachis glabrous; bracts orbicular, centrally peltate, pale green; ovary sub-
globose, the stigma oblique.

In leaf and pubescence characters, the species of this alliance
are variable, and it is difficult to decide whether they should be
divided more finely or whether several of those here recognized should
be united.

Peperomia praetenuis Trelease in Standl. Field Mus. Bot. 12:
406. 1936.

Known only from the type, British Honduras, in a clearing,
Belize, C. L. Lundell 1929.

Plants very small, glabrous, 4 cm. high or less, erect, with fibrous roots or
with a very short rhizome, the internodes short; leaves alternate, pale green and
very thin when dried, pellucid-punctate, the petioles about 3 mm. long; leaf blades
broadly deltoid-ovate-cordate, 5-10 mm. long and wide, subacute, shallowly
cordate at the base, obscurely 3-nerved; peduncles little longer than the petioles,
terminal, simple, naked; spikes mostly 1-1.5 cm. long, 1 mm. thick, remotely
flowered, pale green; bracts rounded, centrally peltate, short-pedicellate; fruits
ellipsoid-globose, only slightly immersed, carnose, the stigma apical.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 265

This is the smallest of all the Guatemalan Piperaceae, the
individual plants so small that they probably have been overlooked
by other collectors. However, the species may be only a stunted
form of P. pellucida.

Peperomia praeteruentifolia Trelease in Yuncker, Field Mus.
Bot. 17: 337. 1938.

Moist or wet, mixed, mountain forest, 1,200-2,000 meters;
Chiquimula; Jalapa; Guatemala; San Marcos. Honduras, the type
from Siguatepeque.

Plants usually terrestrial, sometimes epiphytic, the stems erect or decumbent,
slender, mostly 30 cm. long or less, simple below, branched above, with mostly
elongate internodes, finely puberulent or almost wholly glabrous; leaves alternate,
the petioles slender, 3 cm. long or shorter, puberulent or glabrate; leaf blades thin
when dried, usually green, broadly rhombic-ovate to ovate or elliptic-ovate,
mostly 3-6 cm. long and 2-5 cm. wide, acute or subacute or sometimes obtuse,
obtuse or more often rounded at the base, glabrous or nearly so, paler beneath,
obscurely 5-7-plinerved, the inner nerves arising far above the base of the blade;
peduncles axillary or often terminal and clustered, usually much longer than the
petioles, puberulent or glabrate; spikes very slender, 3-9 cm. long, scarcely 1 mm.
thick, glabrous, very remotely flowered; bracts orbicular, centrally peltate.

Peperomia pseudopereskiifolia C. DC. in DC. Prodr. 16, pt. 1:
448. 1860.

Lowland forest; Alta Verapaz (between Sepacuite" and Secanquim,
H. Pittier, May 18, 1905). Cuba.

Plants glabrous, stout, erect or ascending, with elongate internodes; leaves
mostly opposite, the uppermost ternate, on rather long, stout petioles; leaf blades
lanceolate or elliptic-lanceolate, mostly 5.5-9.5 cm. long and 2-3 cm. wide, long-
acuminate, narrowly cuneate-attenuate to the base, thick and fleshy when fresh,
rather thick when dried, conspicuously 5-nerved or sometimes with 2 obscure
additional nerves; spikes axillary and terminal, about twice as long as the leaves,
densely flowered, the peduncles longer than the petioles; bracts rounded, centrally
peltate; flowers immersed, the ovary ovoid, contracted below the apex, the stigma
apical, puberulent, capitellate; fruits globose, mucronulate at the apex, pedicellate.

The single Guatemalan collection agrees so well with a photo-
graph of the Cuban type that we believe there is little doubt that
it should be referred here.

Peperomia puberula J. G. Baker in Saunders, Ref. Bot. 5:
pi. 302. 1871.

Described from plants cultivated in England, said to be from
Alta Verapaz.

266 FIELDIANA: BOTANY, VOLUME 24

Plants short-caulescent, somewhat succulent, velutinous-puberulent; leaves
alternate, broadly ovate, 4-7 cm. long, 3-6 cm. wide, subacute or abruptly acute,
peltate toward the rounded or subcordate base, the petioles about equaling the
blades; spikes terminal and axillary, 5-10 cm. long, long-pedunculate; fruits
obovoid, the stigmas subsessile, somewhat oblique.

We have seen no material referable to this species, which is not
included in the key. There, apparently, it would run to P. peltata,
for which species, indeed, puberula may be the oldest name.

Peperomia quadrifolia (L.) HBK. Nov. Gen. & Sp. 1: 69.
1815. Piper quadrifolium L. Sp. PI. ed. 2. 43. 1762. Peperomia
edulis Miquel, Linnaea 18: 711. 1844. P. santarosana C. DC. in
Bonn. Smith, Bot. Gaz. 19: 10. 1894 (type from Santa Rosa, Santa
Rosa, Heyde & Lux 3454). P. Luxii C. DC. in Bonn. Smith, Bot.
Gaz. 19: 260. 1894 (type from Ojo de Agua, Santa Rosa, Heyde &
Lux 3828). P. Standleyi Trelease in Standl. Journ. Wash. Acad.
Sci. 13: 366. 1923 (type from Tonacatepeque, Salvador). P. circuli-
folia Trelease in Yuncker, Field Mus. Bot. 17: 331. 1938 (type from
El Achote, near Siguatepeque, Honduras). P. conocarpa Trelease,
op. cit. 332 (type from El Achote, Honduras). Retono; Colchdn
de pena.

On trees in moist or wet forest, often on oaks, 3,000 meters or
lower, most common at middle elevations; Alta Verapaz; Baja
Verapaz; El Progreso; Izabal; Zacapa; Chiquimula; Jalapa; Santa
Rosa; Escuintla; Guatemala; Sacatepe"quez; Chimaltenango; Solola;
Quich^ ; Suchitepequez; Retalhuleu; Quezaltenango; San Marcos;
Huehuetenango. Mexico; El Salvador and Honduras to Panama;
West Indies; South America.

Plants glabrous, very succulent, mostly 20 cm. high or less, often forming
dense bushy clumps on the trunks or branches of trees, the stems sometimes
procumbent and rooting at the nodes, with short or elongate nodes; leaves pale
green, very thick and fleshy, quaternate, on short thick petioles, narrowly to very
broadly cuneate-obovate, mostly 7-13 mm. long but variable in size, broadly
rounded and usually emarginate at the apex, broadly or narrowly cuneate at the
base, obscurely 3-nerved, generally opaque; spikes terminal, solitary, on rather
long and slender peduncles, erect, densely or rather remotely flowered, mostly
1.5-4 cm. long, little more than 1 mm. thick; bracts orbicular, centrally peltate,
short-stipitate; ovary subimmersed, ovoid, attenuate, stigmatiferous at the tip;
fruits oblong-ovoid, emersed, mucronate at the apex. (Fig. 44.)

This is one of the most common of Guatemalan species, and is
represented by a large number of collections that show considerable
variation in shape and size of the leaves and in the length of the
spikes. It is believed, however, that all the not very different forms
may be referred satisfactorily to a single widespread species. The

FIG. 44. Peperomia quadrifolia. A. Habit of portion of flowering stem; X 1.
B. Flower with bract; X 12. C. Portion of inflorescence with flowers in position;
X 3. D. Pistil; X 30.

267

268 FIELDIANA: BOTANY, VOLUME 24

plant has been reported from Guatemala as P. Lindeniana Miquel,
which is probably still another synonym of P. quadrifolia, and as
P. tenerrima Miq. Some 24 sheets that we place here have been
marked by Trelease as new species or varieties. According to
Schiede, collector of the Mexican type of P. edulis, this plant was
eaten, either raw or cooked, by the inhabitants of the mountains of
central Mexico, and he stated that the plant had an aromatic odor
suggestive of coriander. So far as we know, Peperomias are not
eaten in Central America, although they might make a satisfactory
pot herb.

Peperomia reflexa (L. f.) A. Dietr. Sp. PI. 1: 180. 1831. Piper
reflexum L. f. Suppl. PI. 91. 1781. Peperomia opaca Trelease, var.
ciliata Trelease in Yuncker, Field Mus. Bot. 17: 336. 1938 (type
from El Achote, near Siguatepeque, Honduras).

Epiphytic on trunks or branches of trees in wet to rather dry
forest, often on oaks, 1,000-2,500 meters; Alta Verapaz; Baja Vera-
paz; Chiquimula; Jalapa; Guatemala; Sacatepe"quez; Chimaltenango;
Suchitepequez; Quezaltenango; San Marcos; Huehuetenango. Cen-
tral and southern Mexico; widely distributed in Central America;
West Indies; South America; also in the Old World tropics.

Plants perennial, very succulent, usually erect, often forming dense clumps
of numerous stems, generally less than 20 cm. high, the stems sometimes rooting
at the lower nodes, sparsely dichotomous or trichotomous, sulcate, glabrous or
hirtellous; leaves small, in whorls of 3-4, very shortly petiolate, when dry rigid
or somewhat coriaceous, usually lustrous, mostly glabrous, rhombic-elliptic or
broadly elliptic or rarely ovate, spreading or reflexed, obtuse or rounded at the
apex, acute or obtuse at the base, most often glabrous, impressed-punctate beneath,
obsoletely 3-nerved; petioles pubescent; spikes terminal, pale green, long-pedun-
culate, much longer than the leaves, mostly 2-4 cm. long but variable in length,
the peduncles glabrous or hirtellous, the rachis densely hirtellous; bracts orbicular,
centrally peltate, pedicellate; ovary ovoid, acute, the stigma apical, capitellate,
puberulent; fruits ovoid-cylindric, somewhat spreading or ascending.

A common plant in many parts of Guatemala. Around Coban
the crushed leaves of this and doubtless other species are applied
as poultices to relieve the cutaneous irritation produced by Rhus
striata. The specific name P. opaca Trelease has not been validly
published, in spite of the fact that a variety of it was published from
Honduras. Most of the Guatemalan specimens were determined by
Trelease as P. opaca.

Peperomia rotundifolia (L.) HBK. Nov. Gen. & Sp. 1: 65.
1815. Piper rotundifolium L. Sp. PI. 30. 1753. Peperomia Koepperi

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 269

Trelease in Yuncker, Field Mus. Bot. 9: 275. 1940 (type from
Mount Cangrejal near La Ceiba, Honduras). Cuartillo de palo.

On trees in moist or wet forest, 1,700 meters or lower, often
creeping among mosses; Pete*n; Alta Verapaz; Izabal; Chiquimula;
Retalhuleu; Quezaltenango; San Marcos. Southern Mexico; British
Honduras to Panama; West Indies; South America.

Stems very slender, with mostly elongate internodes, repent and rooting at
the nodes, puberulent or glabrate; leaves thin or when fresh thick and lenticular,
orbicular or oval-orbicular, 6-10 mm. long and almost or quite as wide, not pellucid-
punctate, sparsely blackish-puncticulate beneath, broadly rounded at base and
apex, sparsely or rather densely pilose or sometimes glabrate, ciliate, obscurely
3-nerved; spikes terminal, short-pedunculate, longer than the leaves, scarcely
1 mm. thick, the rachis glabrous, the peduncles about as long as the petioles;
bracts orbicular, centrally peltate, subsessile; ovary stigmatiferous slightly below
the apex, obtuse, the stigma minute, glabrous; fruit subglobose.

This has been reported from Alta Verapaz as P. matlaleucaensis
C. DC.

Peperomia Sandersii C. DC., perhaps native of Brazil, is grown
occasionally as a pot plant in Guatemala and the United States.
It is small, with very thick and fleshy, peltate, ovate-rounded leaves,
handsomely striped on the upper surface with green and silver.
Aguilar reports the local name of "cebranita."

Peperomia santa-helenae Trelease in Standl. Field Mus. Bot.
22: 9. 1940.

Moist or wet, mixed, mountain forest, sometimes in Cupressus
forest, epiphytic or rarely terrestrial, rarely in lowland forest little
above sea level; 1,500-3,200 meters; endemic; Alta Verapaz(?);
Izabal; El Progreso; Zacapa; Jalapa; Guatemala; Chimaltenango
(type from Santa Elena, Cerro de Tecpam, J. R. Johnston 421);
Quiche* ; Huehuetenango; Quezaltenango; San Marcos.

Plants erect or decumbent, the stems generally simple below but often or
usually somewhat branched above, stout, rather densely pilose with short, mostly
fulvescent and incurved hairs, the internodes short or elongate; leaves alternate,
rather thick and succulent when fresh, somewhat paler beneath, green or fuscescent
when dried, not pellucid-punctate, generally black-puncticulate beneath, the
petioles 1.5 cm. long or usually much shorter, incurved-pubescent; leaf blades
variable in size and shape, mostly elliptic or lance-elliptic, sometimes rhombic-
ovate to broadly ovate or obovate, mostly 2-5 cm. long and 1.5-3.5 cm. wide, acute
or acuminate with an obtuse tip, rounded to subacute at the base, 3-nerved, pilose
on both surfaces with spreading or subappressed hairs or in age rarely glabrate;
peduncles axillary and terminal, solitary or the terminal ones clustered, simple,
naked, 1.5 cm. long or shorter, sparsely pilose; spikes pale green, slender, mostly

270 FIELDIANA: BOTANY, VOLUME 24

5-10 cm. long and 1.5 mm. thick, glabrous, remotely flowered; bracts centrally
peltate, orbicular, sessile; fruits minute, ovoid-globose, glabrous, the stigma sub-
obliquely apical.

Peperomia sisiana C. DC. in Donn. Smith, Bot. Gaz. 19: 9.
1894.

On trees in lowland forest, 700 meters or lower; endemic; Suchite-
pequez (type from Rio Sis, J. D. Smith 2584; collected also at San
Antonio).

Plants with elongate, probably pendent stems, the stems sometimes rooting
at the lower nodes, sparsely hirsute or glabrate, the internodes elongate; leaves
alternate, on stout petioles 1 cm. long or shorter, oblong-lanceolate, mostly 7-9 cm.
long, acuminate or narrowly long-acuminate, obtuse at the base, glabrous, con-
spicuously ciliate, penninerved or 5-7-plinerved with the inner nerves arising far
above the base of the blade and little below the middle of it, usually fuscescent
when dried, not or obscurely pellucid-punctate; peduncles terminal, simple, naked;
spikes slender, glabrous, densely flowered, about 6 cm. long, in fruit 6 mm. thick;
bracts orbicular, centrally peltate, subsessile; ovary emersed, obliquely rostrate;

fruits cylindric, widely ascending, as much as 2.5 mm. long, somewhat tuberculate.

Peperomia Skutchii Trelease & Standley, sp. nov.

Terrestrial in moist or wet, mixed, mountain forest, 1,500-2,900
meters; endemic; Alta Verapaz; Chimaltenango; Quezaltenango
(type from western slopes of Volcan de Zunil, above Santa Maria
de Jesus, Steyermark 35144); San Marcos; Huehuetenango.

Plants very slender, weak, succulent, and fragile, erect or procumbent, glabrous
almost throughout, the stems simple or often abundantly branched, mostly less
than 20 cm. long, sometimes hispidulous at the nodes, the internodes short or
elongate; leaves very small, alternate but often opposite at the ends of the branches,
translucent when dried but not pellucid-punctate, the slender petioles mostly
3-7 mm. long, glabrous or hispidulous near the apex; leaf blades orbicular, ovate-
rounded, or rounded-rhombic, mostly 8-23 mm. long and 8-16 mm. wide, rounded
or broadly rounded at the apex, rounded at the base, glabrous throughout or
often with a few scattered short white hairs on the upper surface, sometimes
hispidulous beneath near the base, 5-7-plinerved or penninerved, the nerves very
inconspicuous; peduncles filiform, axillary, solitary, simple, naked, glabrous,
generally longer than the petioles; spikes filiform, mostly about 1 cm. long, pale
green, remotely few-flowered, less than 1 mm. thick, glabrous; fruits minute,
stipitate, emersed, obliquely ascending, fusiform, rather long-mucronate, the
stigma broader than the style.

Planta gracilis vulgo ramosa erecta vel procumbente carnosa fere glabra,
caulibus interdum ad nodes hispidulis; folia parva longipetiolata alterna sed ad
apices ramorum saepe opposita, orbicularia, ovato-orbicularia vel rotundo-rhombea,
apice rotundata vel late rotundata, basi rotundata, glabra vel supra pilis paucis
brevibus albis conspersa, subtus interdum prope basin hispidula, 5-7-plinervia

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 271

vel penninervia; pedunculi filiformes axillares singuli plerumque petiolis longiores,
spicis filiformibus vulgo ca. 1 cm. longis laxe paucifloris glabris; fructus stipitatus
fusiformis emersus oblique adscendens longimucronatus.

A small and delicate plant of the high wet mountain forests,
collected numerous times in the Occidente.

Peperomia staminea Trelease, Journ. Wash. Acad. Sci. 19:
328. 1939.

Epiphytic on trees in wet mixed lowland forest, 500 meters or
lower; Izabal. British Honduras; Atlantic coast of Honduras (type
from Lancetilla Valley near Tela).

Plants rather small, glabrous almost throughout but often puberulent on the
petioles and peduncles and at the nodes, the stems slender, elongate, usually
repent and rooting at the nodes, the internodes elongate; leaves mostly quaternate,
on petioles 2-4 mm. long, very thick and succulent when fresh, subcoriaceous when
dry, usually not at all pellucid-punctate, elliptic to oblong-lanceolate, mostly
2-2.5 cm. long and 1-1.5 cm. wide, somewhat narrowed to the very obtuse apex,
acute or obtuse at the base, 3-nerved, the nerves conspicuous beneath but obsolete
above; spikes terminal, on peduncles as much as 3.5 cm. long, densely flowered,
about 3 cm. long and 2 mm. thick, pale green; bracts suborbicular, peltate at the
middle.

Peperomia suchitanensis Trelease & Standley, sp. nov.

Known only from the type, Jutiapa, Volcan de Suchitan, on
wet rocks among mosses and on bark of trees in forest at the summit,
2,050 meters, Steyermark 31940 (type in Herbarium of Chicago
Natural History Museum).

Plants very slender, the stems repent or procumbent, rooting at the lower
nodes, short, simple or sparsely branched, with short internodes, sparsely pilose
with slender spreading hairs; leaves alternate, numerous, the petioles about 1 mm.
long; leaf blades rather thick, elliptic-ovate, 6-11 mm. long, 3-8 mm. wide,
narrowed to the obtuse, often subemarginate, minutely apiculate apex, rounded
at the base, green when dried, not pellucid-punctate, very obscurely 3-nerved,
conspicuously hispidulous on the upper surface with rather long, spreading hairs,
somewhat paler beneath and glabrous; peduncles terminal, solitary, simple, slender,
glabrous, longer than the petioles; spikes (very immature) rather stout, about
6 mm. long, scarcely 1 mm. thick, very densely flowered, glabrous.

Planta parva gracillima, caulibus repentibus vel procumbentibus brevibus,
sparse pilis patentibus pilosis, internodiis brevibus; folia parva alterna brevissime
petiolata elliptico-ovata, 6-11 mm. tantum longa, apicem obtusum saepe sub-
emarginatum minute apiculatum versus paullo angustata, basi rotundata, obscure
3-nervia, supra hispidula, subtus glabra; pedunculi terminales singuli glabri
petiolis longiores, spicis immaturis erectis crassiusculis erectis glabris ca. 6 mm.
longis.

272 FIELDIANA: BOTANY, VOLUME 24

Peperomia tacanana Trelease & Standley, sp. nov.

Epiphytic or terrestrial in moist or wet, mountain forest, some-
times growing on rocks, 1,500-2,700 meters; Zacapa; El Progreso;
Chiquimula; Jalapa; Guatemala; Chimaltenango; Quezaltenango;
San Marcos (type from Volcan de Tacana, along Rio Vega, between
San Rafael and the Mexican boundary, Steyermark 36351; in Her-
barium of Chicago Natural History Museum). El Salvador.

Plants usually very slender, the stems erect or often long-repent, frequently
30 cm. long or more, simple or sparsely branched, densely and finely puberulent,
the internodes mostly elongate; leaves small, alternate, long-petiolate, succulent
when fresh but rather thin when dried, the petioles 2 cm. long, slender, often
longer than the blades, puberulent; leaf blades broadly ovate or rounded-ovate,
acute or subacute or sometimes obtuse, rounded or truncate at the base, mostly
12-25 mm. long and 10-27 mm. wide, finely and usually sparsely puberulent on
both surfaces or sometimes glabrate, paler beneath, sparsely pellucid-punctate,
obscurely 3-nerved; peduncles very slender, terminal, solitary or often ternate,
naked, puberulent, simple, 1.5 cm. long or shorter; spikes pale green, erect, slender,
commonly 2-4 cm. long and 1-1.5 mm. thick, remotely flowered, glabrous; bracts
orbicular, centrally peltate; fruits ovoid-globose, minute, emersed, glabrous, sub-
apiculate at the apex, the stigma slightly oblique.

Caules graciles erecti vel saepius repentes et ad nodos radicantes, simplices
vel sparse ramosi, dense minute puberuli; folia parva longipetiolata late ovata
vel rotundo-ovata, acuta vel subacuta vel interdum obtusa, basi rotundata vel
truncata, utrinque sparse puberula vel glabrata, subtus pallidiora, sparse pellucido-
punctata, obscure 3-nervia; pedunculi terminales saepe ternati nudi puberuli
simplices, spicis gracilibus erectis 2-4 cm. longis 1-1.5 mm. crassis glabris; bracteae
rotundatae peltatae; fructus emersus ovoideo-globosus suboblique apiculatus.

Peperomia tacticana Trelease & Standley, sp. nov.

Terrestrial or epiphytic in moist or wet, mixed, mountain forest,
1,500-2,300 meters; endemic; Alta Verapaz (type from mountains
east of Tactic, on the road to Tamahu, Standley 71389; in Herbarium
oLChicago Natural History Museum) ; Zacapa (Sierra de las Minas) ;
San Marcos; Huehuetenango (Cerro Negro, Sierra de los Cuchu-
matanes).

Plants erect or ascending, glabrous throughout or nearly so, the stems slender,
mostly 40 cm. long or less, simple or sparsely branched, with elongate internodes;
leaves mostly in verticils of 3-4, or often partly opposite, on slender petioles
1 cm. long or shorter, somewhat fuscescent when dried and thin, finely pellucid-
punctate, oblong-lanceolate or oblong-ovate, mostly 6-8.5 cm. long and 2-3 cm.
wide, rather abruptly long-acuminate, acute to subobtuse at the base, 3-nerved or
sometimes 5-nerved; peduncles terminal, solitary, simple, about 1.5 cm. long;
spikes erect, slender, about 6.5 cm. long and 2 mm. thick, densely flowered, the
bracts suborbicular, sessile, centrally peltate; fruits mostly emersed, broadly
ovoid, minute, sessile, somewhat obliquely apiculate at the apex, glabrous.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 273

Caules erecti vel adscendentes graciles, simplices vel sparse ramosi, plantis
glabris; folia plerumque 3-4-nata, interdum pro parte opposita, graciliter petiolata,
in sicco tenuia fuscescentia, pellucido-punctata, oblongo-lanceolata vel oblongo-
ovata, subabrupte longiacuminata, basi acuta vel subobtusa, 3-nervia vel interdum
5-nervia; pedunculi terminales solitarii petiolis longiores simplices nudi; spicae
erectae graciles 2 mm. crassae densi florae; bracteae suborbiculares sessiles centre
peltatae; fructus emersus late ovoideus sessilis suboblique apice apiculatus.

Peperomia Treleasei Standl. & Steyerm., sp. nov.

Type from Guatemala, cultivated in the garden of Don Mariano
Pachecho H., Guatemala, May 14, 1942, Steyermark 46398 (in Her-
barium of Chicago Natural History Museum). Represented also
by Sesse & Mocino 2, 320, and 330, presumably from Mexico (Herb.
Madrid).

Plants slender, erect, the stems simple, 35 cm. high or less, very finely and
densely puberulent, the internodes few, often greatly elongate and as much as
14 cm. long; leaves in verticils of 3-5, thin when dried and green, not pellucid-
punctate, the petioles slender, 3 cm. long or shorter, sometimes longer than the
blades; leaf blades broadly cordate or rounded-cordate, 2.5-3.5 cm. long, about
2.5 cm. wide, subacute to very obtuse at the apex, shallowly cordate at the base,
mostly 7-nerved, densely and minutely puberulent on both surfaces or sometimes
glabrate; peduncles terminal, usually ternate, the central one much longer than
the lateral ones and as much as 7.5 cm. long, minutely puberulent, naked; spikes
very numerous, pedunculate, laxly or densely flowered, the lower ones as much
as 1 cm. long (including the peduncles), forming a dense raceme as much as 4 cm.
long, the upper spikes of the raceme gradually shorter, the racemes thus attenuate-
acute, the secondary peduncles bracteate at the base.

Caulis gracilis erectus simplex dense minuteque puberulus, internodiis elongatis
interdum longissimis; folia verticillata 3-5-nata longe graciliter petiolata tenuia
non pellucido-punctata, late cordata vel rotundo-cordata apice subacuta usque
obtusissima, basi breviter cordata, plerumque 7-nervia, utrinque dense minute
puberula vel interdum glabrata; pedunculi terminales vulgo ternati, centrali
elongate nudo, minute puberuli; spicae numerosae racemose dispositae 1 cm. longae
vel breviores basi bracteatae laxiflorae vel densiflorae patentes.

This is related to P. Fraseri C. DC., which was described from
Ecuador. Casimir De Candolle referred to it also a collection from
"Nova Hispania," collected by Pavon, which is doubtless the
plant that was collected by Sesse* and Mocino, as cited above. The
Ecuador and Mexican plants are evidently closely related, and form
a very distinct group, with inflorescences quite unlike those of other
species of the genus. Two of the Madrid collections were seen by
Trelease, who labeled them P. Fraseri. We have seen a photograph
of the type of that species, and believe that it is quite distinct from
the plant of Guatemala and Mexico. It has leaves that were
described by De Candolle as cordate-lanceolate, and they are

274 FIELDIANA: BOTANY, VOLUME 24

gradually attenuate to a long narrow apex; the leaves of P. Treleasei \
are rather abruptly narrowed to a much broader apex, and decidedly
.different in appearance. All the four collections of P. Treleasei are
uniform in leaf characters, and we have no doubt that they represent
a form quite distinct from the plant of Ecuador. It still is uncertain
where P. Treleasei is native. The plant cultivated in Guatemala
was said to have come from the Department of Jutiapa. It has not
been found wild anywhere in the country, and may well have been
in cultivation in Jutiapa. The Sesse* and Mocino collections, as
usual, bear no locality data.

Peperomia Tuerckheimii C. DC. ex Donn. Smith, Enum. PI.
Guat. 2: 96. 1891, nomen; Ann. Conserv. Jard. Bot. Geneve 2: 279.
1898.

On shaded wet cliffs or rocks, usually in dense forest, 900-1,500
meters; endemic; Alta Verapaz (type from Pansamala, Turckheim
433; also in the region of Chirriacte") ; Huehuetenango (Maxbal).

Plants very small, the stems short, slender, prostrate, rooting at the nodes,
pilosulous; leaves small, thin, the slender petioles as much as 4 cm. long, mostly
shorter; leaf blades thin, rounded-ovate, 2.5 cm. long and 2 cm. wide or smaller,
subacute, rounded at the base, peltate shortly above the base, translucent but
not punctate, pilosulous on both surfaces, ciliate; peduncles axillary, solitary,
simple, about equaling the petioles; spikes very slender, twice as long as the
leaves, the rachis puberulent; bracts orbicular; ovary emersed, ovoid, stigmati-
ferous at the very apex, acute.

The plant seems to be a rare one. It probably is confined to
limestone regions.

Peperomia vegana Trelease & Standley, sp. nov.

Moist or wet, mixed, mountain forest, on shaded banks or some-
times on rocks, 2,500-2,700 meters; endemic; San Marcos (type
from Volcan de Tacana, along Rio Vega, between San Rafael and
the Mexican boundary, Steyermark 36350 (type in Herbarium of
Chicago Natural History Museum; collected also in Barranco
Eminencia, near the base of Volcan de Tajumulco).

Plants slender, the stems elongate, prostrate or repent, rooting at the nodes,
the internodes elongate, sparsely pilose or glabrate, often conspicuously granular;
leaves rather small, alternate, the slender petioles 4.5 cm. long or shorter, pilose
with long spreading hairs; leaf blades thin when dried, fuscescent, not pellucid-
punctate, ovate or broadly ovate, 4.5-7.5 cm. long, 2.5-4.5 cm. wide, rather
abruptly acute or acuminate, broadly rounded at the base, peltate far above
the base, villous-hirsute on both surfaces with long slender spreading multicellular
hairs, about 7-nerved, paler beneath; spikes unknown.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 275

Planta gracilis, caulibus elongatis prostratis vel repentibus ad nodos radi-
cantibus, internodiis elongatis, sparse pilosis vel glabratis, saepe conspicue
granulosis; folia inter minora alterna longipetiolata, in sicco tenuia, non pel-
lucido-punctata, ovata vel late ovata, subabrupte acuta vel acuminata, basi late
rotundata, bene supra basin laminae peltata, utrinque dense pilis longis paten-
tibus laxis plurilocularibus villoso-hirsuta; spicae ignotae.

PIPER L.

Shrubs or small trees, generally terrestrial, sometimes scandent (very rarely
in American species), sometimes herbaceous almost throughout but always woody
about the base, the branches generally nodose; leaves alternate, often very unequal
at the base, palmately nerved or penninerved; stipules sometimes adnate to the
petiole and wing-like, or the two united into one opposite the petiole, sometimes
almost obsolete; flowers perfect or unisexual, adnate to the subtending peltate
bracts, solitary at each bract, sessile or rarely stipitate; spikes generally peduncu-
late, at. first terminal, becoming opposite the leaves, rarely collected on a common
peduncle; stamens 2-4 or rarely more numerous, the filaments short or rarely
exceeding the bracts, the anthers ovate or shorter, usually opening by longitudinal
slits; ovary sessile or nearly so, obtuse or rostrate, the stigmas 2-5, usually 3-4,
distinct, erect or recurved ; ovule erect from the base of the cell ; fruit baccate, very
small, ovoid or globose, smooth, sometimes partly immersed, sometimes sessile
and exserted or rarely long-stipitate, the pericarp very juicy or with scant, only
slightly succulent flesh; testa of the seed thin, the endosperm farinose.

Because of the different treatments of the genus by different
authors, and because no recent monograph of the genus as a whole
has been attempted, or is likely to be undertaken very soon, the
number of species is very uncertain, but the genus is one of the
largest of the plant kingdom.

Flower spikes fasciculate at the apex of a distinct peduncle, or rarely fasciculate
in the leaf axils. Leaves large and thin, ovate-rounded or orbicular, peltate
or epeltate. Subgenus Heckeria.

Leaves not peltate P. umbellatum.

Leaves conspicuously peltate.

Leaves sparsely villous on the upper surface, densely villosulous beneath on

the nerves and veins P. Heydei.

Leaves glabrous or nearly so P. peltatum.

Flower spikes solitary, axillary or opposite the leaves.
Leaves conspicuously peltate.
Branches and lower leaf surface densely pubescent; spikes mostly 3-5 cm.

long P. Tuerckheimii.

Branches and lower leaf surface glabrous or nearly so; spikes 7-12 cm. long
or longer.

Leaves lance-oblong, 5-6.5 cm. wide P. imberbe.

Leaves broadly ovate, mostly 10-20 cm. wide P. grandilimbum.

Leaves not peltate.

Leaves palmate-nerved, the 3 (sometimes more) principal nerves extending
nearly or quite to the apex of the leaf blade.

Flowers conspicuously pedicellate. Subgenus Arctottonia.

276 FIELDIANA: BOTANY, VOLUME 24

Leaf blades conspicuously cordate at the base, pilose beneath on the

nerves P. guazacapanense.

Leaf blades acute to rounded at the base, glabrous or minutely puberulent.
Leaves, at least the upper ones, acute or subacute at the base, thick

and stiff, lustrous on the upper surface P. sempervirens.

Leaves obliquely rounded or very obtuse at the base, membranaceous,

scarcely lustrous P. yucatanense.

Flowers sessile.

Leaf blades acute to rounded at the base, the base usually symmetric

or nearly so, never very conspicuously oblique.
Leaves abundantly pilosulous or puberulent beneath, at least on the

nerves.

Leaves pilose or pilosulous beneath with rather long, spreading hairs.

P. Martensianum.
Leaves finely and densely puberulent beneath on the nerves.

P. vaccinum.
Leaves glabrous or at most granular beneath on the nerves, rarely

very minutely and sparsely puberulent.
Principal leaves oblong or lance-oblong, mostly 9-15 cm. long and

2.5-4.5 cm. wide, little if at all narrowed toward the base.
Leaves densely and prominently reticulate- veined beneath.

P. Schippianum.

Leaves very laxly and inconspicuously reticulate-veined beneath.

P. vergelense.

Principal leaves ovate or broadly ovate to elliptic or oval-elliptic,

if rather narrow then conspicuously narrowed toward the base.

Bracts and rachis of the inflorescence densely or conspicuously

pubescent.

Peduncles several times as long as the petioles .... P. jumayense.
Peduncles little if at all longer than the petioles, sometimes

shorter P. Amalago.

Bracts and rachis of the inflorescence glabrous or essentially so.
Peduncles almost half as long as the spikes; leaves with a very

long and narrow, tail-like acumination P. stillans.

Peduncles short, usually much less than one-fourth as long as
the spikes; leaves mostly acuminate but not with a long
narrow tail-like tip.

Spikes all or mostly 3-5 cm. long P. psilorhachis.

Spikes 7-15 cm. long (except the undeveloped ones).
Peduncles filiform, mostly 2.5-5 cm. long.

P. perlongipedunculum.
Peduncles stouter, scarcely filiform, rarely as much as 1 cm.

long P. Amalago.

Leaf blades all or mostly cordate or subcordate at the base, at least on
one side, often very unequal, one side produced much lower on the
petiole than the other side.

Leaf blades unequal at the base, one side produced lower on the
petiole than the other; leaves lance-oblong to ovate-oblong, all
or most of them 2-5 cm. wide.

Leaves glabrous beneath P. tajumulcoanum.

Leaves more or less pubescent beneath, at least on the nerves, some-
times only minutely puberulent.

Leaves merely puberulent beneath on the nerves, the hairs minute
and scarcely with any appreciable length . . P. oblique-ovatum.
Leaves pilosulous beneath along the nerves, the hairs of very
appreciable length.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 277

Peduncles 2-3 cm. long P. santae-clarae.

Peduncles mostly 1-1.5 cm. long P. Standleyi.

Leaf blades equal and symmetric at the base or nearly so or, if some-
what unequal, the blade produced on each side to the same point
on the petiole.

Leaves abundantly pilose or pilosulous beneath, at least on the
nerves.

Leaves hirsute on the upper surface P. cristinanum.

Leaves not hirsute on the upper surface.

Leaf blades lance-oblong, about 3 times as long as wide.

P. tacananum.
Leaf blades ovate to rounded-ovate, mostly twice as long as

wide or shorter.

Leaves all rather deeply cordate at the base; plants herba-
ceous throughout or nearly so, usually low, weak, and
straggling, or sometimes more or less scandent.

P. relalhuleuense.

Leaves not all cordate at the base, the upper ones, and some-
times also the lower ones, rounded; plants erect shrubs.

P. Martensianum.

Leaves glabrous beneath, or merely puberulent on the nerves, the
hairs very short and scarcely with appreciable length.

Some nerves arising from sides of midrib P. variabile.

All nerves arising palmately from base of blade.

Leaves thin, usually blackish when dry, densely and finely pellu-
cid-punctate P. patulum.

Leaves thick and firm, not blackening when dried, epunctate or

nearly so P. diandrum.

Leaves penninerved, the lateral nerves not extending to the apex of the

blade, the lower ones usually terminating below the middle of the blade.

Leaves glabrous on both surfaces, at most minutely granular beneath on

the costa.
Leaf blades acute to attenuate at the base, 1-2 cm. wide, the nerves

arising at an extremely narrow angle P. flavidum.

Leaf blades cordate to obtuse at the base or, if acute, much more than

2 cm. wide.

Spikes at anthesis 1-2.5 cm. long. Leaves mostly 3.5-4.5 cm. wide.
Leaf blades unequal and shallowly cordate at the base.

P. ixocubvainense.
Leaf blades acute or subacute at the base P. telanum.

Spikes at anthesis more than 2.5 cm. long, mostly 5-10 cm. long or
longer. Leaves often very large and broad.

Leaves large, mostly 9-18 cm. wide, acuminate or abruptly short-
acute.

Petiole vaginate only at base; leaf blades broadest at base.

P. variabile.
Petiole vaginate throughout; leaf blades broadest at or about

middle P. yzabalanum.

Leaves mostly 7 cm. wide or much narrower, if rather wide the
blades narrowly long-acuminate.

Principal leaves cordate or subcordate at the base, at least on

one side.
Leaf blades subequal at the base, the two sides of about equal

length P. inslabilipes.

Leaf blades very oblique or unequal at the base.

278 FIELDIANA: BOTANY, VOLUME 24

Leaves mostly obtuse or rounded at the apex. .P. tuber culatum.
Leaves acuminate at apex.

Leaves mostly 8-11 cm. long, short-acuminate . P. pinetorum.
Leaves mostly 12-20 cm. long, narrowly long-acuminate.

P. pergamentifolium.

Principal leaves acute to rounded at the base, not at all cordate.
Primary nerves more or less impressed on the upper surface,
often deeply so, strongly elevated beneath.

P. xanthostachyum.
Primary nerves not impressed on the upper surface, not very

prominent beneath.
Leaves usually with 3 pairs of principal nerves.

Leaf blades lance-oblong, narrowly long-acuminate, not

blackening when dried P. curvatipes.

Leaf blades ovate or elliptic-ovate, rather abruptly short-
acuminate, blackening when dried. . . .P. brevilimbum.
Leaves all or mostly with 5-6 pairs of conspicuous lateral

nerves.

Leaf blades very unequal at the base, one of the sides
decurrent on the petiole and almost 1 cm. lower than

the other P. geniculatum.

Leaf blades not very unequal at the base or, if so, one side

decurrent but little below the other.
Leaves conspicuously paler beneath, glaucescent.

P. subeburneum.

Leaves not or scarcely paler beneath, not glaucescent.
Spikes short, in fruit as much as 5.5 cm. long, at

anthesis usually shorter.
Leaves lanceolate or ovate-lanceolate, broadest near

the base P. pergamentifolium.

Leaves oblong-elliptic, broadest at or near the

middle P. ixocubvainense.

Spikes elongate, mostly 6-9 cm. long in anthesis.

Leaf blades acute at the base P. coronanum.

Leaf blades rounded or very obtuse at the base.
Leaves rather small, mostly 3-5.5 cm. wide.

P. Donnell-Smithii.
Leaves large, mostly 6-9 cm. wide.

Leaf blades mostly 12-15 cm. long. .P.frioense.
Leaf blades large, mostly 17-22 cm. long.

Petioles 2-3.5 cm. long P. cheyennense.

Petioles 8-15 mm. long P. virginicum.

Leaves evidently pubescent on one or both surfaces, or at least puberulent

or pilosulous beneath on the nerves.

Leaves glabrous on the upper surface or nearly so, smooth to the touch.
Leaf blades conspicuously cordate at the base, at least on one side.
Nerves, and often veins, conspicuously impressed on the upper sur-
face, very prominent beneath, the blades often strongly bullate.

Leaves not bullate, glabrous beneath or sometimes sparsely hirsute
on the nerves toward the base of the blade . P. xanthostachyum.

Leaves conspicuously bullate, densely pubescent beneath on the

veins with short, mostly appressed hairs.

Leaf blades lanceolate or oblong-lanceolate, mostly 11-22 cm.
long, long-attenuate P. Biritak.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 279

Leaf blades oblong-elliptic, 7-10 cm. long, acute or short-
acuminate P. zacapanum.

Nerves and veins not impressed on the upper surface, not strongly

elevated beneath, the blades not at all bullate.
Leaves large, mostly 13-25 cm. wide, shallowly or deeply cordate
at the base, the basal lobes subequal, or sometimes unequal,
but one of the lobes then 5-8 cm. long or larger.
Base of the leaf blade deeply cordate, the sinus 3-5 cm. deep
or more; leaves pubescent beneath over the whole surface.

P. pansamalanum.

Base of the leaf blade shallowly cordate, the sinus usually less
than 2 cm. deep, often very broad and open; leaves pubes-
cent beneath chiefly or wholly on the nerves.

Leaves densely black-punctate beneath P. punctulatum.

Leaves inconspicuously pale-punctate beneath or epunctate.
Leaf blades shallowly and very narrowly cordate at the

base, thin P. exactum.

Leaf blades very broadly and openly cordate at the base,

thick P. philodendroides.

Leaves smaller, mostly 4-7 cm. wide, sometimes as much as 9.5
cm., the base very unequal, one side usually acute or obtuse,
the other cordate, with a small lobe usually much less than
1 cm. long.
Leaf blades dull on the upper surface, not lustrous.

Pubescence of the veins on the lower leaf surface of rather

long, spreading, whitish hairs P. patzulinum.

Pubescence of the veins of the lower surface of very short,

sordid, appressed or subappressed hairs.
Leaves oblong-lanceolate, 2.5-4.5 cm. wide ... P. brujoense.

Leaves elliptic, 6-7 cm. wide P. planadosense.

Leaf blades very lustrous on the upper surface.

P. aeruginosibaccum.

Leaf blades not at all cordate at the base, acute to obtuse or rounded.
Young branchlets densely pubescent or short-pilose.

Leaves very lustrous on the upper surface ... P. aeruginosibaccum.
Leaves dull on the upper surface, at least when dried.

P. uspantanense.
Young branchlets glabrous or sparsely and very minutely puberulent

or granular.

Leaves conspicuously lustrous on the upper surface when dried.

P. subcitrifolium.
Leaves dull on the upper surface when dried.

Leaves strigillose or appressed-pubescent beneath on the costa.

Leaf blades broadest somewhat above the middle, somewhat

rhombic-obovate, bright green when dried P. misantlense.

Leaf blades broadest below or near the middle, lance-oblong

or lance-ovate, darkening when dried.

Leaf blades acute at the base P. Chamissonis.

Leaf blades very obtuse at the base.

Leaf blades lance-oblong P. come.

Leaf blades lance-ovate P. tacticanum.

Leaves pilosulous or villosulous beneath along the costa, the
hairs spreading.

Leaves large, mostly 6-9 cm. wide P. patzulinum.

280 FIELDIANA: BOTANY, VOLUME 24

Leaves relatively small, mostly 3-5 cm. wide.

Leaf blades very acute on both sides at the base.

P. lanciferum.

Leaf blades obtuse or rounded on both sides at the base.

P. Hermes.

Leaves obviously pubescent on the upper surface, at least along the lower
part of the costa, or often almost invisibly scabrous or scaberulous,
but then more or less rough to the touch, often very scabrous and
rough.

Leaves smooth to the touch on the upper surface, either glabrous
between the veins or softly pubescent, not at all scabrous or
scaberulous.

Spikes normally conspicuously curved or recurved P. aduncum.

Spikes straight or nearly so.

Leaves with about 10 pairs of primary nerves, oblong-lanceolate.

P. barriosense.

Leaves with mostly 3-4 pairs of primary nerves, rarely with as

many as 7 pairs, but the leaves then broadly oval or elliptic.

Leaves deeply and narrowly cordate at the base, with a sinus

3-7 cm. deep or deeper, velutinous-pilosulous on the upper

surface, mostly 20-30 cm. wide P. auritum.

Leaves shallowly and often very broadly cordate at the base,

or else not cordate, the sinus, if any, very short, the blades

not velutinous-pilosulous on the upper surface.

Leaf blades acute on both sides at the base. . . .P. georginum.

Leaf blades cordate to very obtuse at the base, at least on

one side.
Pubescence of the nerves on the lower leaf surface of closely

appressed or subappressed hairs P. oradendron.

Pubescence of the nerves of the lower leaf surface pilosulous

or villosulous with spreading hairs.
Leaves lanceolate or lance-oblong, mostly 4-6 cm. wide,

conspicuously bullate P. jactatum.

Leaves broadly ovate to oval or broadly elliptic, mostly

8-20 cm. wide.

Leaves with 5-7 pairs of principal lateral nerves.
Leaf blades mostly 7-11 cm. wide, very oblique at

the base P. tecutlanum.

Leaf blades mostly 15-20 cm. wide, almost equal

at the base P. calophyllum.

Leaves with 3-4 pairs of principal nerves.

Petioles less than 1.5 cm. long P. minarum.

Petioles 2-5 cm. long or longer P. adamatum.

Leaves rough to the touch on the upper surface, usually very rough,
obviously scabrous or scaberulous between the veins, at least on
the younger leaves.
Nerves of the lower leaf surface strigose with closely appressed or

t subappressed hairs.

Leaves lanceolate or lance-oblong, 2-5 cm. wide.

Leaves conspicuously bullate, the nerves and veins impressed

' on the upper surface P. cayoense.

Leaves not obviously bullate, the nerves and veins little or not

at all impressed on the upper surface P. phaeophyllum.

Leaves broadly ovate to ovate or elliptic, mostly 5-11 cm. wide,
if less than 5 cm. wide broader than lanceolate or lance-
oblong P. scabrum.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 281

Nerves of the lower leaf surface pubescent, pilosulous, or hispidulous

with spreading hairs.
Leaves, at least when young, villous-pilose on the upper surface

with long spreading multicellular hairs P. fallens.

Leaves short-pilose or scabrous on the upper surface, never with
long spreading hairs.

Spikes, at least when young, with a conspicuous tail-like naked
tip P. Luxii.

Spikes obtuse or rounded at the apex, not with a naked tail-
like tip.

Leaves, at least the older ones, conspicuously bullate, very
rough to the touch on the upper surface.

Leaves very finely and closely areolate beneath, the ultimate
veinlets very strongly elevated P. alveolati folium.

Leaves coarsely bullate, the ultimate veinlets not or but

slightly elevated.
Leaves lanceolate or ovate-lanceolate, broadest near the

base P. cayoense.

Leaves variable in shape but mostly elliptic-ovate or
somewhat rhombic-elliptic, broadest at or near the
middle P. pseudoasperi folium.

Leaves not at all bullate, even in age, or, if rarely slightly
bullate, soft to the touch on the upper surface, or at
least not very scabrous.

Leaves, at least the young ones, densely hispidulous with
[ subappressed hairs over the whole upper surface, or

else densely and finely pubescent and then rather
[ soft and velvety to the touch, neither scabrous nor

scaberulous.

Upper leaf surface very densely hispidulous throughout,

both surfaces grayish when dried and almost con-

colorous P. indignum.

Upper leaf surface finely and densely pubescent, almost
velvety to the touch, usually green when dried or
sometimes blackish P. atlantidanum.

Leaves scabrous or scaberulous on the upper surface, rough
to the touch.

Leaf blades acute or short-acuminate, 3-4 cm. wide.

P. achoteanum.

Leaf blades abruptly acuminate or long-acuminate,
usually much wider.

Young branches finely short-hispidulous, the hairs
stiff, spreading or often in part subreflexed.

P. fraguanum.

Young branches villous-pilosulous with long spreading
multicellular hairs.

Leaf blades acute at the base on both sides.

P. chiquihuitense.

Leaf blades rounded or subcordate on at least one
side of the base P. pogonioneuron.

Piper achoteanum Trelease, Journ. Wash. Acad. Sci. 19: 328.
1929 (type from El Achote, near Siguatepeque, Honduras). P.
pictamentum Trelease in Yuncker, Field Mus. Bot. 17: 352. 1938

282 FIELDIANA: BOTANY, VOLUME 24

(type from Siguatepeque, Honduras). P. Yunckeri Trelease, op.
cit. 355. 1938 (type from El Achote, Honduras). Cola de rat6n.

At 400-1,100 meters, probably in moist or dry thickets; Alta ;
Verapaz; Guatemala (Amatitlan). Mountains of Honduras.

A rather stout, erect shrub 1-3 meters high, the branches strongly nodose,
with short internodes, densely hirtellous with spreading, whitish or fulvescent
hairs; petioles stout, not winged, 5 mm. long or usually shorter, often concealed
by the basal lobes of the blade; leaf blades lance-oblong or oblong, mostly 9-13
cm. long and 3-5 cm. wide, acute or subobtuse, very thick, yellowish green when
dry, densely and rather coarsely pellucid-punctate, very unequal and oblique at
the base, rather deeply cordate on one side, cordate to obtuse on the other, coarsely !
granular and very scabrous on the upper surface, not bullate, coarsely granular '
beneath and scabrous, usually hispidulous on the nerves and veins with some-
what spreading or subappressed hairs, penninerved, the nerves 3-5 on each side,
elevated, arcuate-ascending, the veins prominent and coarsely reticulate; peduncles
opposite the leaves, stout, 1 cm. long or somewhat shorter, densely hirsutulous;
spikes mostly 5-9 cm. long, 2.5-3 mm. thick, obtuse or subacute and submucronate;
bracts rounded-peltate at the apex, densely ciliate, pale.

A very distinct species, whose foliage reminds one somewhat of
certain species of Celtis.

Piper adamatum Trelease & Standley, sp. nov.

Dense, moist or wet, mixed, mountain forest, 1,500-2,900 meters;
so far as known, endemic, but doubtless extending to Chiapas; Alta
Verapaz; Guatemala; Chimaltenango; Quezaltenango; San Marcos
(Volcan de Tacana, along Rio Vega, near the Mexican boundary,
Steyermark 36354; type in Herbarium of Chicago Natural History
Museum).

An erect, sparsely branched shrub 1-3 meters high, the branches stout or
rather slender, with short or mostly elongate internodes, densely villous-pilose
with spreading, fulvous or brownish hairs; petioles stout or slender, 2.5-7 cm.
long, dilated and clasping at the base, not winged, densely villous-pilose or some-
times glabrate in age; leaf blades large, thin or rather thick, usually fuscescent
when dried, minutely pellucid-punctate or almost wholly opaque, conspicuously
and finely bullate, at least in age, broadly ovate to rounded-ovate, rarely lance-
ovate, mostly 14-19 cm. long and 7-14 cm. wide, abruptly acuminate, subequal
at the base and shallowly cordate, the two sides of about the same size, dull above,
villous-pilosulous over almost the whole surface or in age glabrate, often brownish
or fulvescent beneath, villous-pilosulous throughout, often between as well as
upon the veins, penninerved, the nerves 3-4 on each side, arcuate-ascending at
an angle of 45 degrees or more, the veins closely and conspicuously reticulate,
impressed on the upper surface, very prominent beneath; peduncles opposite the
leaves, stout, densely hirsutulous with spreading fulvescent hairs, at anthesis
1 cm. long or shorter; young spikes very stout, about 3.5 cm. long, 5-6 mm. thick,
mucronate at the apex with a tail-like tip 3 mm. long, the bracts very densely
f ul vous-hirsutul ous .

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 283

Frutex, ramis crassiusculis vel gracilibus dense pilis patentibus fulvis vel
brunnescentibus villoso-pilosis; petioli exalati basi dilatati 2.5-7 cm. longi, dense
villoso-pilosi vel interdum glabrati; lamina magna in sicco fuscescens tenuis vel
crassiuscula minute pellucido-punctata vel fere omnino epunctata, late ovata
usque rotundo-ovata, abrupte acuminata, basi subaequali cordata, lateribus fere
aequalibus, supra ubique villoso-pilosa, nervis venisque impressis, subtus ubique
saepe quoque inter venas villosopilosula, penninervia, nervis utroque latere vulgo
3-4 angulo semirecto vel latiore arcuato-adscendentibus, venis prominentibus arete
reticulatis, lamina inter venas profunde areolata; pedunculi oppositifolii dense
hirsutuli 1 cm. longi vel breviores; spicae juveniles crassae 3.5 cm. longae 5-6 mm.
crassae, apice longe caudato-mucronatae, bracteis dense fulvo-hirsutulis.

Piper aduncum L. Sp. PI. 29. 1753. P. multinervium Mart.
& Gal. Bull. Acad. Brux. 10, pt. 2: 130. 1843. P. Stevensonii Trelease
ex Standl. Field Mus. Bot. 12: 104. 1936, without Latin description
(type from Toledo District, British Honduras, N. S. Stevenson 93).
P. multinervium var. amplum Trelease in Yuncker, Field Mus. Bot.
17: 347. 1938 (type from La Libertad, Pete*n, C. L. Lundell 2556).
P. multinervium var. kantelolense Trelease, op. cit. 348 (type from
La Libertad, Pete"n, Lundell 3008). P. multinervium var. Skutchii
Trelease, op. cit. 350 (type from Colomba, A. F. Skutch 1299).
Cordoncillo; Cuturo (North Coast, fide Blake); Cordoncillo bianco;
Biritac (Coban, Quecchi).

Wet to dry thickets, often in second growth, sometimes in pine
forest, 1,600 meters or lower; Pete'n; Alta Verapaz; Baja Verapaz;
Izabal; Zacapa; Chiquimula; Jalapa; Jutiapa; Santa Rosa; Escuintla;
Guatemala; Chimaltenango; Suchitepequez; Retalhuleu; Quezal-
tenango; San Marcos; Huehuetenango. Southern Mexico; British
Honduras to El Salvador and Panama and probably farther south-
ward; West Indies; widely distributed in South America.

An erect shrub or small tree, 1-5 meters high or perhaps even larger, often
with a rather thick and clean, distinct trunk and a rather narrow and elongate
crown, the branchlets mostly straight and elongate, strigillose or hirtellous with
very short, white, spreading hairs, glabrate in age, the larger branches often
somewhat pendent; petioles short, usually much less than 1 cm. long, strigose
or short-hispidulous; leaves narrowly to broadly lance-oblong, mostly 13-20 cm.
long and 4.5-8 cm. wide, rather abruptly and narrowly long-acuminate, very
scabrous and rough to the touch on the upper surface, often somewhat lustrous,
slightly paler beneath, rather densely scabrous or appressed-hispidulous beneath,
strigose on the nerves and veins, the primary lateral nerves usually 3-4, sometimes
5, on each side, ascending at a very narrow angle, the uppermost arising near the
middle of the blade, the nerves and veins plane on the upper surface, the lateral
nerves strongly elevated beneath, the ultimate veins scarcely at all elevated,
inconspicuous; spikes solitary opposite the leaves, on slender or stout peduncles
1.5 cm. long or shorter, slender, normally very conspicuously curved, when mature

284 FIELD IANA: BOTANY, VOLUME 24

mostly 10-13 cm. long, about 3 mm. thick; flowers crowded in dense verticils,
the bracts puberulent, the spikes in anthesis pale green or greenish white.

Known in British Honduras by the names "Spanish elder,"
"Spanish Ella," and "cow's-foot." This species is one of the most
common ones of Guatemala and other parts of Central America,
abundant in many localities, easily recognized by its combination
of penninerved leaves and strongly curved flower spikes. It has
been reported from the region as P. elongatum Vahl, a synonym of
P. aduncum. The mature flower spikes .-of this, and doubtless of
other species, are used in Guatemala and British Honduras for
flavoring food, having more or less the properties of black pepper.
Schipp reports that in the latter country they are sometimes boiled
and eaten. The leaves of P. aduncum are of a lighter green than
those of most Central American species of Piper.

Piper aeruginosibaccum Trelease, Journ. Wash. Acad. Sci.
19: 328. 1929 (type from La Ceiba, Honduras). P. citrifolium var.
Cookii C. DC. Bot. Gaz. 70: 186. 1920 (type collected near Finca
Sepacuite", Alta Verapaz, 0. F. Cook & R. F. Griggs 697). P. onerosum
Trelease, Journ. Wash. Acad. Sci. 19: 335. 1929 (type from Tela,
Honduras). P. praeterlatum Trelease, loc. cit. (type from Lancetilla
Valley near Tela, Honduras). P. dimorphophyllum Trelease in
Standl. Field Mus. Bot. 12: 407. 1936 (type from Craig Point, Sibun
River, British Honduras, P. H. Gentle 1387). P. Gentlei Trelease,
loc. cit. (type from San Andre's, Corozal District, British Honduras,
Gentle 1077). P. nitidulifolium Trelease in Standl. op. cit. 408
(type from Tower Hill, British Honduras, J. S. Karling 26). P.
kantelulense Trelease, in Standl. loc. cit. (type from Kantelul, Pete"n,
C. L. Lundell 3173). P. kantelulense var. Gentlei Trelease in Standl.
loc. cit. (type from Prospect, Northern River, British Honduras,
Gentle 947). P. discolor Trelease in Standl. Field Mus. Bot. 17:
231. 1937 (type from La Libertad, Pete"n, Lundell 2565). P. emanci-
pationis Trelease in Standl. loc. cit. (type from La Libertad, Pete"n,
iMndell 3658). P. emancipations var. longum Trelease in Standl.
loc. cit. (type collected near La Libertad, Pete"n, Lundell 3052).
P. nitidulilaminum Trelease in Standl. op. cit. 232 (type from
Remate, Pet^n, Lundell 2077). P. plumbeicolor Trelease in Standl.
op. cit. 233 (type from La Libertad, Lundell 2554). Cordoncillo;
Pooczuyaax (Pete"n, Maya, fide Lundell).

Moist or wet thickets or forest, sometimes in pine forest or in
Manicaria swamps, 900 meters or lower; Pete"n; Alta Verapaz;
Izabal. Campeche; British Honduras.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 285

A shrub, commonly about 2 meters high, sometimes a small tree, the young
branches densely hispidulous or hirsutulous, sometimes glabrate in age or occa-
sionally almost glabrous from the first; petioles mostly 1 cm. long or less, some-
times longer in the lower leaves, stout, densely hispidulous or rarely glabrate;
leaf blades ovate-oblong or ovate-elliptic, mostly 12-20 cm. long and 4.5-9 cm.
wide, rather abruptly acuminate or long-acuminate, very unequal at the base
and more or less oblique, usually rounded or more or less cordate on one side and
obtuse on the other, one side much lower decurrent than the other, thick and
firm, very lustrous on the upper surface and often lustrous beneath, slightly paler
beneath, drying grayish green or sometimes blackish, finely pellucid-punctate,
glabrous above, smooth to the touch, hispidulous beneath, especially on the nerves,
with short sordid subappressed hairs, rather harsh to the touch, penninerved,
the nerves usually 3 on each side, arcuate-ascending, the upper ones arising at
or above the middle of the blade, the veins prominent beneath, laxly reticulate;
peduncles short, thick, densely puberulent or hispidulous; spikes rather stout,
mostly 5-7 cm. long and 3-4 mm. thick, erect, obtuse, the bracts densely pubescent.

The material we refer here is somewhat variable but not more
so than is to be expected in a tropical species. Some of the characters
by which the various species were separated by Trelease are more
or less obvious, but apparently they are individual variations. Seven
or eight other species, not listed here but described by Trelease,
are clearly referable to the same species, for which it is quite possible
that a still earlier name may be found.

Piper alveolatifolium Trelease, Journ. Wash. Acad. Sci. 19:
329. 1929 (type from Siguatepeque, Honduras). P. faviculiferum
Trelease in Yuncker, Field Mus. Bot. 17: 345. pi. 11. 1938 (type
from El Achote, above Siguatepeque, Honduras).

Moist or wet forest, often in pine-oak forest, 1,000-1,600 meters;
Zacapa; Chiquimula; Jalapa; Guatemala; Quiche*. Honduras.

A shrub or small tree 2-7 meters high, the young branches very densely fulvous-
hirsute with short spreading hairs, the internodes mostly short; petioles stout,
commonly 1-1.5 cm. long, densely hispidulous; leaf blades ovate or lance-ovate,
mostly 16-23 cm. long and 6.5-9 cm. wide, gradually acuminate to very long-
acuminate, very unequal at the base, rounded on one side, cordate or subcordate
at the other, opaque, very finely and deeply Initiate, not blackening when dry,
very scabrous on the upper surface, densely hirtellous beneath with spreading
whitish hairs, penninerved, the lateral nerves 3-4 on each side, very prominent,
the veins exceedingly prominent and very closely and finely reticulate, the lower
surface of the blade deeply alveolate; peduncles stout, equaling or shorter than the
petioles, densely fulvous-hispidulous; spikes stout, erect, greenish white, mostly
6-10 cm. long, 4 mm. thick, the bracts very densely hispidulous.

One of the most distinct and clearly marked of all local species,
easily recognized by its finely and closely bullate leaves, the veins
of the lower surface being greatly elevated and areolate, with deep

286 FIELDIANA: BOTANY, VOLUME 24

depressions between them. Three of the Guatemalan collections
were indicated as new species by Trelease, but all of them appear
to be identical with the two Honduran plants described as distinct
species.

Piper Amalago L. Sp. PL 29. 1753. P. medium Jacq. Icon. PI.
Rar. 1: 2. 1781. P. cubilquitzianum C. DC. Bot. Gaz. 70: 174.
1920 (type from Cubilgiiitz, Alta Verapaz, Turckheim 11.1440).
P. Gaumeri Trelease, Journ. Wash. Acad, Sci. 19: 332. 1929 (type
from Yucatan). P. Lundellii Trelease in Standl. Field Mus. Bot.
12: 408. 1936 (type from Honey Camp, British Honduras, C. L.
Lundell 570). P. lundelliifolium Trelease, op. cit. 17: 232. 1927 (type
from La Libertad, Pete"n, Lundell 3524). Cordoncillo.

Moist or wet thickets or mixed forest, 2,600 meters or lower;
Pete"n; Alta Verapaz; Baja Verapaz; Izabal; Zacapa; Escuintla;
Guatemala; Sacatepe"quez; Chimaltenango; Solola; Suchitepequez;
Retalhuleu; Quezaltenango; San Marcos; Huehuetenango. Southern
Mexico; British Honduras and southward, probably to Panama;
West Indies; South America.

A rather slender, often much branched shrub or small tree, commonly 1.5-3
meters high or sometimes as much as 6 meters, the branches glabrous or minutely
and often only sparsely puberulent; petioles about 1 cm. long, or often longer or
shorter, glabrous, vaginate only at the base; leaf blades green or blackish when
dried, minutely pellucid-punctate or almost wholly epunctate, rather thin but
usually rather firm and stiff, variable in shape, lance-elliptic to ovate-elliptic or
rounded-ovate, mostly 7-14 cm. long and 3.5-7 cm. wide, gradually or abruptly
acuminate or long-acuminate, obtuse to broadly rounded at the base, palmately
5-nerved, the nerves slender, prominent on both surfaces, the lateral ones some-
times much weaker than the others and inconspicuous or very irregular, the blades
symmetric at the base or only slightly oblique, the veins generally prominulous,
laxly reticulate, the leaves glabrous or sometimes minutely puberulent beneath on
the nerves; peduncles slender, equaling or usually somewhat longer than the
petioles, glabrous or puberulent; spikes cream-colored or greenish, slender, 3-7 cm.
long or longer, 2.5 mm. thick, the rachis puberulent or hirtellous, the bracts ciliate,
sometimes hirtellous; stamens 4, the anthers reniform; fruits glabrous.

It is quite possible that the material we refer here represents
more than a single species, but we do not find fixed characters by
which the forms, if there are any definite ones, may be distinguished.
Trelease has annotated many of the sheets as representing new
species. The shrub is known in British Honduras by the names
"Spanish elder" and "cordoncillo chico."

Piper atlantidanum Trelease, Journ. Wash. Acad. Sci. 19:
329. 1929 (type from La Ceiba, Honduras). P. praemollitum Tre-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 287

lease in Yuncker, Field Mus. Dot. 17: 352. 1938 (type from Lake
Yojoa, Honduras). P. viae-marginis Trelease, op. cit. 355. 1938
(type from Siguatepeque, Honduras). P. coyolense Trelease in
Yuncker, Field Mus. Bot. 9: 278. 1940 (type from Coyoles, Hon-
duras). P. elasmophyllum Trelease, op. cit. 278. 1940 (type from
Mount Cangrejal, Honduras). P. subaequilaterum Trelease, op. cit.
280. 1940 (type from region of La Ceiba, Honduras). P. atlanti-
danum var. sibunense Trelease in Standl. Field Mus. Bot. 12: 406.
1936 (type from Craig Point, Sibun River, British Honduras, P. H.
Gentle 1381). P. quadratilimbun Trelease in Standl. Field Mus. Bot.
17: 233. 1937 (type from El Paso, Pete"n, C. L. Lundell 1488).

Moist or wet thickets or mixed forest, 1,000 meters or lower;
Pete"n; Alta Verapaz. British Honduras; Honduras.

An erect shrub 1.5-3.5 meters high, the branches rather slender, with mostly
short internodes, densely pilose with short, spreading or somewhat reflexed, usually
fulvescent hairs; petioles stout, about 1 cm. long, sometimes longer or shorter,
densely short-pilose, not winged, dilated at the base; leaf blades mostly green
when dried, not bullate, rather thin, minutely pellucid-punctate, elliptic to ovate
or obovate-elliptic, mostly 10-16 cm. long and 5-8.5 cm. wide, rather abruptly
acuminate or long-acuminate, conspicuously unequal and oblique at the base,
generally cordate to rounded on one side, rounded to acute on the other, densely
scabrous-puberulent on the upper surface but soft and almost velvety to the
touch, densely velutinous-pubescent beneath, penninerved, the nerves 3-4 on
each side, very slender but prominent, ascending at an angle of 45 degrees or less,
slightly arcuate or almost straight, the veins usually not conspicuous, rather laxly
reticulate; peduncles opposite the leaves, stout, 8 mm. long or usually shorter,
densely hirtellous; spikes slender, erect-spreading, mostly 6-7 cm. long, 2.5 mm.
thick, cream-colored; bracts triangular-peltate at the apex, ciliolate; anthers
short-exserted; fruits mostly trigonous, somewhat depressed at the apex, glabrous
or nearly so.

Piper auritum HBK. Nov. Gen. & Sp. 1: 54. 1816. Santa
Maria (the usual name); Cordoncillo; Hoja de jute; Juniapra (fide
Pittier); Xaclipur (reported as Quecchi name); Obet (Coban,
Quecchi); Cana de oro (Quezaltenango).

Moist or wet thickets or forest, often in second growth, 1,800
meters or lower, most common at 900 meters or below; Peteii;
Alta Verapaz; Izabal; Zacapa; Santa Rosa; Escuintla; Guatemala;
Sacatepe"quez (probably only introduced); Chimaltenango; Solola;
Suchitepequez; Retalhuleu; Quezaltenango; San Marcos. Southern
Mexico; British Honduras to Salvador and Panama; Colombia;
described originally from Mexico.

A large, coarse, somewhat succulent herb, sparsely branched, or rarely woody
below and becoming tree-like, commonly about 2 meters high but occasionally

288 FIELDIANA: BOTANY, VOLUME 24

as much as 6 meters, the branches stout, sparsely pubescent or glabrate; leave;
on short or elongate, stout petioles, the petioles broadly winged, more or less*
dilated and clasping at the base; leaf blades very large, thin and soft, usually
drying bright yellowish green, broadly ovate or oblong-ovate, as much as 60 cm.
long and 35 cm. wide but usually much smaller, acute or abruptly short-acuminate,
deeply and narrowly cordate at the base, the basal lobes rounded, one of them
extending 1.5-3 cm. lower on the costa than the other, slightly paler beneath,
softly puberulent or short-pilosulous on both surfaces, with usually 3 pairs of
nerves above the basal ones; peduncles simple, opposite the leaves, about 3 cm.;
long; spikes pale green, 4 mm. thick, commonly 20-25 cm. long, the pale peltate
scales finely puberulent.

In British Honduras called "bullhoof" and "maculan" (Maya);:
"matarro" (Honduras); "momo" (Yucatan). This is one of the
commonest and most widespread species of the genus, found in the
lowlands almost throughout Central America, at least where there
is abundant shade. The plants often form dense thickets in aban-
doned land, partly, no doubt, on account of the great abundance
of seeds produced. The fruits of many or most of the Piperaceae
are extremely viscid, and so small that they can be dispersed by all
kinds of animals, even insects. This plant is a conspicuous and rather
handsome one because of its large leaves, but these become limp
immediately, if they are broken from the branch. Large plants,
particularly those that become really woody, often shed their lower
branches, so that only a small number of them are left at the very
top of the trunk. The crushed leaves and stems have a strong
odor similar to that of sarsaparilla, and the leaves are much used
to flavor food of all kinds, particularly meat dishes, and especially
the common local snails or jutes of the small streams.

Piper barriosense Trelease & Standley, sp. nov.

Moist or wet, lowland thickets or mixed forest, at or near sea
level; endemic; Izabal (type from Entre Rios, Standley 72611, in
Herbarium of Chicago Natural History Museum; also at Puerto
Barrios).

A shrub 2 meters high, the branches stout or slender, with short or elongate
internodes, conspicuously pale-lenticellate, densely and softly sordid-pilosulous
when young, conspicuously nodose; petioles strongly dilated at the base, when
young broadly winged throughout, the shorter side 7-12 mm. long, the longer
side as much as 2 cm. long, densely pilosulous; leaf blades thin, brownish and
blackish when dried, minutely pellucid-punctulate, dull, lanceolate or narrowly
lanceolate, mostly 14-22 cm. long and 3.5-7 cm. wide, gradually long-acuminate,
sparsely short-pilosulous above when young but soon glabrate, rather densely
and softly short-pilosulous beneath over the whole surface, very unequal at the
base, rounded or obtuse on either side, penninerved, the nerves about ten on each

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 289

side, divergent at wide angle, arcuate, very slender and inconspicuous, the veins
very slender and inconspicuous, laxly reticulate; spikes unknown.

Frutex erectus, ramis pallido-lenticellatis, statu juvenili dense molliter sordido-
pilosulis; petioli late alati crassi; lamina lanceolata vel anguste lanceolata longe
sensimque attenuato-acuminata, basi insigniter inaequalis utroque latere obtusa
vel rotundata, rarius latere altero subacuta, penninervia, nervis utroque latere
ca. 10, supra sparse pilosula vel glabrata, subtus sat dense breviter molliterque
pilosula.

In general appearance this is like P. geniculatum Swartz, of which
it may be only a variety, but it differs conspicuously from other
material of that species in its abundant pubescence.

Piper Biritak Trelease & Standley, sp. nov. Biritak (Coban,
Quecchi).

Moist or wet, mixed or pine, mountain forest, sometimes on lime-
stone, 900-1,500 meters; endemic; Alta Verapaz (type from Coban,
Standley 69345; in Herbarium of Chicago Natural History Museum);
Huehuetenango (Maxbal).

A slender shrub 1-2 meters high, often with long straggling branches, the upper
internodes usually short, sometimes elongate, very densely hispidulous with short
sordid spreading hairs; petioles stout, mostly 5-10 mm. long, sometimes longer,
vaginate only at the base, densely hispidulous; leaf blades strongly bullate, at
least in age, grayish or sometimes fuscescent when dry, lustrous on the upper
surface, finely pellucid-punctate, mostly lanceolate or narrowly lanceolate, some-
times ovate-lanceolate, mostly 13-25 cm. long and 4-7.5 cm. wide, narrowly and
gradually very long-attenuate, strongly unequal and oblique at the base, usually
cordate or subcordate on one side and rounded or obtuse on the other, glabrous
above or nearly so, the nerves and veins strongly impressed, at least in age, densely
hispidulous or strigose beneath on the nerves and veins, glabrous or nearly so
between them and strongly granular, penninerved, the nerves 3 or 4 on each side,
strongly elevated, the veins conspicuously elevated and closely reticulate; spikes
unknown.

Frutex, ramis gracilibus interdum elongatis, internodiis plerumque brevibus
pilis brevibus sordidis patentibus dense hispidulis; folia breviter petiolata, petiolo
vulgo 5-10 mm. longo dense hispidulo, basi tantum vaginante, insigniter bullata,
supra lucida, minute pellucido-punctata, anguste lanceolata usque ovato-lanceo-
lata modica vel magna subcoriacea, sensim longeque attenuato-acuminata, basi
inaequalia et obliqua, uno latere cordata vel subcordata, altero obtusa vel rotund-
ata, penninervia, nervis utroque latere 3-4, venis supra impressis, subtus elevatis
arete reticulatis; spicae ignotae.

A rather common and well-marked plant of the forests of Alta
Verapaz, represented by about a dozen collections.

Piper brevilimbum C. DC. Candollea 1: 128, 238. 1923; ex
Schroeder, Candollea 3: 136. 1926.

290 FIELDIANA: BOTANY, VOLUME 24

Wet, mixed, mountain forest, 1,250-1,500 meters; endemic; Alta
Verapaz (type from Coban, Turckheim II. 1625; collected several
times in the Coban region); Chiquimula (Cerro Tixixi).

A slender branched erect shrub 1.5-2.5 meters high, the branches slender,
with short internodes, sparsely and inconspicuously pubescent or glabrate; petioles
slender, mostly 5 mm. long or less; leaf blades ovate-elliptic or lance-elliptic,
5.5-12 cm. long, 3-5.5 cm. wide, rather abruptly acuminate, acute at the base
and slightly unequal, blackish when dried, dull, opaque, penninerved, the nerves
usually three on each side, slender and very inconspicuous, arcuate-ascending,
arising from the middle of the blade or lower, the blades glabrous throughout;
peduncles slender, glabrous, about 1 cm. long; spikes rather stout, 3.5-6 cm.
long, 3 mm. thick; bracts rounded-subpeltate, ciliate.

Piper brevipedunculatum C. DC. Ann. Cons. Jard. Bot.
Geneve 2: 266. 1898.

Known only from the type, Suchitepequez, banks of the Rio
Grande, Mazatenango, Bernoulli 470.

Branches glabrous; leaves glabrous, on petioles 1-2.5 cm. long, firm-membra-
naceous, ovate, as much as 15 cm. long and 9 cm. wide, rather long-acuminate,
unequally rounded at the base, pellucid-punctate, the costa emitting four lateral
nerves on each side; peduncles shorter than the petioles, 1 cm. long or less, the
spikes about half as long as the leaves, 5 mm. thick in anthesis; bracts truncate
at the apex, sparsely ciliolate; fruits trigonous-obpyramidal, glabrous.

We have seen no material of this species and have not attempted
to include it in the key. It probably antedates some species listed
here, but from the description alone we could not determine its
relationship.

Piper brujoense Trelease & Standley, sp. nov. Santa Maria
(Chiquimula).

Moist or wet, mixed, mountain forest, 1,700-2,500 meters;
endemic; Zacapa (Sierra de las Minas); Chiquimula (type from
middle slopes of Montana Norte to El Jutal, on Cerro Brujo, south-
east of Conception de las Minas, Steyermark 31049; in Herbarium
of Chicago Natural History Museum).

A shrub or small tree 1.5-6 meters high, the branches slender, rather densely
hispidulous with spreading hairs or glabrate, the internodes mostly short; petioles
stout, scarcely more than 3 mm. long, densely hispidulous; leaf blades rather
thick, grayish green when dry, densely and minutely pellucid-punctate, oblong-
lanceolate, 10-17 cm. long, 3-4.5 cm. wide, narrowly attenuate-acuminate, con-
spicuously unequal and oblique at the narrow ba'se, shallowly cordate on one
side, acute or obtuse on the other, dull and glabrous above, much paler beneath,
hispidulous with short spreading hairs on the nerves and veins, glabrous or nearly
so between them, penninerved, the nerves 2-3 on each side, arcuate-ascending

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 291

at an angle of about 45 degrees, the veins prominulous or obscure, laxly reticulate;
peduncles opposite the leaves, rather stout, 12 mm. long or shorter, glabrous;
spikes rather stout, erect, about 7.5 cm. long and 3 mm. thick, obtuse; bracts
pale, densely puberulent; ovaries glabrous or nearly so.

Frutex, ramis gracilibus pilis patentibus subdense hispidulis vel glabratis;
petioli vix ultra 3 mm. longi dense hispiduli; lamina crassiuscula in sicco griseo-
viridis dense minute pellucido-punctata, oblongo-lanceolata anguste longeque
attenuato-acuminata, basi manifeste inaequali et obliqua uno latere breviter
cordata, altero acuta vel obtusa, supra glabra, subtus pallidior ad nervos venasque
dense hispidula, penninervia, nervis utroque latere 2-3 angulo angusto arcuato-
adscendentibus; pedunculi oppositifolii usque ad 12 mm. longi, glabri, spicis
crassiusculis ca. 7.5 cm. longis et 3 mm. crassis.

Piper calophyllum C. DC. in Bonn. Smith, Bot. Gaz. 33: 257.
1902.

Moist or wet, mixed, lowland forest, sometimes or perhaps always
on limestone, 900 meters or lower; Alta Verapaz (type from Cubil-
giiitz, Turckheim 7959; collected also about Chirriacte*).

A rather coarse shrub 2-3.5 meters high, with few branches, the young branches
very thick and stout, villous-hirsute, as much as 11 mm. thick; petioles very stout,
about 4 cm. long, vaginate throughout, sparsely villous-hirsute; leaf blades very
large, rather thick, somewhat blackish when dried, ovate, 25-35 cm. long, 15-20
cm. wide, abruptly acute or short-acuminate, rounded at the base and shallowly
and narrowly cordate, with small rounded lobes, green above, somewhat lustrous,
villous-pilose along the nerves and veins or glabrate, somewhat paler beneath,
brownish when dried, densely and softly short-pilose, penninerved, the nerves
8-9 on each side, prominent beneath, the veins prominent and laxly reticulate;
spikes white or greenish white, erect, borne on a very thick, pilose or glabrate
peduncle about 1 cm. long, very stout, conspicuously caudate at the apex, 5-6.5
cm. long, 6 mm. thick, the bracts truncate-triangular, glabrous; stamens 4, the
anthers reniform, much shorter than the filaments; ovary glabrous, somewhat
narrowed at the apex, the 3 stigmas sessile, linear, short, recurved.

Piper cayoense Trelease in Standl. Field Mus. Bot. 12: 407.
1936 (type from San Antonio, El Cayo District, British Honduras,
H. H. Bartktt 13070). P. tikalense Trelease in Standl. Field Mus.
Bot. 17: 234. 1937 (type from Tikal, Pet<n, H. H. Bartlett 12595).
P. yalochanum Trelease in Standl. loc. cit. (type collected between
Dos Arroyos and Yaloch, Pete"n, Bartlett 12849). Biritak.

Moist or wet forest or thickets, sometimes in pine forest, often
on limestone, 1,200-1,450 meters or lower; endemic; Pete*n (Laguna
de Mexico); Alta Verapaz. British Honduras.

A slender shrub 1-2.5 meters high, sparsely branched, often with elongate
and rather weak branches, the young branches densely pale-hispidulous, the
internodes short or often elongate; petioles very stout, 7 mm. long or shorter,
densely hispidulous, not winged, dilated at the base; leaf blades rigid, subcoriaceous,

292 FIELDIANA: BOTANY, VOLUME 24

strongly and rather closely bullate, opaque or nearly so, mostly lanceolate or
oblong-lanceolate, rarely ovate, usually 7-14 cm. long and 2.5-5 cm. wide, narrowly
long-acuminate or attenuate-acuminate, very unequal and oblique at the base,
cordate on both sides or cordate on one side and rounded on the other, densely
pustulate on the upper surface, scabrous and very rough to the touch, dull or
slightly lustrous, often fuscescent when dried, the nerves and veins, strongly
impressed, densely appressed-pilose beneath on the nerves and veins, or rarely
pilose with spreading hairs, usually short-pilose with spreading hairs between
them, penninerved, the nerves 3-4 on each side, very prominent, arcuate-ascending
at an angle of 45 degrees or usually more, the veins strongly elevated and closely
reticulate, the surface areolate between them; peduncles opposite the leaves,
stout, about 6 mm. long, hispidulous; spikes erect-ascending, rather stout, very
obtuse, at anthesis about 4.5 cm. long and 3 mm. thick; bracts pale, densely
hispidulous.

Piper Chamissonis (Miq.) Steud. Norn. Bot. ed. 2. 2: 340.
1841; ex C. DC. in DC. Prodr. 16, pt. 1: 283. 1869. Artanthe Cha-
missonis Miq. Syst. Piper. 457. 1844.

Dense, moist or wet, mixed, mountain forest, 900-2,500 meters;
Alta Verapaz; Quezaltenango; San Marcos. Southern Mexico.

A shrub 1.5-2.5 meters high, sparsely branched, erect, the branches glabrous
or glabrate, rather stout, often granular; petioles stout, 1.5 cm. long or shorter,
often much less than 1 cm. long, glabrous or sparsely strigose or hirtellous, vaginate
only at the base; leaf blades usually thin, generally dark green when dry, minutely
pellucid-punctate or almost wholly opaque, dull when dried, oblong-elliptic to
elliptic or sometimes ovate-elliptic, mostly 13-22 cm. long and 4.5-9.5 cm. wide,
rather abruptly acuminate or long-acuminate, rather conspicuously unequal and
often oblique at the base, one side usually much longer decurrent than the other,
acute on both sides, green above, glabrous, somewhat paler beneath, more or less
strigose or hirtellous beneath on the nerves and veins or almost wholly glabrous,
penninerved, the nerves 3-4 on each side, arcuate-ascending at an angle of about
45 degrees or less, very slender but prominent, the veins mostly inconspicuous,
very laxly reticulate; peduncles opposite the leaves, rather slender, mostly 10-14
mm. long, glabrous or nearly so; spikes erect-spreading, rather slender, white or
greenish, mostly 7-10 cm. long when fully developed and 3 mm. thick, obtuse;
bracts triangular-peltate at the apex, densely ciliate, pale; fruit glabrous.

Piper cheyennense Trelease & Standley, sp. nov.

Wet mixed forest, at or little above sea level; endemic; Izabal
(type collected along Rio Juyama, southeast of Cheyenne, about
15 miles southwest of Bananera, Steyermark 39154; in Herbarium of
Chicago Natural History Museum).

A shrub 1.5-2.5 meters high, glabrous throughout, the branches rather slender,
with short internodes; petioles stout, 2-3.5 cm. long, vaginate only at the base;
leaf blades rather thick and firm, green when dried, opaque, oblong-ovate, 18-23
cm. long, 7-10.5 cm. wide, abruptly acuminate, subequal and rounded or broadly
rounded at the base, concolorous, penninerved, the nerves mostly 5 on each side,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 293

slender, prominent on both surfaces, ascending at an angle of 45 degrees or more,
slightly arcuate, the veins prominent beneath, laxly reticulate; peduncles about
8 mm. long or even shorter, the spikes erect-spreading, 10-11.5 cm. long, 3 mm.
thick, mucronate at the apex; bracts with an ovate apex, pale green, glabrous,
ciliate.

Frutex omnino glaber, ramis subgracilibus, internodiis brevibus; folia magna
2-3.5 cm. longe petiolata, petiolo basi tantum vaginante, oblongo-ovata abrupte
acuminata, basi subaequali et symmetrica rotundata vel late rotundata, penni-
nervia, opaca, in sicco viridia, nervis utroque latere saepius 5 arcuato-adscendenti-
bus; spicae oppositifoliae breviter pedunculatae graciles 10-11.5 cm. longae 3 mm.
crassae.

Besides the type, we have seen one other collection, gathered
between Dartmouth and Morales, which Trelease designated as the
type of another new species, but the two collections are identical.

Piper chiquihuitense Trelease & Standley, sp. nov.

Moist or wet thickets or mixed mountain forest, 1,200-1,500
meters; endemic; Quezaltenango (type from Chiquihuite, Standley
68146; in Herbarium of Chicago Natural History Museum; also in
the region of Finca Pirineos).

An erect shrub about 3 meters high, densely pilose with very unequal hairs,
some of the hairs long, soft, and spreading, fulvescent; petioles mostly 1.5-2 cm.
long, not winged, dilated at the base, densely pilose with soft spreading hairs;
leaf blades thin, scarcely fuscescent when dried, opaque or nearly so but with
translucent veins, elliptic or ovate-elliptic, mostly 14-25 cm. long and 6-10 cm.
wide, long-acuminate, somewhat unequal and conspicuously oblique at the base,
acute on each side, densely scabrous and very rough to the touch, hispidulous
along the nerves, not at all rugose, densely scabrous beneath, pilose on the nerves
and veins, part of the hairs long, soft, and spreading, penninerved, the nerves
3-5 on each side, ascending at an angle of 45 degrees or less, slightly arcuate,
very slender, the veins very slender, not conspicuous, laxly reticulate; peduncles
opposite the leaves, 10-18 mm. long, scabrous and often short-hispidulous; spikes
erect or ascending, 8-9.5 cm. long, 4 mm. thick, obtuse; bracts rounded-peltate
at the apex, pale, densely pubescent; anthers short-exserted ; fruits obtusely
angulate, about 1 mm. long, somewhat depressed at the apex, sparsely puberulent
or granulate above.

Frutex, ramis dense pilis inaequalibus pro parte longis mollibus patentibus
pilosis, internodiis plerumque elongatis; petioli 1.5-2 cm. longi crassiusculi exalati
basi dilatati dense patenti-pilosi ; lamina sat magna opaca, venis pellucidis, in
sicco vix fuscescens, elliptica vel ovato-elliptica, longiacuminata, basi inaequali
atque obliqua utroque latere acuta, supra dense scabra et asperrima, ad nervos
venasque hispidula, subtus dense scabra, ad nervos venasque pilis pro parte
longis mollibus patentibus pilosa, penninervia, nervis utroque latere 3-5 angulo
angusto adscendentibus subarcuatis, venis laxe reticulatis; pedunculi oppositifolii
10-18 mm. longi, spicis 8-9.5 cm. longis 4 mm. crassis.

294 FIELDIANA: BOTANY, VOLUME 24

Piper come Trelease & Standley, sp. nov.

Known only from the type, Jalapa, Volcan de Jumay, moist
or wet forest, about 2,000 meters, Steyermark 32350 (in Herbarium
of Chicago Natural History Museum).

A shrub 3 meters high, erect, the branches stout, with short internodes, almost
glabrous but when young with sparse appressed hairs; petioles stout, about 1 cm.
long, glabrous or nearly so, not winged, dilated at the base; leaf blades dark green
when dried, dull, densely and finely pellucid-punctate, lance-oblong, 13-22 cm.
long, 5-7.5 cm. wide, gradually long-acuminate, conspicuously unequal and some-
what oblique at the base, rounded or obtuse on either side, not at all bullate,
glabrous above, the nerves inconspicuous, scarcely paler beneath, sparsely strigose
along the nerves and principal veins, glabrous elsewhere, penninerved, the nerves
3-4 on each side, arcuate-ascending at an angle of more than 45 degrees, the veins
inconspicuous, often brownish, laxly reticulate; spikes unknown.

Frutex, ramis crassiusculis internodiis brevibus in statu juvenili sparse
strigosis cito glabratis; petioli crassi ca. 1 cm. longi exalati, basi dilatati; lamina
tenuis in sicco obscure viridis pellucido-punctata lanceolato-oblonga modica,
sensim longiacuminata, basi inaequali atque obliqua utroque latere rotundata vel
obtusa, supra glabra, subtus ad nervos venasque sparse strigosa, penninervia,
nervis utroque latere 3-4 arcuato-adscendentibus, venis non conspicuis laxe
reticulatis; spicae ignotae.

Piper coronanum Trelease & Standley, sp. nov.

Known only from the type, San Marcos, moist mixed forest,
Volcan de Tajumulco, between Finca El Porvenir and Loma Corona,
nine miles northwest of El Porvenir, 1,300-2,000 meters, Steyermark
37741 (in Herbarium of Chicago Natural History Museum).

A shrub, glabrous throughout, the upper internodes short; petioles stout,
8-10 mm. long, vaginate only at the base; leaf blades thin, dull green when dried,
very minutely pellucid-punctate or almost opaque, elliptic-oblong, 13-20 cm.
long, 5.5-8.5 cm. wide, broadest at or slightly below the middle, acuminate,
acute at the base and somewhat decurrent on the petiole, equal or nearly so,
penninerved, the nerves mostly five on each side or sometimes only four, arcuate-
ascending at an angle of 45 degrees or slightly more, very slender, prominent
beneath, the veins obscure, very laxly reticulate; peduncles very stout, about
1 cm. long; spikes green, spreading, stout, straight, about 9 cm. long and 4 mm.
thick, obtuse or subapiculate; bracts small, densely pale-ciliate; ovaries glabrous.

Frutex erectus omnino glaber; folia breviter petiolata tenuiter membranacea
in sicco sordide viridia, minutissime punctata vel fere epunctata, elliptico-oblonga,
majuscula, acuminata, basi subaequali acuta et subdecurrentia, penninervia,
nervis utroque latere saepius 5 arcuato-adscendentibus tenerrimis, venis obscuris
laxissime reticulatis; spicae oppositifoliae breviter crasseque pedunculatae ca. 9
cm. longae atque 4 mm. crassae.

Piper cristinanum Trelease & Standley, sp. nov.
Moist or wet, brushy hillsides, 225 meters or lower; endemic;
Izabal (type collected between Milla 49.5 and Cristina, about 70

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 295

meters, Steyermark 38677; in Herbarium of Chicago Natural History
Museum).

A branched shrub 1.5-2.5 meters high, the branches slender, very sparsely
short-hirsute or glabrous, the upper internodes short; petioles stout, 10 mm. long
or mostly shorter, vaginate for their whole length, densely hirsute; leaf blades
drying grayish, sparsely and minutely pellucid-punctate or mostly opaque, rather
thick and firm, lance-oblong or ovate-oblong, 9-15 cm. long, 3.5-5.5 cm. wide,
rather abruptly and narrowly caudate-acuminate, equal or somewhat unequal
at the base, rounded or broadly rounded and subcordate, hirsute on the upper
surface and very rough to the touch, or most of the hairs deciduous in age, con-
colorous beneath, hirsute or short-hispid beneath, at least on the nerves and veins,
palmately 5-nerved, the nerves slender but prominent, the veins divaricate at a
right angle from the costa, at least the principal ones strongly elevated, coarsely
reticulate; spikes unknown.

Frutex erectus ramosus, ramis gracilibus sparse breviter hirsutis vel glabris;
petioli crassiusculi 10 mm. longi vel breviores tota longitudine vaginantes; lamina
in sicco griseo-viridis crassiuscula sparse minute pellucido-punctata vel fere opaca,
lanceolato-oblonga vel ovato-oblonga, modica, subabrupte angusteque caudato-
acuminata, basi aequali vel paullo inaequali rotundata vel late rotundata atque
subcordata, supra hirsuta et asperrima, subtus concolor saltern ad nervos venasque
hirsuta vel breviter hispida, palmatim 5-nervia; spicae ignotae.

Among local species with palmate nervation this may be recog-
nized at once by the coarse and very rough pubescence of the leaves.

Piper curvatipes Trelease in Standl. Field Mus. Bot. 17: 231.
1937.

Moist or wet thickets or forest, 150 meters or lower; Pete"n (type
from Nictun, Lago de Pete"n, C. L. Lundell 3122). British Honduras

(Vaca).

A shrub, glabrous throughout, the branches rather slender, nodose, often
tortuous, the internodes short, somewhat granular; petioles stout, 8 mm. long or
shorter, slightly dilated at the base but scarcely vaginate; leaf blades firm, drying
green, coarsely pellucid-punctate, lance-oblong, often narrowly so, 8-12 cm. long,
2.5-4.5 cm. wide, rather abruptly long-acuminate, obtuse or subacute at the
base, conspicuously unequal, one of the sides decurrent, lower than the other,
slightly lustrous above, the nerves prominent, scarcely paler beneath, the nerves
usually three on each side, ascending at a very narrow angle, the veins prominent,
laxly reticulate; peduncles rather slender, mostly less than 1 cm. long; spikes rather
stout, 3-4 cm. long, 3-4 mm. thick, short-pointed at the apex, erect or recurved,
straight or nearly so; bracts rounded and subpeltate, densely pubescent; stigmas 3.

Piper diandrum C. DC. Ldnnaea 37: 364. 1871-73.

Moist or wet to rather dry forest or sometimes on brushy hill-
sides, 1,650 meters or lower, most common below 900 meters;
Pete"n; Alta Verapaz; Baja Verapaz; El Progreso; Izabal; Zacapa;

296 FIELDIANA: BOTANY, VOLUME 24

Santa Rosa; Escuintla; Guatemala; Suchitepequez; Retalhuleu;
Quezaltenango; San Marcos; Huehuetenango. Southern Mexico.
British Honduras; Salvador.

A shrub or tree, commonly 2-6 meters high, rather sparsely branched, appear-
ing glabrous to the naked eye, the young branches stout, glabrous; petioles rather
stout, very variable in length, mostly 2.5-7 cm. long but sometimes even longer,
usually glabrous; leaf blades broadly ovate to orbicular-ovate, mostly 15-22 cm.
long and 8.5-19 cm. wide, rather thick and firm, obscurely if at all pellucid-
punctate, abruptly acuminate, subcordate to rather deeply and sometimes narrowly
cordate at the base, or sometimes rounded in the upper leaves, not blackening
when dried, green above, glabrous or practically so, very slightly paler beneath,
glabrous or often puberulent on the nerves, usually palmately 9-nerved, the
nerves slender but prominent, the costa emitting numerous veins, these all slender
and equally prominent, divergent at a right angle or some of the uppermost often
arcuate and upcurved; peduncles opposite the leaves, mostly 1.5-2.5 cm. long,
glabrous or nearly so ; spikes very slender, often equaling or exceeding the leaves,
mostly 12-25 cm. long, 2-3 mm. thick or in fruit as much as 4 mm., often curved
or pendent, the rachis pubescent; bracts truncate-peltate at the apex, hirtellous
on the margins; stamens 2; stigmas 4, short; fruit glabrous, very small.

Much of the material we place here was referred by Trelease to
P. patulum or its unpublished varieties. It is, however, fully distinct
from that species, or at least is easily separated by its general appear-
ance, although definite characters for separating the two are not so
obvious. Some of the specimens were referred by Trelease to P.
variabile to which P. diandrum also is closely related, but the two
can be separated fairly well by the characters given in the key.

Piper Donnell-Smithii C. DC. ex Bonn. Smith, Enum. PL
Guat. 2: 95. 1891, nomen; in Bonn. Smith, Bot. Gaz. 19: 259. 1894;
P. eldoradense Trelease ex Standl. Field Mus. Bot. 12: 103. 1936,
without Latin description (type from Eldorado Road, British Hon-
duras, W. A. Schipp 1005).

Moist or wet, mixed forest, 1,600 meters or lower; Pete"n; Alta
Verapaz (type from Pansamala, Turckheim 975); Izabal; Huehue-
tenango (region of Ixcan). British Honduras.

A slender branched erect shrub or small tree, 1.5-6 meters high, glabrous
throughout; petioles slender, 1 cm. long or usually shorter, scarcely vaginate but
slightly dilated at the base; leaf blades lanceolate to ovate-oblong, usually some-
what blackish and dull when dry, scarcely at all lustrous, opaque, mostly 11-15
cm. long and 3.5-5.5 cm. wide, very narrowly long-acuminate, acute or subacute
at the base and subequal, slightly paler beneath, penninerved, the nerves 3-5 on
each side, arcuate-ascending, the veins prominent beneath, laxly reticulate;
peduncles about as long as the petioles, the spikes whitish or dull green, erect,
mostly 4-8 cm. long, 3 mm. thick, very obtuse; bracts truncate-peltate at the apex,
hirtellous on the margin; stamens 3; fruit obpyramidal-trigonous, sparsely hirtellous.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 297

Piper exactum Trelease & Standley, sp. nov.

Known only from the vicinity of the type locality, Quezaltenango,
moist or wet forest, slopes of Volcan de Zunil at or above Aguas
Amargas, 2,450-2,850 meters (type, Standley 65462 in Herbarium of
Chicago Natural History Museum).

An erect, sparsely branched shrub about 2.5 meters high, the branches rather
slender, sparsely hirtellous or almost wholly glabrous; petioles mostly 5-7 cm.
long, rather stout, thinly hirtellous or glabrate, vaginate only at the base; leaf
blades large, thin, blackening when dried, very obscurely and minutely if at all
pellucid-punctate, deep green and glabrous above, smooth, somewhat paler
beneath, rather sparsely or densely strigillose and hispidulous beneath on the
nerves and veins, glabrous or nearly so between the veins, broadly ovate or rounded-
ovate, mostly 14-20 cm. long and 9.5-12 cm. wide, abruptly acuminate, broadly
rounded at the base and shallowly and narrowly cordate, with subequal, broadly
rounded basal lobes, penninerved, the principal nerves 3-4 on each side, very
slender but rather prominent, strongly arcuate, ascending at an angle of 45 degrees
or slightly more, the veins rather prominent beneath, laxly reticulate; spikes
unknown.

Frutex altus, ramis subgracilibus sparse hirtellis vel fere glabris; petioli
crassiusculi 5-7 cm. longi basi tan turn vaginantes; lamina tenuis magna in sicco
fuscescens obscure minuteque pellucido-punctata vel fere epunctata, supra intense
viridis glabra, subtus paullo pallidior ad nervos venasque sat dense strigillosa
et sparse hirtella, abrupte acuminata, basi subaequali late rotundata et breviter
angusteque cordata, penninervia, nervis utroque latere 3-4 arcuato-adscendentibus;
spicae ignotae.

Piper fallens Trelease, Journ. Wash. Acad. Sci. 19: 332. 1929.

Moist or wet, mixed, mountain or lowland forest, 1,500 meters
or lower; Alta Verapaz; Izabal; Chiquimula; Solola; Quezaltenango;
San Marcos. Honduras, the type from Lancetilla Valley near Tela.

An erect shrub 1.5-3 meters high, sparsely branched, the branches rather
slender, densely villous-pilose with long, slender, spreading, multicellular, mostly
brownish hairs, the internodes usually elongate; petioles stout, about 7 mm.
long, densely villous-pilose, not winged; leaf blades rather thin and often flaccid,
dull green or fuscescent when dried, dull, minutely pellucid-punctate, lance-oblong
to lance-elliptic or ovate-elliptic, mostly 14-24 cm. long and 4-9 cm. wide, narrowly
long-attenuate or attenuate-acuminate, conspicuously unequal and oblique at the
base, cordate or rounded on one side, rounded or obtuse on the other, densely
long-pilose on the upper surface, at least when young, with rather weak, mostly
spreading, multicellular hairs, densely villous-hirsute beneath with long spreading
hairs, at least on the nerves and veins, penninerved, the nerves 3-4 on each side,
ascending at an angle of 45 degrees or less, somewhat arcuate or almost straight,
the veins not very conspicuous, laxly reticulate; peduncles stout or rather slender,
opposite the leaves, mostly 1-1.5 cm. long, sometimes 2.5 cm. long, densely
hirsute; spikes slender, mostly 8-10 cm. long, almost 3 mm. thick, mucronate or
obtuse; bracts triangular-subpeltate at the apex, pale, densely hirtellous, at least

298 FIELDIANA: BOTANY, VOLUME 24

on the margins; fruits depressed and papillose at the apex; stigmas 2, slender,
borne on a short style.

Piper flavidum C. DC. ex Bonn. Smith, Enum. PL Guat. 2:
66. 1891, nomen; Bot. Gaz. 19: 258. 1894. Monte de agua.

Usually on rocks in the edges of swift streams, 1,350 meters or
lower; endemic; Alta Verapaz (type from Barranco de Rubelcruz,
J. D. Smith 1744); Izabal; Quiche"; Huehuetenango.

An erect shrub, commonly a meter high or less, with rather few, erect branches,
the branches strongly nodose, glabrous, with short internodes; leaves on petioles
6 mm. long or shorter, linear-lanceolate, mostly 8-13 cm. long and 1.5-2 cm. wide,
narrowly long-attenuate, attenuate to the base, glabrous, usually yellowish green
when dried, the costa and nerves impressed on the upper surface, prominent be-
neath, the lateral nerves 2-3 on each side, ascending at an extremely narrow angle,
the upper ones extending nearly or quite to the apex, the veins prominent, laxly
reticulate; peduncles glabrous, equaling or longer than the petioles; spikes mostly
3-4 cm. long, 3 mm. thick, the bracts pubescent on the margins; berries some-
what tetragonous, short-hirtellous at the apex.

One of the best marked of Guatemalan species, and the only
local Piper that grows in its peculiar habitat large rocks in the
edges of swift mountain streams. It is one of several plants confined
to this habitat, its associates being Cuphea, Aster, and representatives
of a few other genera of diverse families.

Piper fraguanum Trelease, Journ. Wash. Acad. Sci. 19: 332.
1929 (type from La Fragua, Atlantida, Honduras). P. hispidiseptum
Trelease, loc. cit. (type from Lancetilla Valley near Tela, Honduras).
P. imperspicuibracteum Trelease, loc. cit. (type collected near Tela,
Honduras). P. obsessum Trelease, op. cit. 334. 1929 (type from
Lancetilla Valley, Honduras). P. prodigum Trelease, op. cit. 335.
1929 (type from Lancetilla Valley, Honduras). P. scabriseptum
Trelease, op. cit. 336. 1929 (type from Lancetilla Valley, Honduras).
P. speratum Trelease, loc. cit. (type from Tela, Honduras). P. tes-
seraespicum Trelease in Yuncker, Field Mus. Bot. 17: 354. 1938
(type from Siguatepeque, Honduras). P. puentealtoanum Trelease
in Yuncker, Field Mus. Bot. 9: 280. 1940 (type collected near La
Ceiba, Honduras). P. cocquericotense Trelease in Standl. Field Mus.
Bot. 12: 407. 1936 (type from Little Cocquericot, Belize River,
British Honduras, C. L. Lundell 3829).

Moist or wet thickets or mixed forest, 1,650 meters or lower,
most frequent at rather low elevations; Alta Verapaz; Izabal;
Escuintla; Guatemala; Suchitepequez; San Marcos. Honduras;
British Honduras.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 299

An erect shrub 1.5-2.5 meters high, the branches slender, often flexuous or
somewhat geniculate, densely hispidulous with short, whitish or fulvous, spreading
or somewhat reflexed hairs; petioles 1 cm. long or usually shorter, densely hispi-
dulous, not winged, dilated at the base; leaf blades thin, usually green or dark
green when dried, finely pellucid-punctate, dull, mostly elliptic or ovate-elliptic,
sometimes lance-oblong or lance-ovate, mostly 12-23 cm. long and 5.5-11 cm.
wide, rather narrowly and abruptly long-acuminate, conspicuously unequal and
oblique at the base, cordate to obtuse on one side, rounded to acute on the other,
conspicuously granular on the upper surface and very scabrous, harsh to the touch,
not at all bullate, granular beneath, finely hispidulous with spreading hairs on
the nerves and veins, often also between them, rough to the touch, penninerved,
the nerves 3-5 on each side, slender, prominent, arcuate-ascending at an angle
of about 45 degrees, the veins not conspicuous but often prominulous, laxly
reticulate; peduncles opposite the leaves, stout, 1 cm. long or shorter, hirtellous
or glabrate; spikes spreading or ascending, rather slender, cream-colored or green-
ish, mostly 6-10 cm. long and 3-3.5 mm. thick, obtuse or submucronate; bracts
rounded-peltate at the apex, puberulent or ciliate.

This has been reported as P. jalapense C. DC. from both Guate-
mala and Honduras, where it is a rather common species. The species
of this general alliance are all much alike, and it is probable that
several of those tentatively recognized in this treatment ultimately
will have to be combined.

Piper Friedrichsthalii C. DC. (in DC. Prodr. 16, pt. 1: 327. 1869)
was based on a collection by Friedrichsthal, ascribed by De Candolle
to Guatemala, but the plant was actually from Orosi, Costa Rica.

Piper frioense Standl. & Steyerm., sp. nov.

Known only from the type, Alta Verapaz, wet forest near Tactic,
above the bridge across Rio Frio, 1,500 meters, Standley 90328
(in Herbarium of Chicago Natural History Museum).

A shrub 2 meters high, glabrous throughout, the branches rather slender, with
short internodes; petioles stout, 1-2.5 cm. long, vaginate only near the base;
leaf blades rather thick, brownish when dried, minutely pellucid-punctate or almost
wholly opaque, not at all bullate, ovate or broadly ovate, mostly 10-15 cm. long
and 5-8 cm. wide, abruptly short-acuminate, equal at the base and symmetric
or nearly so, rounded to subacute, penninerved, the nerves 5-6 on each side,
prominent beneath, arcuate-ascending at an angle of about 45 degrees, with a few
stout intermediate nerves, the veins prominent beneath, laxly reticulate; peduncles
opposite the leaves, rather slender, about 1 cm. long; spikes slender, probably
erect, about 6.5 cm. long and 2 mm. thick, subacute; bracts glabrous, ciliolate;
anthers slightly exserted.

Frutex fere omnino glaber; petioli crassi 1-2.5 cm. longi tantum prope basin
vaginantes; lamina crassiuscula in sicco brunnescens minute pellucido-punctata
vel fere omnino opaca, ovata vel late ovata, majuscula, abrupte breviterque

300 FIELDIANA: BOTANY, VOLUME 24

acuminata, basi fere aequali et symmetrica roundata usque subacuta, penninervia,
nervis utroque latere 5-6, aliis minus prominentibus interjectis, venis laxe reticu-
latis; pedunculi oppositifolii, spicis gracilibus ca. 6.5 cm. longis 2 mm. crassis.

Piper geniculatum Swartz, Prodr. Veg. Ind. Occ. 15. 1788.

Dense, wet, mixed, lowland forest, 350 meters or lower; Alta
Verapaz (Cubilgiiitz) ; Izabal (Cerro San Gil). British Honduras,
and probably along much of the Atlantic coast of Central America;
West Indies; South America.

A shrub or small tree of 2-6 meters, sometimes with a trunk 7 cm. in diameter,
glabrous throughout or nearly so; petioles stout, 1-2 cm. long, vaginate through-
out and at least when young broadly winged; leaf blades thin, blackening when
dried, dull, densely and minutely pellucid-punctate, lance-oblong or ovate-oblong,
mostly 12-20 cm. long and 3.5-6.5 cm. wide, gradually or rather abruptly acumi-
nate, usually very unequal at the base with one side often 1 cm. longer than the
other, rounded or obtuse on each side, slightly paler beneath and often black-
puncticulate, penninerved, the nerves usually 5-6 on each side, very slender and
inconspicuous, arcuate-ascending, irregular, scarcely at all elevated, the veins
very obscure and distant; peduncles stout, about 1 cm. long; spikes cream-colored
or greenish, very slender, about 12-13 cm. long and scarcely 3 mm. thick, obtuse;
bracts lunulate-peltate, hirtellous on the margins; fruit glabrous, laterally com-
pressed.

Although one of the Guatemalan collections was designated (but
not published) as a new species by Trelease, the mainland material
seems to agree well enough with the West Indian P. geniculatum.
Called "cordoncillo" in British Honduras.

Piper georginum Trelease & Standley, sp. nov.

Moist or wet, mixed, mountain forest, 2,400-2,850 meters;
endemic; Escuintla; Quezaltenango (type from Fuentes Georginas,
western slope of Volcan de Zunil, Standley 67355; type in Herbarium
of Chicago Natural History Museum); San Marcos.

A shrub of 2-3.5 meters, the branches rather slender, sparsely villous or in
age glabrate, the internodes elongate; petioles stout, 2-3.5 cm. long, densely
villous-pilose with rather long, spreading, brownish hairs, not winged, scarcely
dilated at the base; leaf blades thin, large, fuscescent when dry, very minutely
pellucid-punctate, dull, elliptic, broadly elliptic, or broadly ovate-elliptic, mostly
20-25 cm. long and 10-13 cm. wide, cuspidate-acuminate, acute at the base and
only slightly unequal, villous-pilosulous on the upper surface along the costa,
elsewhere glabrous or nearly so, not paler beneath, villous-pilose beneath on the
nerves and veins, glabrous between them, penninerved, the nerves 3-4 or some-
times 5 on each side, very slender, prominent, arcuate-ascending at an angle of
45 degrees or less, the veins very slender but often conspicuous, very laxly reticu-
late; peduncles opposite the leaves, 13-20 mm. long, villosulous, slender or stout;
spikes erect, at maturity stout, 5.5-6 cm. long, 5 mm. thick at anthesis, bearing a

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 301

tail-like apical mucro 4-6 mm. long; bracts densely pubescent; fruits densely
puberulent about the apex.

Frutex, ramis sparse villosis vel glabratis, internodiis elongatis; petioli 2-3.5
cm. longi dense villoso-pilosi non alati; lamina magna tenuis in sicco fuscescens
minute pellucido-punctata, late elliptica vel ovato-elliptica, cuspidato-acuminata,
basi vix inaequali acuta, supra ad costam villoso-pilosula, subtus ad nervos
venasque villoso-pilosula, penninervia, nervis utroque latere 3-5; pedunculi
oppositifolii 13-20 mm. longi; spicae erectae crassiusculae ca. 6 cm. longae et 5
mm. crassae, apice 4-6 mm. longe mucronatae; bracteae dense pubescentes;
fructus apice dense puberulus.

Trelease assigned a distinct name to each of the four sheets we
refer here, but all seem to be conspecific.

Piper grandilimbum C. DC. Bot. Gaz. 70: 172. 1920.

Moist or wet, mixed forest or thickets, 150-1,150 meters; endemic;
Pete"n; Alta Verapaz (type collected near Cubilgiiitz, Turckheim
11.1400); Huehuetenango (regions of Barillas and Ixcan).

A coarse shrub 1-2.5 meters high, the young branches glabrous; petioles rather
short or much elongate, often vaginate well above the base, glabrous; leaf blades
peltate far above the base or in some of the leaves epeltate, broadly ovate, mostly
18-36 cm. long and 11-22 cm. wide, rather abruptly acuminate, shallowly and
broadly cordate or subcordate at the base, with broadly rounded basal lobes,
glabrous above and pale-granular, the nerves rather prominent, almost con-
colorous beneath, minutely hirtellous or puberulent on the nerves and veins,
palmately nerved at the base with 13 or fewer nerves, penninerved above the
base with 3-4 nerves on each side, these arcuate-ascending at a wide angle;
peduncles hirtellous or glabrate, 2 cm. long or shorter, several times shorter than
the petioles; spikes slender, straight or slightly curved, mostly 11-14 cm. long,
4 mm. thick, the bracts hirtellous on the margins; stamens 3; fruit glabrous,
obpyramidal-globose.

One of the very distinct local species, easily recognized by its
large peltate leaves, which appear glabrous or nearly so to the naked
eye.

Piper guazacapanense Trelease & Standley, sp. nov.

Known only from the vicinity of the type locality, Santa Rosa,
along the Avellana road, south of Guazacapan, on wooded plains
or in quebradas, 150 meters, Standley 79461 (type in Herbarium of
Chicago Natural History Museum).

Plants chiefly herbaceous, much branched, the stems procumbent and rooting
at the nodes, glabrous; stipules lanceolate, about 8 mm. long, free, hirsute; petioles
slender, very unequal, 10-15 mm. long, hirsute; leaf blades thin, drying grayish
green, orbicular-ovate, 5-7.5 cm. long, 3.5-6.5 cm. wide, abruptly acuminate or
long-acuminate, deeply but openly cordate at the base with rounded basal lobes,
epunctate, sometimes obscurely bullate, palmately 5-nerved, with the inner nerves

302 FIELDIANA: BOTANY, VOLUME 24

protracted almost to the apex, deep green on the upper surface, puberulent on
the nerves, paler beneath, sparsely puberulent or hirtellous on the nerves, the
veins almost obsolete, epunctate; inflorescences opposite the leaves, racemose,
often few-flowered, 8-18 mm. long, the rachis short-hirsute, the pedicels short,
stout, divaricate; fruit globose, glabrous, white at maturity, when dry about 2
mm. in diameter.

Herba vel suffrutex, caulibus procumbentibus et radicantibus glabris; stipulae
lanceolatae liberae hirsutae; folia graciliter petiolata, petiolo hirsute, orbiculari-
ovata subito acuminata, basi profunde aperte cordata, palmatim 5-nervia, utrinque
ad nervos sparse puberula vel breviter hirtella; flores racemosi, racemis oppositi-
foliis saepe paucifloris, rhachi breviter hirsuta, pedicellis crassis divaricatis; fructus
glaber.

Referable to Trelease's genus Arctottonia, under which it was
designated as a new species. It is quite different from any other
Central American species of that group but is rather similar to some
of the Mexican ones, although clearly distinct from all of them that
we have seen. The plant is distinctive in habit, being herbaceous
almost throughout, and forming dense colonies of procumbent stems
that take root at many of the nodes. It formed a rather dense ground
cover in the lowland forest where it was collected. The small fruits,
white when ripe, have a rather faint but distinct flavor of black
pepper.

Piper Hermes Trelease & Standley, sp. nov.

Known only from the type, San Marcos, Volcan de Tajumulco,
between Todos Santos and Finca El Porvenir, 1,300 meters or higher,
Steyermark 37009 (type in Herbarium of Chicago Natural History
Museum).

A shrub 3-4.5 meters high, the branches rather stout, with very short inter-
nodes, glabrous or nearly so; petioles about 1.5 cm. long, winged for their whole
length, hirsute dorsally; leaf blades thick, dull, grayish green when dried, opaque,
lance-oblong or narrowly elliptic-oblong, widest at or slightly below the middle,
8-11 cm. long, 3-4 cm. wide, acute or short-acuminate, very unequal and slightly
oblique at the base, one side 5-6 mm. longer than the other, obtuse or rounded
on each side, glabrous above, the costa and nerves sometimes slightly impressed,
brownish beneath when dried, sparsely villous-hirsute on the costa, elsewhere
glabrous, penninerved, the nerves usually 3 on each side, very slender, prominent,
arcuate-ascending at an angle of 45 degrees or slightly more, the veins incon-
spicuous, laxly reticulate; very young spikes (about 1 cm. long) mucronate, borne
on a slender peduncle 7 mm. long, the peduncles sparsely hispidulous or glabrate,
opposite the leaves; bracts glabrous, ciliolate.

Frutex, ramis crassiusculis, internodiis brevibus glabris; petioli tota longi-
tudine alati ca. 1.5 cm. longi dorso hirsuti; lamina crassa non lucida opaca lanceo-
lato-oblonga vel anguste elliptico-oblonga, acuta vel breviter acuminata, basi
solemniter inaequalis et obliqua, utroque latere obtusa vel rotundata, supra glabra,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 303

subtus ad costam villoso-hirsuta, penninervia, nervis utroque latere vulgo 3;
spicae apice mucronatae.

Piper Heydei C. DC. in Bonn. Smith, Bot. Gaz. 19: 7. 1894.
Palo de jute (fide Aguilar) ; Santa Maria.

Dense, wet, mixed, mountain forest, 1,800-2,850 meters; endemic;
Zacapa; El Progreso; Chimaltenango; Quiche" (type from San Miguel
Uspantan, Heyde & Lux 3461); Quezaltenango; San Marcos.

A coarse stout herb 1-2.5 meters high, simple or with few branches, sometimes
suffrutescent below, the branches thick and somewhat succulent, villous-hirsute;
leaves very large and thin, the petiole long and stout, vaginate for most of its
length, 15-20 cm. long or more; leaf blades ovate-rounded, mostly 25-40 cm. long
and nearly or quite as wide, rounded at the apex and abruptly short-acute or at
least pointed, broadly rounded and rather obscurely cordate at the base, deep
green above, sparsely villous, much paler beneath, densely short-villous along the
nerves and veins, peltate far above the base, radiately palmate-nerved at the base,
the costa emitting usually 4 lateral nerves above the basal ones; peduncles rather
slender, erect, mostly 13-17 cm. long, bearing 2-12 spikes, these on secondary
peduncles 3-6 cm. long, erect; spikes green, rather slender, mostly 6-15 cm. long,
in anthesis 4 mm. thick, very obtuse, the bracts densely puberulent; stamens 3,
the anthers subglobose; ovary ovoid, attenuate at the apex, glabrous.

This is a distinct species, related to P. peltatum but very different
in appearance when growing, and readily distinguishable by the
abundant pubescence of the leaves and by the villous-hirsute stems.
We refer here, not without some question, one collection that
Trelease indicated as a new species. It is distinguished by having
the slender secondary peduncles all arising from the axil of the leaf,
and as much as 15 cm. long. In the type of the species the secondary
peduncles are much shorter and are borne at the apex of a long
primary peduncle. Since the plants are exactly alike in their leaves
and pubescence, we suspect that these differences are only individual
variations, especially since both forms have been collected in the
same general region.

Piper imberbe Trelease & Standley, sp. nov.

Known only from the type, Izabal, wet mixed forest, between
Escobas and the waterfall, near sea level, across the bay from
Puerto Barrios, Steyermark 39857 (in Herbarium of Chicago Natural
History Museum).

Stems spreading or pendent, very slender, green, sparsely and obscurely
puberulent when young or practically glabrous, a meter long or shorter, suffrutes-
cent, with elongate internodes; petioles very slender, 4-4.5 cm. long, glabrous;
leaf blades thin, lance-oblong, 17.5-23 cm. long, 5-6.5 cm. wide, attenuate at the
apex into a very long and slender, tail-like acumen, rounded or obscurely sub-

304 FIELDIANA: BOTANY, VOLUME 24

cordate at the base, peltate about 1.5 cm. above the base, palmately few-nerved
about the point of attachment, penninerved above the base, with about 5 nerves
on each side, these very slender, divergent at a wide angle, arcuate, prominent
beneath, scarcely elevated above, the veins very slender and inconspicuous, laxly
reticulate, the blades glabrous on both surfaces; spikes opposite the leaves, the
peduncle very slender, about 1 cm. long, glabrous; spikes very slender, 7.5 cm.
long, scarcely more than 2 mm. thick, the bracts suborbicular, glabrous or practi-
cally so.

Frutex, ramis obscure puberulis vel fere omnino glabris; folia tenues longe
graciliter petiolata, bene supra basin peltata, lanceolato-oblonga, apice anguste
longissime attenuato-acuminata, basi rotundata vel obscure subcordata, basi
palmatinervia, costa supra basin utroque latere nervos 3-4 emittente; spicae
breviter graciliter pedunculatae, elongatae, gracillimae.

Piper indignum Trelease, Journ. Wash. Acad. Sci. 19: 333.
1929. P. micoense Trelease, op. cit. 334 (type from Sierra del Mico,
Izabal, W. A. Kellerman 6715).

Moist or rather dry thickets or forest, 1,400 meters or lower;
Izabal ; Jutiapa ; Santa Rosa. Honduras, the type from Siguatepeque.

A stout erect shrub 1.5-2 meters high, the branches very densely short-hirsute
with spreading whitish hairs, the internodes usually short; petioles stout, mostly
7-10 mm. long, not winged, somewhat dilated at the base, very densely short-
hirsute; leaf blades rather thick, pale grayish green or somewhat fulvescent when
dry, minutely pellucid-punctate, broadly elliptic to ovate-elliptic or obovate-
elliptic, mostly 10-16 cm. long and 4-8 cm. wide, rather abruptly acute to long-
acuminate, conspicuously oblique and unequal at the base, usually cordate or
subcordate on one side and rounded or obtuse on the other, very densely and rather
softly pilose on the upper surface with short, spreading or subappressed, fulvescent
or whitish hairs, not at all bullate, or only slightly so, densely and softly pilose
beneath between as well as upon the nerves and veins, penninerved, the nerves
3-4 on each side, slender, prominent, ascending at an angle of about 45 degrees,
slightly arcuate or almost straight, the veins often prominent, sometimes rather
closely reticulate; peduncles opposite the leaves, stout, about 1 cm. long or some-
what shorter, densely hirtellous; spikes erect or ascending, commonly 3.5-5.5 cm.
long, short-mucronate, 2.5 mm. thick; bracts triangular-peltate at the apex,
densely fulvescent-puberulent; fruits trigonous, glabrous, slightly depressed at
the apex.

Piper instabilipes Trelease in Standl. Field Mus. Bot. 17: 232.
1937.

Known only from the type, Pete"n, high upland forest near Santa
Teresa, Rio Subin, C. L. Lundell 2776.

A rather slender shrub, glabrous throughout, the internodes short or elongate;
petioles slender, 1-1.5 cm. long, vaginate only at the base; leaf blades thin, darken-
ing when dried, oblong-ovate to elliptic-oblong, mostly 12-16 cm. long and 4.5-
7.5 cm. wide, rather abruptly caudate-acuminate, rounded to subtruncate at the

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 305

base, the two sides equal or nearly so, penninerved, the nerves 4-6 on each side,
very slender, prominulous on both surfaces, arcuate, ascending at an angle of 45
degrees or more, the veins prominulous but not very conspicuous beneath, laxly
reticulate, the blades epunctate or nearly so, dull when dried, concolorous;
peduncles 5 mm. long or more, shorter than the petioles; young spikes about 8 cm.
long and 2 mm. thick, obtuse, slender, the bracts rounded-subpeltate, ciliolate;
ovary glabrous; stigmas 3, sessile.

Piper ixocubvainense Standl. & Steyerm., sp. nov.

Dense, wet, mixed forest, 300-1,600 meters; endemic; Alta Vera-
paz (type from Montana Ixocubvain, 2.5 miles west of Cubilgiiitz,
'Steyermark 44987; in Herbarium of Chicago Natural History Mu-
seum; collected also in the Tactic region).

A rather densely branched shrub 1-1.5 meters high, glabrous throughout, the
upper internodes short, the lower ones elongate; petioles slender or stout, mostly
2 cm. long or slightly shorter, vaginate for all or most of their length, narrowly
winged; leaf blades drying dark green but scarcely fuscescent, rather thick,
minutely pellucid-punctate, dull, elliptic-oblong, mostly 9.5-12 cm. long and
3.5-5.5 cm. wide, acute or short-acuminate, slightly unequal and somewhat
oblique at the base, narrowly rounded or shallowly cordate or subcordate, penni-
nerved, the nerves mostly 5 on each side, sometimes only 4 on one side, arcuate-
ascending at an angle of slightly more than 45 degrees, very slender, prominulous
beneath, the veins obscure, very laxly reticulate; spikes opposite the leaves, small,
on peduncles 3-4 mm. long, rather stout, at anthesis 1.5-2.5 cm. long, about 2.5
mm. thick, log-mucronate at the apex; bracts small, glabrous, ciliate; anthers
slightly exserted.

Frutex erectus glaber; folia ca. 2 cm. longe petiolata crassiuscula in sicco
intense viridia vix fuscescentia minute pellucido-punctata, elliptico-oblonga, acuta
vel breviter acuminata, basi paullo inaequali et plus minusve obliqua anguste
rotundata vel breviter cordata, penninervia, nervis saepius utroque latere 5
tenerrimis subtus prominulis, arcuato-adscendentibus, venis obscuris laxissime
reticulatis; spicae oppositifoliae parvae ad anthesin 1.5-2.5 cm. tantum longae
longe apice mucronatae.

Piper jactatum Trelease & Standley, sp. nov.

Dense, wet, mixed, mountain forest, sometimes in wooded swamps,
1,400-1,600 meters; endemic; Alta Verapaz (type collected in moun-
tains along the road between Tactic and the divide on the road to
Tamahu, Standley 90614; in Herbarium of Chicago Natural History
Museum; known only from the Tactic region).

A sparsely branched, erect shrub about 2 meters high, the branches very
densely hispidulous with short spreading sordid hairs, the upper internodes short;
petioles stout, about 8 mm. long, densely hispidulous, not winged, somewhat
dilated at the base; leaf blades fuscous when dried, thick, not or scarcely lustrous,
densely and minutely pellucid-punctate, lanceolate or oblong-lanceolate, mostly
12-18 cm. long and 4.5-6 cm. wide, very narrowly and gradually long-attenuate,

306 FIELDIANA: BOTANY, VOLUME 24

slightly unequal and oblique at the base or sometimes subequal, rounded or usually
somewhat cordate on each side, conspicuously bullate, hispidulous above along
the costa and nerves but elsewhere glabrous or nearly so and smooth to the touch,
the nerves and veins more or less impressed, brownish beneath, densely and rather
softly hispidulous or hirtellous almost throughout, penninerved, the nerves mostly
3-4 on each side, slender but very prominent, arcuate-ascending at a narrow angle,
the veins very prominent and rather stout, openly reticulate; peduncles opposite
the leaves, about 1 cm. long, stout, densely hispidulous; spikes (perfect ones
not seen) rather stout, spreading or erect-spreading, about 5.5 cm. long and 3 mm.
thick; bracts densely pubescent; fruits depressed at the apex, puberulent above.

Kami pilis brevibus patentibus sordidis dense hispiduli; petioli ca. 8 mm.
longi non alati dense hispiduli; lamina in sicco fusca crassa non lucida dense,
minute pellucido-punctata, lanceolata vel oblongo-lanceolata modica, sensim
longeque attenuata, basi paullo inaequali et obliqua vel fere symmetrica utroque
latere plus minusve cordata, insigniter bullata, supra ad costam nervosque hispi-
dula, aliter glabra vel fere glabra, subtus ubique dense molliter hispidula vel
hirtella, penninervia, nervis utroque latere 3-4 angulo angusto arcuato-adscen-
dentibus elevatis, venis prominentibus conspicue reticulatis, supra saepe impressis;
pedunculi oppositifolii ca. 1 cm. longi dense hispiduli, spicis (perfectis non visis)
crassiusculis ca. 5.5 cm. longis et 3 mm. crassis; bracteae dense pubescentes;
fructus apice depressus superne puberulus.

Piper jumayense Trelease & Standley, sp. nov.

Known only from the type, Jalapa, south-facing slopes of bar-
ranco, Volcan de Jumay, 2,000 meters, Steyermark 32449 (Herbarium
of Chicago Natural History Museum).

A slender shrub 3 meters high, glabrous throughout or nearly so, the internodes
mostly elongate; petioles slender, 5-6 mm. long, vaginate only at the base; leaf
blades thin but rather stiff, minutely pellucid-punctate, drying somewhat blackish,
ovate or elliptic-ovate or lance-ovate, mostly 8-10 cm. long and 4-5.5 cm. wide,
gradually or somewhat abruptly long-acuminate, obliquely rounded or very obtuse
at the base, somewhat paler beneath, palmately 5-nerved, the nerves very slender
but prominent beneath, the outermost somewhat irregular but conspicuous, the
veins pale, obscure or prominent, laxly reticulate; peduncles almost filiform,
2.5 cm. long, sparsely puberulent or glabrate, rather laxly flowered, the rachis
densely puberulent; spikes about 2.5 cm. long, in fruit fully 4 mm. broad; fruits
oval, 2 mm. long, minutely puberulent or almost glabrate; stigmas 3, large, sessile.

Frutex gracilis fere omnino glaber; folia graciliter petiolata, petiolo basi
tantum vaginante, in sicco nigrescentia, minute pellucido-punctata, ovata usque
elliptico-ovata vel lanceolato-ovata, modica, sensim vel subabrupte longiacuminata,
basi oblique rotundata vel obtusissima, palmatim 5-nervia, venis inconspicuis vel
prominentibus laxe reticulatis; pedunculi fere filiformes 2.5 cm. longi, puberuli
vel glabrati, rhachi dense puberula; spicae sublaxiflorae breves; fructus ovalis
minute puberula vel glabrata; stigmata 3 sessilia.

This may be only a form of P. Amalago, but the collection is
noteworthy for the very long and slender peduncles.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 307

Piper lanciferum Standl. & Steyerm., sp. nov.

Known only from the type, Suchitepequez, Volcan de Santa
Clara, between Finca El Naranjo and the upper slopes, 1,300-2,600
meters, Steyermark 46696 (type in Herbarium of Chicago Natural
History Museum).

A shrub 3 meters high, the branches rather stout, with very short upper inter-
nodes, glabrous or at first sparsely short-hispidulous; petioles rather stout, mostly
8-14 mm. long, sparsely hispidulous or glabrate, not winged; leaf blades lanceolate
or elliptic-lanceolate, broadest near the middle, mostly 10-14 cm. long and 3-4.5
cm. wide, narrowly long-acuminate, gradually attenuate to the base, somewhat
unequal and oblique at the base, very acute on both sides, fuscescent when dried,
rather thick, minutely pellucid-punctate or almost wholly opaque, dull on the
upper surface, glabrous, sometimes minutely granular, somewhat brownish
beneath, villous-pilosulous on the nerves and veins with rather long, soft, spread-
ing hairs, penninerved, the nerves 3-4 on each side, slightly arcuate, ascending
at an angle of less than 45 degrees, the veins somewhat prominent but not very
conspicuous, laxly reticulate; peduncles opposite the leaves, stout, glabrous, about
8 mm. long; spikes erect-ascending, stout, mostly 7-8 cm. long and 4-5 mm.
thick; bracts triangular-peltate at the apex, densely pale-pubescent on the margins;
fruits small, subglobose, glabrous.

Frutex, ramis crassiusculis, internodiis brevibus glabris vel primo sparse
breviter hispidulis; folia modica crassiuscula in si ceo fuscescentia minute pellucido-
punctata vel fere omnino opaca, non lucida, lanceolata vel anguste elliptico-
lanceolata, anguste longiacuminata, basi inaequali et obliqua utroque latere acuta,
supra glabra, subtus in sicco brunnescentia, ad nervos venasque pilis longiusculis
mollibus villoso-pilosula, penninervia, nervis utroque latere 3-4 angulo angusto
arcuato-adscendentibus; pedunculi oppositifolii crassi glabri ca. 8 mm. longi,
spicis erecto-patentibus plerumque 7-8 cm. longis, 4-5 mm. crassis.

Piper Luxii C. DC. in Donn. Smith, Bot. Gaz. 19: 5. 1894.

Moist or wet, mixed forest, 1,800-2,400 meters; endemic; Chimal-
tenango; Quich6 (type from San Miguel Uspantan, Heyde & Lux
3462).

A rather stout, erect shrub 1.5-2.5 meters high, the young branches densely
short-villous with brownish or sordid, spreading hairs, the internodes short or
elongate, often glabrate in age; petioles rather stout, mostly 2-4 cm. long, brownish-
villous, vaginate at the base; leaf blades large, lance-elliptic to elliptic or obovate-
elliptic, mostly 15-20 cm. long and 9-10.5 cm. wide, abruptly short-acuminate,
often broadly narrowed to the base and rounded or obtuse, sometimes narrowly
subcordate, opaque, often somewhat bullate, villous-hirsute above when young
but in age merely scabrous and rough to the touch, rather densely villosulous
beneath, especially on the nerves and veins, blackish when dry, penninerved, the
nerves 3-5 on each side, arising below the middle of the blade, ascending at a very
narrow angle, the veins prominent beneath, laxly or closely reticulate, often deeply
impressed on the upper surface; peduncles erect, villosulous, sometimes 2.5 cm.
long but generally shorter; spikes stout, mucronate at the apex, 5-6 cm. long,
4-5 mm. thick; bracts obovate-oblong, rounded at the apex, densely villosulous;

308 FIELDIANA: BOTANY, VOLUME 24

stamens 4, the filaments exserted, the anthers elliptic; stigmas 3; ovary sparsely
pilosulous.

Piper Martensianum C. DC. in DC. Prodr. 16, pt. 1: 251.
1869. P. santarosanum C. DC. ex Dorm. Smith, Enum. PI. Guat.
2: 96. 1891, nomen; in Donn. Smith, Bot. Gaz. 19: 258. 1894 (type
from Santa Rosa, Baja Verapaz, Tilrckheim 1174.) ?P. Savagii
C. DC. Ann. Cons. Jard. Bot. Geneve 2: 252. 1898 (type said to
have been collected in Guatemala by Savage). P. andresense
Trelease in Standl. Field Mus. Bot. 17: 230. 1937 (type from San
Andre's, Lago de Pete"n, Pete"n, C. L. Lundell 3203). P. andresense
var. subclavescens Trelease in Standl. op. cit. 231 (type from Yaxha,
Pete"n, Lundell 2011). P. rematense Trelease in Standl. op. cit. 233
(type from Remate, Lago de Pete"n, Lundell 2087). P. simulhabitans
Trelease in Standl. op. cit. 233 (type from Remate, Pete"n, Lundell
2073). P. yaxhanum Trelease in Standl. op. cit. 234 (type from
Yaxha-Remate road, Pete"n, Lundell 3286). Cordoncillo.

Wet to dry thickets or forest, usually in open forest, sometimes
in pine-oak forest, ascending from sea level to about 2,300 meters,
most common at middle elevations; Pete"n; Alta Verapaz; Baja
Verapaz; Zacapa; Chiquimula; Jalapa; Jutiapa; Santa Rosa; Es-
cuintla; Guatemala; Sacatepe"quez; Chimaltenango; Suchitepequez;
Retalhuleu; Quezaltenango; Huehuetenango; Quiche". Southern
Mexico; El Salvador and probably farther southward.

An openly branched shrub or rarely a small tree, 1.5-6 meters high, the
branches slender, densely hirtellous, at least when young; leaves on hirtellous
petioles 1 cm. long or often slightly longer, rather thin and flaccid, very minutely
pellucid-punctate, rounded-ovate to oblong-ovate, mostly 10-15 cm. long and
5-14 cm. wide, abruptly acute or acuminate, at least the larger leaves cordate
at the base, the upper ones often merely rounded, slightly or conspicuously oblique,
deep green above and more or less puberulent, somewhat paler beneath, usually
densely velutinous-pubescent, often with yellowish hairs, palmately 5-7-nerved,
the veins very slender, laxly reticulate; peduncles opposite the leaves, densely
hirtellous, 1-2 cm. long; spikes whitish or pale green, slender, variable in length,
mostly 7-10 cm. long, 3 mm. thick; bracts spatulate, rounded at the apex, densely
hirtellous; ovary hirtellous; stamens 4, the anthers reniform-globose; fruit small,
globose-ovoid, hirtellous at the apex.

Sometimes called "candelillo" in El Salvador. This is one of the
most common species of Piper in Guatemala. It grows in some-
what drier and less shaded habitats than most species of the genus.
The plant is so common and widely distributed that some older
name than P. Martensianum may eventually be found for it among
the names based on Mexican collections. A synonym here is

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 309

P. decrescens (Miq.) C. DC. var. ovatum C. DC. (in DC. Prodr. 16,
pt. 1: 251. 1869), reported as Guatemalan but actually from
Nicaragua.

Piper minarum Standl. & Steyerm., sp. nov.

Known only from the type, Zacapa, Sierra de las Minas, slopes
of Monte Virgen, 2,200-2,400 meters, Steyermark 42667 (in Her-
barium of Chicago Natural History Museum).

A shrub of 3 meters, the branches slender, with short internodes, when young
densely villous-pilosulous with soft, spreading, fulvescent or sordid hairs; petioles
stout, 1.5 cm. long or shorter, densely villous-pilosulous, vaginate only at the
base; leaf blades thin, fuscous when dried, elliptic or obovate-elliptic, about 21
cm. long and 9-10 cm. wide, rather long-acuminate, somewhat narrowed to the
somewhat oblique and unequal base, subcordate or rounded on one side, obtuse
on the other, villosulous-pilosulous above along the costa and when young often
sparsely pilosulous, in age glabrous or nearly so, slightly lustrous, not rugose,
slightly paler beneath, villous-pilose with soft spreading hairs along the costa,
very shortly pilosulous on the veins, glabrous or nearly so between them, penni-
nerved, the nerves usually 4 on each side, arcuate-ascending at an angle of 45
degrees or less, the veins prominulous, laxly reticulate; spikes (only very young
ones seen) borne on stout villous-pilosulous peduncles 6-7 mm. long, stout, about
3.5 cm. long and 4 mm. thick, erect, rounded at the apex, the bracts densely sordid-
pubescent.

Rami graciles dense villoso-pilosuli, internodiis brevibus; petioli crassiusculi
1.5 cm. longi vel breviores dense villoso-pilosuli basi tan turn vaginantes; lamina
tenuis in sicco fuscescens minutissime pellucido-punctata, elliptica vel obovato-
elliptica, majuscula, longiuscule acuminata, basin versus aliquanto angustata, basi
ipsa inaequali atque obliqua uno latere subcordata vel rotundata, altero obtusa,
supra ad costam villoso-pilosula, non bullata, subtus ad costam pilis longis patenti-
bus villoso-pilosa, ad venas minute pilosula, inter venas glabra vel glabrata, penni-
nervia, nervis utroque latere vulgo 4 angulo angusto arcuato-adscendentibus;
spicae (juveniles tantum visae) crassae apice rotundatae, bracteis dense sordido-
pubescentibus.

Piper minutantherum C. DC. Bot. Gaz. 70: 171. 1920.

Known only from the type, Alta Verapaz, Cubilgiiitz, 350
meters, Turckheim 11.1441.

Branchlets glabrous, minutely verruculose; leaves glabrous, on petioles 2 cm.
long, membranaceous when dried, minutely pellucid-punctate, elliptic-lanceolate,
as much as 21 cm. long and 7.7 cm. wide, rather long-acuminate, slightly unequal
at the base and acute on each side, the costa emitting 4-5 lateral nerves on each
side; peduncles glabrous, shorter than the petioles, as much as 13 mm. long;
spikes about 10 cm. long and almost 4 mm. thick, short-acute at the apex, the
rachis glabrous; bracts hirtellous; stamens 3; fruits obovoid, glabrous, 1.5 mm.
long, narrowed to the style.

We have seen no material of this species, which is not included
in the key.

310 FIELDIANA: BOTANY, VOLUME 24

Piper misantlense C. DC. in DC. Prodr. 16, pt. 1: 286. 1869.

Moist forest or in second growth thickets, 1,100-1,600 meters;
Quich^ ; Huehuetenango. Southern Mexico.

A shrub about 3 meters high, the branches rather slender, glabrous, with
short or somewhat elongate internodes; petioles slender, 6-15 mm. long, glabrous,
not winged, dilated and sheathing at the base; leaf blades rather thick and firm,
green when dried, finely pellucid-punctate, elliptic-oblong to elliptic or obovate-
elliptic, mostly broadest at or above the middle, 11-14 cm. long, 4-7.5 cm. wide,
abruptly acuminate, oblique and somewhat unequal at the narrow base, obtuse
on each side, green above, glabrous, slightly paler beneath, puberulent or strigil-
lose on the nerves and veins, glabrous between them, penninerved, the nerves
3-4 on each side, slender, prominent, arcuate-ascending at an angle of about 45
degrees, the veins very slender, usually inconspicuous, very laxly reticulate;
peduncles opposite the leaves, about 1 cm. long or somewhat shorter, glabrous;
spikes erect or spreading, slender, 5.5-10.5 cm. long, at anthesis little more than
2 mm. thick, obtuse; bracts triangular-peltate at the apex, ciliolate; fruit glabrous.

One of the two Guatemalan collections was marked by Trelease
as a new species, but it agrees perfectly with specimens from Mexico
that he determined as P. misantlense.

Piper oblique-ovatum Trelease in Standl. Field Mus. Bot. 22:
10. 1940. Cordoncillo.

Moist or wet, mixed forest or thickets, 1,250 meters or lower;
endemic; Alta Verapaz; Izabal; Santa Rosa; Escuintla; Sacatepe"-
quez; Chimaltenango (type from Conception, J. R. Johnston 557);
San Marcos.

A branched erect shrub 1-2.5 meters high, the young branches densely and
very minutely puberulent, slender, with short or elongate internodes; petioles
stout, 1 cm. long or usually shorter, puberulent, narrowly vaginate below; leaf
blades thick and firm, usually grayish when dried, epunctate, the veins translucent
against strong light, oblong-ovate to lance-ovate or broadly ovate, mostly 8-12 cm.
long and 3-6 cm. wide, acute or acuminate, very unequal at the base, the shorter
side rounded or subcordate, the longer side deeply cordate, often equaling and
concealing the petiole, glabrous on the upper surface, slightly lustrous, some-
what paler beneath, finely puberulent on the veins, palmately 5-nerved, the
nerves slender but prominent, the veins slender, not very conspicuous, laxly
reticulate, the blades often slightly bullate in age; peduncles slender, mostly 1.5-2
cm. long, minutely puberulent; spikes slightly curved, 5-6 cm. long, 2.5 mm. thick,
obtuse, the bracts glabrous except on the margins.

This is closely related to P. Lindenii (Miq.) C. DC. of Tabasco
and may not be distinct, although, to judge from a photograph of
the type of that species, it has much more slender flower spikes, and
somewhat different leaf venation. We place here three forms that
have been indicated by Trelease as new species, but most of the
specimens we have referred here are sterile, and it is possible that

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 311

fertile specimens will show that more than one species really is
represented by the dozen collections now at hand.

Piper oradendron Trelease & Standley, sp. nov. Cordoncillo.

Wet to dry thickets or mixed forest, 1,200 meters or lower; so
far as known, endemic; Izabal; Santa Rosa; Escuintla (type col-
lected below Las Lajas, Standley 64817; in Herbarium of Chicago
Natural History Museum); Guatemala; Sacatepe*quez; Retalhuleu;
San Marcos.

A shrub 1-2.5 meters high, the branches slender, densely hispidulous with
short, spreading or reflexed, usually fulvous hairs, the upper internodes short;
petioles slender, 1-2 cm. long, not winged, hispidulous, dilated at the base; leaf
blades thin, usually green or dark green when dry, densely and minutely pellucid-
punctate, not or very slightly lustrous, ovate or ovate-elliptic, mostly 13-18 cm.
long and 6-9 cm. wide, abruptly acuminate or long-acuminate, oblique and con-
spicuously unequal at the base, usually acute on one side and obtuse or even
rounded on the other, not bullate, scabrous or hirtellous above along the costa,
elsewhere glabrous or nearly so, usually quite smooth to the touch, sparsely
strigillose beneath on the nerves and veins or in age glabrate, smooth to the touch,
penninerved, the nerves 3-4 on each side, ascending at an angle of usually less than
45 degrees, slightly arcuate or almost straight, very slender, prominent, the veins
prominent, usually pale, laxly reticulate; peduncles opposite the leaves, rather
stout, about 6 mm. long, hispidulous or glabrate; spikes slender, the immature
ones 5-6 cm. long and 2 mm. thick, obtuse; bracts densely pubescent.

Frutex, ramis gracilibus dense pilis brevibus patentibus vel reflexis fulvis
hispidulis; petioli 1-2 cm. longi exalati hispiduli basi tantum dilalati; lamina in
sicco viridis vel obscure viridis tenuis dense minuteque pellucido-punctata ovata
vel ovato-elliptica modica, abrupte acuminata vel longiacuminata, basi inaequali
et obliqua uno latere acuta, altero obtusa vel rotundata, supra ad costam hirtella
vel scabra, tactu laevis, subtus ad costam venasque strigillosa vel glabrata, penni-
nervia, nervis utroque latere 3-4 anguste adscendentibus subarcuatis vel fere
rectis, venis manifestis laxe reticulatis; spicae bene evolutae non visae, juveniles
5-6 mm. longae 2 mm. crassae erectae vel paten tes; bracteae dense pubescentes.

A common shrub of the foothills and plains of the Pacific slope.
It may not be fully distinct from some of the other forms here
treated as species, and probably an older name will be found for
it among the described Mexican species.

Piper pansamalanum C. DC. Bot. Gaz. 70: 177. 1920. P.
mayanum Lundell, Phytologia 1: 337. 1939 (type from Stann Creek
District, Middlesex, British Honduras, P. H. Gentle 2866).

Dense, moist or wet, mixed, mountain forest, 150-1,500 meters;
Alta Verapaz (type from Pansamala, Turckheim 940); Izabal;
Huehuetenango. British Honduras.

312 FIELDIANA: BOTANY, VOLUME 24

A coarse shrub or a small tree, 2.5-6 meters high, with few stout branches,
the young branches densely short- villous with sordid multicellular hairs; petioles
stout, 6 cm. long or shorter, vaginate throughout or nearly so, villosulous; leaf
blades very large, rather thick, blackening when dried, oval-ovate to somewhat
rhombic-ovate, mostly 25-40 cm. long and 17-26 cm. wide or even larger, abruptly
short-acuminate, deeply and narrowly cordate at the base, with large, rounded,
often overlapping basal lobes, glabrous above, densely sordid-villosulous beneath
at least on the nerves and veins and often over the whole surface, palmately about
7-nerved at the base, the costa very stout, with about 3 nerves on each side above
the basal ones, the nerves scarcely elevated on the upper surface but very coarse
and prominent beneath, the veins laxly reticulate, the larger ones elevated;
peduncles villosulous or glabrate, in the type 5.5 cm. long but sometimes as much
as 10 cm.; spikes nutant, brick-red, as much as 40 cm. long or even longer and
8 mm. or less in diameter, equaling or longer than the leaves; bracts triangular-
peltate, hirtellous on the margins; stamens 4, adnate to the base of the fruit;
stigmas 3, sessile; fruit somewhat tetragonous, glabrous.

Easy of recognition because of the much elongate spikes and the
very large, deeply cordate leaves, abundantly pubescent beneath
but glabrous on the upper surface. Piper species of this type are
rare in Central America, but they are more plentiful in Panama
and Costa Rica than in Guatemala. The plant has been reported
from Guatemala under the name Piper gigantifolium var. cubil-
quitzanum C. DC.

Piper patulum Bertol. Fl. Guat. 407. pi. 36. 1840. P. quiri-
guanum Trelease in Standl. Field Mus. Bot. 10: 160. 1931 (type
from Quirigua, Izabal). Cordoncillo.

Moist to wet or rather dry, mountain or lowland forest or thickets,
1,200 meters or lower; Alta Verapaz; Jutiapa; Santa Rosa; Escuintla;
Chimaltenango; Suchitepequez; Retalhuleu; Quezaltenango. South-
ern Mexico; British Honduras; Salvador; Honduras; perhaps even
farther southward.

A rather slender shrub, commonly 2-3 meters high, erect, rather sparsely
branched, glabrous throughout or practically so; petioles slender, mostly 2.5-7
cm. long, vaginate for half their length or more, often almost throughout; leaf
blades thin and when fresh often flaccid, finely and densely pellucid-punctate,
broadly cordate-ovate to rounded-cordate, commonly 10-20 cm. long and 7.5-
15 cm. wide, rather abruptly acuminate or long-acuminate, shallowly or rather
deeply and openly cordate at the base, with rounded basal lobes, deep green above,
somewhat paler beneath, palmately about 9-nerved, the nerves slender, prominent
beneath, the veins rather prominent, laxly reticulate; peduncles opposite the leaves,
rather stout, about 1 cm. long or less; spikes very long and slender, often or usually
somewhat curved, about 12-16 cm. long and 2.5-3 mm. thick, the bracts densely
pilosulous or tomentulose.

A very common plant in many regions, especially on the Pacific
plains and foothills. It is one of the well-marked and easily recog-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 313

nized local species. This is presumably the Piper that has been
reported from Guatemala as P. marginatum Jacq.

Piper patzulinum Trelease & Standley, sp. nov. Cordoncillo.

Moist or wet, mixed, usually dense, mountain forest, sometimes
on white-sand slopes, 650-2,600 meters; Chimaltenango; Retalhuleu;
Quezaltenango (type from lower southern slopes of Volcan de Santa
Maria, along the barranco between Finca Pirineos and Patzulin,
Steyermark 33629; in Herbarium of Chicago Natural History Mu-
seum); San Marcos.

Usually an erect shrub of 1.5-2.5 meters, the branches slender, thinly villo-
sulous or glabrate, the internodes mostly short; petioles rather slender, mostly
1-2 cm. long, villosulous or glabrate, naked or nearly so, slightly dilated at the
base; leaf blades thin and often flaccid, drying green or blackish green, minutely
pellucid-punctate or opaque, narrowly elliptic to elliptic-oblong or ovate-oblong,
mostly 15-20 cm. long and 4.5-8.5 cm. wide, rather abruptly long-acuminate,
conspicuously unequal and oblique at the base, usually cordate or subcordate on
one side and obtuse or acute on the other, or sometimes rounded or obtuse on both
sides, green and glabrous above, somewhat paler beneath, villosulous or pilosulous
with soft spreading hairs on the nerves and veins, glabrous or nearly so between
them, penninerved, the nerves 3-4 on each side, arcuate-ascending at an angle
of 45 degrees or slightly more, very slender, prominent, the veins very slender,
little elevated, often inconspicuous, very laxly reticulate; peduncles opposite the
leaves, rather stout, about 8 mm. long, glabrous or nearly so; spikes erect, white
or greenish white, slender, mostly 8-9 cm. long and 2.5-3 mm. thick, obtuse;
bracts very densely pubescent, pale.

Rami graciles sparse villosuli vel glabrati; petioli vulgo 1-2 cm. longi villosuli
vel glabrati non alati ; lamina tenuis et saepe flaccida in sicco viridis vel fuscescens,
minute pellucido-punctata vel opaca, anguste elliptica usque elliptico-oblonga vel
ovato-oblonga modica, subabrupte longiacuminata, basi manifeste inaequali et
obliqua uno latere cordata vel subcordata, altero obtusa vel acuta, supra glabra,
subtus paullo pallidior ad nervos venasque pilis mollibus patentibus villosula vel
pilosula, penninervia, nervis utroque latere 3-4; pedunculi oppositifolii ca. 8 mm.
longi glabri vel glabrati, spicis erectis gracilibus vulgo 8-9 cm. longis et 2.5-3 mm.
crassis.

Piper peltatum L. Sp. PI. 30. 1753. Heckeria peltata Kunth,
Linnaea 13: 565. 1839. Pothomorphe peltata Miq. Comm. Phyt. 37.
1840. P. peltata var. hypoleuca Trelease in Standl. Field Mus. Bot.
12: 408. 1936 (type from Craig Point, Sibun River, British Honduras,
P. H. Gentle 1400). Santa Maria.

Moist or wet, lowland forest or thickets, sometimes in second
growth, 600 meters or lower; Alta Verapaz; Izabal; Santa Rosa;
Suchitepequez; Retalhuleu; San Marcos. Southern Mexico, south-
ward to Panama; West Indies.

314 FIELDIANA: BOTANY, VOLUME 24

Plants erect, rather stout, herbaceous throughout or nearly so, generally about
1.5 meters high, sparsely branched, the stems minutely puberulent or almost
wholly glabrous, succulent; leaves large, long-petiolate, the petioles broad, long-
vaginate; leaf blades very thin, ovate-orbicular or almost orbicular, mostly 20-30
cm. long, sometimes broader than long, abruptly acute at the rounded apex,
shallowly cordate at the base or rounded, peltate far above the base, sparsely
and minutely puberulent or almost glabrous, deep green above, paler and puncticu-
late beneath, palmate-nerved from the point of insertion, the principal nerve with
usually 2 divergent nerves arising on each side above the base; common peduncle
shorter than the petiole, 1-7 cm. long, glabrous or sparsely puberulent, bearing
4-10 spikes, these on peduncles 7-12 mm. long; spikes pale green, mostly 8-9 cm.
long, 3.5 mm. thick, very obtuse, the bracts puberulent.

Sometimes called "ombligo" in Honduras. In the Botany of
Porto Rico and the Virgin Islands, Britton and Wilson treat this
and P. umbellatum as conspecific, whether on the basis of any
authority other than that of the authors we do not know. It is
conceivable that P. peltatum is only a form of P. umbellatum, but
this we are inclined to doubt. In Central America it is very rare
indeed to find the two growing together. Usually in any one part
of the lowlands where there is a suitable habitat one or the other
may be found, but almost never both in the same general region.
P. peltatum is much rarer than P. umbellatum. Among the Guate-
malan collections, Trelease has indicated two new species under the
genus Pothomorphe. One of the specimens consists of very frag-
mentary material unsuited for description, and both the forms
designated as new seem to differ in no respect from P. peltatum.
This species has been reported from Guatemala as Piper cordil-
lerianum C. DC.

Piper pergamentifolium Trelease & Standley, sp. nov. Cor-
doncillo.

Moist or wet, mixed, mountain forest, sometimes in second growth
thickets, 600-2,200 meters; so far as known, endemic, but to be
expected in southern Mexico; Alta Verapaz; Santa Rosa; Escuintla;
Sacatepe*quez; Chimaltenango; Solola; Retalhuleu; Quezaltenango
(type from Volcan de Zunil, probably near Santa Maria de Jesus,
A. F. Skutch 876; in Herbarium of Chicago Natural History Mu-
seum); San Marcos.

Commonly a shrub of 2-4.5 meters, glabrous throughout, often much branched,
the branches slender, with mostly short internodes; petioles slender or stout,
commonly 1-1.5 cm. long, vaginate below; leaf blades thin but rather stiff and
firm, grayish green when dried, minutely pellucid-punctate, variable in size and
somewhat variable in shape, mostly lanceolate or ovate-lanceolate, sometimes
rhombic-lanceolate, mostly 11-20 cm. long and 3.5-8.5 cm. wide but sometimes

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 315

larger or smaller, gradually or abruptly acuminate or attenuate-acuminate, equal
or nearly so at the base but generally very oblique, one of the sides acute, the
other obtuse to shallowly cordate, the two sides of the leaf thus very unequal in
breadth and shape, penninerved, the nerves generally about five on each side, very
islender, little elevated on either surface and often very inconspicuous, ascending
at a wide or narrow angle, the veins very inconspicuous, laxly reticulate; peduncles
Istout or slender, equaling or often shorter than the petioles, commonly 1 cm. long
!or less; spikes erect or spreading, rather stout, most of them 4-7 cm. long and
about 3 mm. thick, obtuse or subacute at the apex, whitish or pale green; anthers
short-exserted.

Frutex ramosus glaber; folia modica in sicco griseo-viridia minute pellucido-
punctata vulgo 1-1.5 cm. longe petiolata, plerumque lanceolata vel ovato-lanceo-
lata, sensim vel abrupte acuminata vel attenuato-acuminata, basi vulgo insigniter
obliqua subaequalia, uno latere acuta, altero obtusa usque subcordata, penninervia,
nervis utroque latere ca. 5 tenerrimis vix elevatis, venis inconspicuis laxe reticu-
latis; pedunculi petioles aequantes vel breviores, spicis erectis vel patentibus
crassiusculis plerumque 4-7 cm. longis atque 3 mm. crassis.

A very common shrub in the forests of the Pacific foothills and
at middle elevations in the mountains. The numerous collections
are variable in shape and size of the leaves, but we believe that all
or most of them are conspecific, and there is no evident good basis
for separating any of the forms as distinct species. Among the sheets
that he examined, Trelease designated nine or more additional
species, but some of these are clearly conspecific, and the rest of
them are probably so.

Piper perlongipedunculum Trelease & Standley, sp. nov.
Cordoncillo.

Dense moist mixed mountain forest, 1,300-1,500 meters; endemic;
Quezaltenango (type from lower southern slopes of Volcan de Santa
i Maria, along the barranco between Finca Pirineos and Patzulin,
\Steyermark 33628; in Herbarium of Chicago Natural History Mu-
seum; known also from two other collections from Finca Pirineos;
[also collected below San Martin Chile Verde).

A slender, sparsely branched, erect shrub 1.5-2 meters high, glabrous through-
I out, the upper internodes rather short; petioles rather stout, mostly 6-7 mm.
[' long or sometimes slightly longer; leaf blades rather thin but firm, drying dark
green or grayish green, opaque or usually so, lance-oblong or narrowly ovate-
oblong, mostly 11-15 cm. long and 4-6 cm. wide, narrowly very long-acuminate,
I rounded at the base or sometimes subcordate, paler beneath, subequal and sym-
metric, palmately 5-nerved, the nerves very slender but prominent beneath, the
I lateral ones usually irregular and more slender than the others, the principal
| veins divergent at a right angle from the costa and straight or nearly so or the
upper ones arcuate-ascending, laxly reticulate; peduncles opposite the leaves,
very slender, almost filiform, mostly 1.5-2.5 cm. long, glabrous; spikes very slender,

316 FIELDIANA: BOTANY, VOLUME 24

pendulous, about 10 cm. long and little more than 2 mm. thick, white, the rachu
glabrous or minutely puberulent; bracts glabrous, ciliolate.

Frutex gracilis fere omnino glaber; folia breviter petiolata, petiolo basi tantum
vaginante, in sicco non fuscescentia, opaca, lanceolato-oblonga vel anguste ovato-
oblonga modica, anguste longissime acuminata, basi subaequali late rotundata
vel interdum subcordata, palmatim 5-nervia; pedunculi oppositifolii fere filiformes,
vulgo 1.5-2.5 cm. longi; spicae graciles vix ultra 2 mm. crassae ca. 10 cm. longae;
bracteae glabrae ciliolatae.

A relative of P. Amalago L., but apparently fully distinct from
that species.

Piper phaeophyllum Trelease & Standley, sp. nov.

Moist or wet forest or thickets, 1,200 meters or lower; endemic;
Pete"n (Cerro Ceibal); Verapaz (type collected above Tamahu,
Standley 70911; in Herbarium of Chicago Natural History Museum).

A shrub 1.5-3 meters high, the branches short-hispidulous, pale, rather slender,
with short or elongate internodes; petioles stout, 4-6 mm. long, exalate, dilated '
at the base, densely puberulent; leaf blades thick and stiff, small, fuscescent when
dry, minutely pellucid-punctate or more often punctate, lanceolate or lance- i
oblong, mostly 7-10 cm. long and 2.5-4 cm. wide, narrowly long-acuminate,
unequal and slightly oblique at the base, rounded or obtuse on each side, minutely
scabrous above, very rough to the touch, not rugose, somewhat paler beneath,
densely strigose on the nerves, scaberulous between them and rough to the touch,
penninerved, the nerves usually 3 on each side, arcuate-ascending at an angle of
about 45 degrees, the veins mostly very prominent, laxly reticulate; peduncles
opposite the leaves, stout, about 6 mm. long, scabrous-puberulent; spikes erect-
spreading, slender, about 8.5 cm. long, slightly more than 2 mm. thick, obtuse;
bracts puberulent; anthers short-exserted; fruit puberulent about the apex.

Kami breviter hispiduli, graciles, internodiis brevibus vel elongatis; petioli
crassi 4-6 mm. longi exalati basi dilatati, dense puberuli; lamina parva in sicco
fuscescens minute pellucido-punctata, lanceolata vel lanceolato-oblonga, anguste
longiacuminata, basi subaequali subobliqua utroque latere rotundata vel obtusa,
supra minute scabra tactu aspera, non bullata, subtus paullo pallidior ad nervos
dense strigosa inter nervos scaberula, penninervia, nervis utroque latere vulgo 3
arcuato-adscendentibus, venis subtus prominentibus laxe reticulatis; pedunculi
oppositifolii 6 mm. longi, spicis gracilibus ca. 8.5 cm. longis 2 mm. vel paullo
ultra crassis.

Piper philodendroides Standl. & Steyerm., sp. nov.

Known only from the type, Izabal, Cerro San Gil, wet forested
slope, 300-900 meters, Steyermark 41928 (in Herbarium of Chicago
Natural History Museum).

A stout shrub of 3-4.5 meters, glabrous throughout, the young branches stout
and thick, brown, with short internodes; petioles stout, 7-12 cm. long, scarcely
vaginate; leaf blades triangular-ovate or broadly oblong-ovate, about 28 cm. long
and 13-16 cm. wide, acute or short-acuminate, almost equal at the base but one

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 317

of the sides slightly larger than the other, very shallowly and broadly cordate,
somewhat fuscescent when dry, dull, densely and minutely pellucid-punctate,
penninerved, the nerves 3-4 on each side, arcuate-ascending at an angle of 45
I degrees or more, prominent beneath, the veins prominent, very laxly and openly
reticulate; spikes unknown.

Frutex erectus glaber, ramis crassis, internodiis brevibus; petioli crassi 7-12
cm. longi fere omnino nudi ; lamina crassiuscula triangulari-ovata vel late oblongo-
ovata magna acuta vel breviter acuminata, basi lata subaequali breviter lateque
cordata, lobis basalibus paullo inaequalibus, dense minute pellucido-punctata,
penninervia, nervis utroque latere 3-4 arcuato-adscendentibus subtus prominenti-
bus, venis subtus prominentibus laxe reticulatis; spicae ignotae.

Apparently a well-marked species, the large long-petiolate leaves
somewhat suggestive of those of certain species of Philodendron.

Piper pinetorum Standl. & Steyerm., sp. nov.

Known only from the type, Huehuetenango, in forest of Liquid-
amber and Pinus Ayacahuile, Cerro Victoria, near Barillas, Sierra
de los Cuchumatanes, 1,800-2,000 meters, Steyermark 49712 (in
Herbarium of Chicago Natural History Museum).

A shrub 3 meters high, apparently densely branched, glabrous throughout or
nearly so, the youngest parts sometimes with a few scattered inconspicuous hairs,
the internodes mostly short; petioles stout, about 2 cm. long, winged nearly or
quite their full length; leaf blades thick, more or less bullate, slightly blackening
or brownish when dry, opaque, oblong or elliptic-oblong, about 11 cm. long and
4 cm. wide, acute or abruptly short-acuminate with an obtuse tip, very unequal
at the base, one side 6-7 mm. longer than the other, cordate or subcordate at the
base, at least on one side, the other side rounded, dull when dried, the nerves and
veins somewhat impressed on the upper surface, the blades penninerved, the
nerves usually 3 on each side, ascending at a narrow angle, slightly arcuate,
prominent beneath, the veins scarcely elevated, laxly reticulate; peduncles stout,
about 1 cm. long; spikes pendent, with alternating rings of white and pale green,
very slender, mostly about 16 cm. long and 2 mm. thick; scales very small, ciliate:
ovary glabrous.

Frutex trimetralis fere omnino glaber; petioli crassi fere tota longitudine
vaginantes; lamina crassa epunctata in sicco fuscescens vel subtus brunnescens
plus minusve bullata oblonga vel elliptico-oblonga modica, acuta vel breviter
obtuso-acuminata, basi valde inaequalis, saltern uno latere cordata vel subcordata,
altero rotundata, pinnatinervia, nervis utroque latere vulgo 3; spicae ca. 1 cm.
longe pedunculatae graciles pendulae ca. 16 cm. longae et 2 mm. crassae.

Piper planadosense Trelease & Standley, sp. nov.

Known only from the type, Zacapa, Sierra de las Minas, cloud
forest on summit, vicinity of Finca Planados, 2,500 meters, Steyer-
mark 29994 (in Herbarium of Chicago Natural History Museum).

An erect shrub 4.5 meters high, the young branches rather stout, densely
puberulent or hispidulous with spreading hairs, the internodes short; petioles

318 FIELDIANA: BOTANY, VOLUME 24

stout, 1 cm. long or shorter, densely hispidulous, vaginate only at the base; leaf
blades somewhat blackish when dried, dull, thin but firm, densely pellucid-punctate,
elliptic or ovate-elliptic, mostly 13-15 cm. long and 6-7 cm. wide, rather abruptly
acuminate, conspicuously unequal and oblique at the base, shallowly cordate or
subcordate on one side, rounded or obtuse on the other, glabrous above, often
brownish beneath, densely strigillose or puberulent on the nerves and veins,
between them glabrous or nearly so and finely granular, penninerved, the nerves
3-4 on each side, very slender, prominent, ascending at an angle of 45 degrees
or less, arcuate, the veins prominent, laxly reticulate; peduncles opposite the
leaves, stout, about 1 cm. long, puberulent or glabrate; spikes in fruit about 7 cm.
long and 5 mm. thick; fruits glabrous or nearly so, almost 2 mm. long, subglobose.

Frutex, ramis crassiusculis dense puberulis vel hispidulis; petioli crassi ca.
1 cm. longi vel breviores hispiduli basi tantum vaginantes; lamina in sicco plus
minusve fuscescens dense pellucido-punctata elliptica vel ovato-elliptica modica,
subabrupte acuminata, basi manifeste inaequalis et obliqua, uno latere cordata
vel subcordata, altero rotundata vel obtusa, supra glabra opaca, subtus ad nervos
venasque dense strigillosa vel puberula, penninervia, nervis utroque latere 3-4
angulo angusto arcuato-adscendentibus; pedunculi oppositifolii ca. 1 cm. longi,
spicis fructiferis ca. 7 cm. longis atque 5 mm. crassis.

Piper pogonioneuron Trelease & Standley, sp. nov. Cordoncillo.

Moist or wet thickets or dense mixed forest, chiefly in the moun-
tains, sometimes on white-sand slopes, 2,250 meters or lower;
endemic; Alta Verapaz; Izabal; Sacatepe'quez; Chimaltenango; Que-
zaltenango (type from western slopes of Volcan de Zunil, opposite
Santa Maria de Jesus, Steyermark 35174; in Herbarium of Chicago
Natural History Museum); San Marcos.

An erect shrub 1.5-3 meters high, the branches rather slender, densely pilose
or villous-pilose with rather long, spreading, mostly fulvescent, multicellular
hairs, the upper internodes short; petioles mostly 1-2 cm. long, slender or stout,
not winged, dilated at the base, villous-pilose; leaf blades rather thin, dull, scarcely
fuscescent when dried, generally opaque but with translucent veins, elliptic to
ovate-elliptic or oblong-elliptic, mostly 16-22 cm. long and 6.5-10 cm. wide,
rather abruptly and narrowly long-acuminate, somewhat unequal and conspic-
uously oblique at the base, shallowly cordate or rounded on one side, usually
obtuse on the other, densely scabrous above and very rough, villous-hirsute along
the costa, not or very obscurely rugose, villous-pilose or hirsute beneath on the
nerves and veins, scaberulous between them, penninerved, the nerves 3-5 on
each side, ascending at an angle of 45 degrees or more, arcuate, the veins often
prominent, laxly reticulate; peduncles opposite the leaves, usually about 1.5 cm.
long, scabrous and scabrous-hirtellous; spikes erect or spreading, mostly 7-10 cm.
long, 3 mm. thick, obtuse or subacute; bracts rounded-peltate at the apex, pale,
densely pubescent; anthers subexserted; fruits about 1 mm. long, somewhat
depressed at the apex, obtusely angulate.

Frutex, ramis dense pilis longiusculis patentibus fulvescentibus multilocu-
laribus pilosis; petioli 1-2 cm. longi exalati basi dilatati villoso-pilosi; lamina
majuscula non lucida in sicco vix fuscescens opaca, venis pellucidis, elliptica usque

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 319

ovato-elliptica vel oblongo-elliptica, subabrupte angusteque longiacuminata, basi
inaequalis obliqua uno latere cordata vel rotundata, altero obtusa, supra dense
scabra et asperrima, non rugosa, ad costam villoso-hirsuta, subtus ad nervos
venasque villoso-pilosa vel hirsuta, inter venas scaberula, penninervia, nervis
utroque latere 3-5, angulo semirecto vel latiore arcuato-adscendentibus; pedunculi
oppositifolii ca. 1.5 cm. longi, spicis plerumque 7-10 cm. longis 3 mm. crassis obtusis
vel acutiusculis; bracteae apice rotundo-peltatae dense pubescentes; fructus apice
subdepressus obtuse angulatus.

Among the material referred here, some of which was not seen
by Trelease, there are twelve sheets that he has indicated as types
of new species. One or two of these may have some claims to specific
rank but we are inclined to believe that all are conspecific.

Piper pseudoasperifolium C. DC. in DC. Prodr. 16, pt. 1:
318. 1869. P. vestitifolium C. DC. Bot. Gaz. 70: 183. 1920 (type
collected near Cajabon, Alta Verapaz, 0. F. Cook & R. F. Griggs 651).
P. sibunense Trelease in Standl. Field Mus. Bot. 12: 408. 1936 (type
from Gracie Rock, Sibun River, British Honduras, P. H. Gentle 1562).
Cordoncillo; Biritac (Coban, Quecchi) ; Cuturro (Izabal).

Moist or wet or sometimes rather dry, mixed, mountain or low-
land forest, or in thickets, frequently in pine-oak forest, 2,300 meters
or lower, most common at middle elevations; Pete"n; Alta Verapaz;
Baja Verapaz; Izabal; Chiquimula; Suchitepequez; Sacatepe"quez;
Chimaltenango; Quiche"; Huehuetenango; Quezaltenango; San
Marcos. Southern Mexico, and probably farther southward in
Central America.

An erect, rather stout, often much branched shrub, usually 1.5-5 meters
high, the branches very densely hispidulous with whitish or yellowish, spreading
or often reflexed, harsh hairs, the internodes mostly short; leaves usually grayish
when dried, pellucid-punctate or almost opaque, thick, the stout petioles 1.5 cm.
long or shorter, vaginate only at the base, densely hispidulous; leaf blades lance-
elliptic or ovate-elliptic, mostly 11-22 cm. long and 4-8 cm. wide, rather abruptly
acuminate or long-acuminate, conspicuously oblique at the base, rounded or obtuse
on each side or sometimes subcordate on one side, penninerved, the nerves usually
3-4 on either side, sometimes as many as 5-6, arising at or below the middle of
the blades, very prominent beneath, the blades densely scabrous on the upper
surface and very rough to the touch, usually very conspicuously bullate, the veins
strongly impressed, at least on the older leaves, densely hispidulous beneath, at
least on the nerves and veins, the veins very prominent and closely reticulate;
peduncles stout, equaling or shorter than the petioles, densely hispidulous; spikes
erect, rather stout, whitish, cream-colored, or pale green, at maturity about 9 cm.
long and 4 mm. thick, obtuse; bracts peltate and truncate, densely hispidulous;
stamens 4; fruits glabrous or nearly so.

This is a very common shrub in some parts of Guatemala and
one of the commonest of all the local species. Trelease designated

320 FIELDIANA: BOTANY, VOLUME 24

as new species twenty or more plants that we have placed here.
Called "Spanish elder" in British Honduras. The plant has been
reported from Guatemala as P. hirsutum Swartz and P. mexicanum
C. DC.

Piper psilorhachis C. DC. Ann. Cons. Jard. Bot. Geneve 2:
253. 1898 (type supposed to have been collected in Guatemala,
Friedrichsthal 1538, the locality unknown). P. colaphitolerans Tre-
lease in Standl. Field Mus. Bot. 17: 231. 1937 (type from Ixlu,
Pete"n, C. L. Lundell 3827). Cucsub (Pete*n, Maya).

Moist or wet, mixed forest, 1,800 meters or lower, often descend-
ing nearly or quite to sea level; Pete"n; Alta Verapaz; Escuintla;
Guatemala; San Marcos. Tabasco; British Honduras.

A slender shrub or small tree, rarely 7.5 meters high, usually much lower,
glabrous throughout; leaves on slender petioles 1 cm. long or usually shorter,
often blackening when dried, pellucid-punctate, lance-ovate or lance-elliptic,
mostly 7-10.5 cm. long and 3-4.5 cm. wide, rather abruptly and narrowly long-
acuminate, obtuse and only slightly unequal at the base, palmately 5-nerved,
slightly paler beneath, the veins prominulous, very laxly reticulate; peduncles
about equaling the petioles or somewhat longer; spikes slender, 5-10 cm. long,
scarcely 3 mm. thick, the rachis glabrous, the bracts minute, ovate, glabrous;
stamens 5-6; fruits oblong, 5-6 mm. long or less at maturity, very minutely
puberulent; stigmas 3, sessile, minute.

The names used in British Honduras are "Spanish elder" and
"chucsuc" (Maya).

Piper punctulatum Standl. & Steyerm., sp. nov.

Dense, moist or wet, mixed, mountain forest, 1,500-2,600 meters;
endemic; Alta Verapaz; El Progreso (type from Sierra de las Minas,
between Finca Piamonte and top of Montana Piamonte, Steyermark
43622; in Herbarium of Chicago Natural History Museum); Zacapa
(Sierra de las Minas).

A shrub 1-3 meters high, sometimes herbaceous almost throughout, sparsely
branched, erect, the branches with elongate internodes, glabrous or nearly so;
petioles rather slender, mostly 3-5 cm. long, winged for almost their whole length,
dilated and clasping at the base, glabrous or puberulent, especially above; leaf
blades large, thin, usually fuscescent when dry, densely and rather coarsely
pellucid-punctate, broadly ovate or rounded-ovate, sometimes rhombic-ovate,
mostly 12-23 cm. long and 9.5-16 cm. wide, abruptly short-acute, sometimes
rounded and short-cuspidate, shallowly and broadly cordate at the base, the two
sides somewhat unequal, deep green and glabrous above, dull when dried, slightly
paler beneath, densely blackish-puncticulate, densely puberulent on the nerves
and veins and sometimes between them, penninerved, the nerves 3-4 on each
side, very slender, prominulous, ascending at an angle of 45 degrees or less, slightly

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 321

arcuate, the veins very slender and openly reticulate; peduncles opposite the leaves,
very slender, about 3.5 cm. long, glabrous, or puberulent above; spikes short and
stout (only one well-developed one seen), about 7 cm. long and 5 mm. thick,
attenuate upward, the bracts pale, densely pubescent; ovaries glabrous.

Frutex erectus sparse ramosus, ramis glabris vel glabratis; petioli 3-5 cm.
longi in fere tota longitudine vaginati glabri vel superne puberuli; lamina magna
tenuis in sicco fuscescens dense et subgrosse pellucido-punctata, late ovata vel
rotundo-ovata interdum rhombeo-ovata, abrupte breviter acuta vel rotundata
et breviter cuspidata, basi breviter lateque cordata, lobis basalibus paullo inae-
qualibus, supra viridis, glabra, subtus paullo pallidior ubique dense fusco-puncticu-
lata, ad nervos venasque et interdum inter venas sat dense puberula, penninervia,
nervis utroque latere 3-4 angulo angusto adscendentibus, venis prominulis laxis-
sime reticulatis; pedunculi oppositifolii ca. 3.5 cm. longi glabri vel superne puberuli,
spicis crassis ca. 6 cm. longis et 5 mm. basi crassis sursum sensim attenuatis.

We have made about eight collections of this species, most of
them in various regions of Alta Verapaz.

Piper ret a 1 h uleuense Trelease & Standley, sp. nov. Corri-
miento.

Moist or rather dry thickets or lowland forest, 325 meters or
lower; endemic; Santa Rosa; Retalhuleu (type from the region of
Las Delicias, south of Retalhuleu, Standley 88047; in Herbarium of
Chicago Natural History Museum).

Plants chiefly herbaceous but often more or less woody about the base, some-
times suberect and 1.5 meters high or less, occasionally scandent, commonly
procumbent and rooting at the lower nodes, the branches pale green, rather stout,
striate, glabrous or very obscurely puberulent, strongly nodose; petioles 1.5-2.5 cm.
long, short-hirsute, vaginate only at the base; leaf blades thin and rather flaccid,
green when dried, cordate-orbicular or ovate-orbicular, 4-8 cm. long and 4-8.5
cm. wide, rather abruptly acute or short-acuminate with an obtuse tip, deeply
and narrowly cordate at the base, hirtellous above along the nerves, paler beneath,
hirtellous on the nerves and veins, palmately 7-nerved, minutely pellucid-punctate,
minutely blackish-puncticulate beneath; peduncles opposite the leaves, 7-10 mm.
long, short-hispidulous or glabrate; spikes erect, 1.5-3.5 cm. long, 3 mm. thick
in fruit; fruits oval-globose, 1.5 mm. long, obscurely puberulent or granular.

Plantae suffruticosae et erectae vel scandentes, saepius fere omnino herbaceae
et procumbentes, internodiis saepe valde elongatis, ramis striatis glabris vel obscure
puberulis; folia longipetiolata, petiolo breviter hirsute basi tantum vaginante;
lamina cordato-orbicularis vel ovato-orbicularis in sicco viridis subabrupte acuta
vel acuminata, acumine obtuso, profunde angusteque basi cordata, supra secus
nervos hirtella, subtus ad nervos venasque hirtella, palmatim 7-nervia, pellucido-
punctata; pedunculi oppositifolii breviter hispiduli vel glabrati, spicis erectis
breves; fructus ovali-globosus obscure puberulus vel granulosus.

Piper santae-clarae Standl. & Steyerm., sp. nov.
Known only from the type, Suchitepequez, Volcan de Santa
Clara, between Finca El Naranjo and the upper slopes, 1,250-2,600

322 FIELDIANA: BOTANY, VOLUME 24

meters, Steyermark 46624 (in Herbarium of Chicago Natural History
Museum).

A slender branched shrub about a meter high, the branches rather sparsely
puberulent or short-pilosulous, the internodes short; petioles short, the longest
5 mm. long, rather stout, puberulent; leaf blades thin, somewhat blackish when
dried, minutely pellucid-punctate, lanceolate or oblong-lanceolate, mostly 8-10.5
cm. long and 2-3 cm. wide, very narrowly attenuate-acuminate, strongly unequal
at the base, rounded or subcordate on each side, one side much longer decurrent
than the other, glabrous above, somewhat paler beneath, minutely hirtellous on
the nerves with pale spreading hairs, palmately 3-nerved or obscurely 5-nerved,
the veins divergent from the costa at a wide angle, laxly reticulate; peduncles
very slender, divaricate and perhaps pendulous, mostly 2.5-3 cm. long, thinly
pilosulous or glabrate; spikes small and slender, white, at anthesis 3-3.5 cm. long,
2 mm. thick, obtuse; anthers subexserted; bracts ciliate; ovaries glabrous.

Frutex metralis, ramis gracillimis sparse puberulis vel breviter pilosulis; folia
tenuiter membranacea breviter petiolata in sicco fuscescentia minute pellucido-
punctata, lanceolata vel oblongo-lanceolata, parva, anguste longeque attenuato-
acuminata, basi valde inaequalia, utroque latere rotundata vel subcordata, palma-
tim 3-nervia vel obscure 5-nervia, supra glabra, subtus at nervos minute hirtella;
spicae oppositifolii patentes vel pendulae 3-3.5 cm. longae, pedunculis gracillimis
spicis fere aequilongis.

A relative of P. Standleyi Trelease, which has normally very much
shorter peduncles.

Piper scabrum Swartz, Fl. Ind. Occ. 59. 1797. P. Chanekii
Trelease in Standl. Field Mus. Bot. 12: 407. 1936 (type from El Cayo,
British Honduras, Mercedes Chanek 10). Cordoncillo.

Moist or wet or sometimes rather dry thickets or forest, often
in open forest, sometimes in second growth, 1,900 meters or lower,
most common below 1,000 meters; Pete"n; Alta Verapaz; Izabal;
Santa Rosa; Escuintla; Sacatepe"quez ; Chimaltenango; Suchite-
pequez; Retalhuleu; Quezaltenango; San Marcos; Huehuetenango;
Quiche". Southern Mexico; British Honduras; probably throughout
Central America; West Indies; probably also in South America.

An erect, branched, rather stout shrub, commonly 1.5-3 meters high, the
young branches densely hispidulous with short, whitish, spreading or somewhat
reflexed, harsh hairs, the internodes short or elongate; petioles stout, 1 cm. long
or usually shorter, vaginate near the base, hispidulous; leaf blades rather thick
and firm, usually drying green or grayish green, rarely blackish, little if at all
lustrous, pellucid-punctate, sometimes very obscurely so, green on the upper
surface, scabrous and very rough to the touch, not bullate, somewhat paler beneath,
inconspicuously scabrous over the whole surface, strigillose or appressed-hispidulous
on the nerves and veins, lance-elliptic to broadly ovate-elliptic, mostly 10-20 cm.
long and 5-8 cm. wide, abruptly acuminate or cuspidate-acuminate, very unequal
at the base, rounded or obtuse on each side or the shorter side sometimes acute;

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 323

peduncles stout, about equaling the petioles, hispidulous; spikes erect, slender,
whitish, cream-colored, or grayish green, obtuse, mostly 8-10 cm. long and about
3.5 mm. thick; bracts densely pubescent; anthers exserted.

This is easily the most common species of Piper found in Guate-
mala, although perhaps not the widest in range. It is an abundant,
often somewhat weedy shrub in many parts of the country, especially
on the Pacific slope. We have collected numerous specimens, and
they are almost uniform in their characters. Despite this, Trelease
has designated among our collections at least seventy new but
fortunately unpublished species. It is in this group of the genus
that he has in the past indicated and often published the greatest
number of species, in Panama, Costa Rica, Honduras, and else-
where, and the number of such published names that will have to
be relegated to synonymy when the genus is carefully monographed
probably will reach more than a hundred, when the South American
species are considered. In treating the Guatemalan Piper species
we have divided this group perhaps too finely, although we have
recognized a number of related species that seem to have some
claim to specific rank. West Indian material of P. scabrum usually
has slightly larger leaves than mainland specimens. This species
has been reported from Guatemala as P. hispidum Swartz and P.
hirsutum Swartz.

Piper Schippianum Trelease & Standley, sp. nov. (P. Schip-
pianum Trelease ex Standl. Field Mus. Bot. 12: 104. 1936, without
Latin description).

Known only from the type, British Honduras, wet forest, 19-mile,
Stann Creek Railway, 90 meters, W. A. Schipp 316; collected also
at Honey Camp; type in Herbarium of Chicago Natural History
Museum.

A tree 9 meters high with a trunk 12 cm. in diameter, glabrous throughout,
the branches strongly nodose; leaves rather small, thin but firm, epunctate, on
slender petioles 5-7 mm. long, narrowly oblong or lance-oblong, mostly 9.5-11.5
cm. long and 3-3.5 cm. wide, abruptly and narrowly long-acuminate, at the base
very acute on one side, obtuse on the other, the obtuse side decurrent lower on
the petiole than the other side, palmately 3-nerved or obscurely 5-nerved, the 3
principal nerves extending to the apex of the blade, the veins transverse, very
slender, closely prominulous-reticulate; spikes very slender, opposite the leaves,
on slender glabrous peduncles about 13 mm. long, straight or somewhat flexuous,
7-14 cm. long, 2-3 mm. thick, very densely flowered; immature fruits globose-
ovoid, 1.5 mm. long, glabrous.

Arbor glabra; folia inter minora firme membranacea epunctata anguste oblonga
vel lanceolato-oblonga longe anguste caudato-acuminata, basi inaequali uno latere

324 FIELDIANA: BOTANY, VOLUME 24

acutissima, altero obtusa, latere obtuso longius decurrente, palmatim 3-nervia
vel obscure 5-nervia, venis utrinque prominulis arete reticulatis; spicae breviter
pedunculatae gracillimae foliis longiores; fructus glaber.

Piper sempervirens (Trelease) Lundell, Carnegie Inst. Wash.
Publ. 478: 208. 1937. Arctottonia sempervirens Trelease in Standl.
Field Mus. Bot. 12: 405. 1936.

Moist or wet, mixed, lowland forest, 300 meters or lower; Pete"n
(type from Uaxactun, H. H. Bartlett 12563); Alta Verapaz (Cerro
Chinaja). British Honduras (Chalilla Crossing, El Cayo District);
Yucatan Peninsula of Mexico.

A slender shrub or small tree, commonly 1.5-5 meters high, the trunk some-
times 15 cm. in diameter, glabrous throughout; leaves on slender petioles 8 mm.
long or short, chartaceous and stiff, usually very lustrous, grayish green when
dried, elliptic or oblong-elliptic, mostly 6-9 cm. long and 3-4.5 cm. wide, abruptly
long-acuminate, acute and only slightly unequal at the base, epunctate, palmately
3-5-nerved, the inner nerves ending a short distance below the apex, concolorous,
the veins prominulous and rather closely reticulate on both surfaces; inflorescences
terminal, racemose, 1.5-4 cm. long, on peduncles 14 mm. long or shorter, lax, the
pedicels divaricate, about equaling the flowers; fruits ovoid, slightly more than
2 mm. long.

Piper Standleyi Trelease in Standl. Journ. Wash. Acad. Sci.
13: 366. 1923. Cordoncillo.

Wet to rather dry, mixed, mountain forest or thickets, sometimes
in rocky places, 1,200-2,400 meters; Guatemala; Chimaltenango;
Suchitepequez ; Quezaltenango. El Salvador, the type from Volcan
de San Salvador.

A very slender, erect, branched shrub 1-2.5 meters high, the branches densely
puberulent or hirtellous; petioles very short, 4 mm. long or less, densely hirtellous,
vaginate only at the base; leaf blades drying grayish green, very minutely pellucid-
punctate or almost wholly opaque, narrowly lanceolate to lance-oblong, mostly
7-10 cm. long and 2-3 cm. wide, occasionally somewhat larger, very narrowly
long-attenuate, somewhat oblique and unequal at the base, rounded or very obtuse,
usually shallowly and narrowly cordate, the auricles unequal in size, glabrous
above or practically so, dull, paler beneath, minutely hirtellous on the nerves,
palmately 3-nerved or obscurely 5-nerved, the veins conspicuous, pale, very openly
reticulate; peduncles opposite the leaves, 7-15 mm. long, puberulent; spikes
spreading or pendulous, small, mostly 2-3 cm. long, or sometimes 5-7 cm. long,
in fruit about 4 mm. thick; bracts very small, densely pubescent on the margins;
stigmas 3, short and thick, sessile; fruits globose, 1.5 mm. long, puberulent above
or granular.

The material referred here is somewhat variable and may be
separable into two species, except that some specimens are inter-
mediate between the extreme forms that might be considered distinct
species.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 325

Piper stillans Trelease & Standley, sp. nov.

Known only from the type, Quiche", Cerro Putul, Zona Reina,
1,200 meters, A. F. Skutch 1822 (in Herbarium of Chicago Natural
History Museum).

A slender shrub about a meter high, glabrous throughout; petioles stout,
about 3 mm. long, vaginate only at the very base; leaf blades drying green, densely
and minutely pellucid-punctate, thin, oblong-elliptic or oblong-ovate, 7.5-10 cm.
long, 2.5-4 cm. wide, caudate-acuminate, with a very long and narrow, attenuate,
obtuse acumen, rounded to subacute at the base, dull when dried, palmately 3-
nerved, or 5-nerved but the outer nerves very obscure and irregular, the veins
divergent from the costa at almost a right angle, straight or somewhat curved;
peduncles opposite the leaves, very slender and almost filiform, 2-2.5 cm. long;
spikes small, very slender, probably pendulous, 3-4.5 cm. long, scarcely 2 mm.
thick, obtuse; bracts glabrous, ciliate.

Frutex gracilis humilis omnino glaber; folia inter minora breviter petiolata,
petiolo 3 mm. tantum longo basi breviter vaginante, in sicco viridia dense minute
pellucido-punctata, oblongo-elliptica vel oblongo-ovata, anguste longeque caudato-
acuminata, acumine obtuso, basi rotundata usque subacuta, palmatim 3-nervia;
pedunculi oppositifolii fere filiformes 2-2.5 cm. longi; spicae gracillimae breves,
3-4.5 cm. tantum longae vix 2 mm. diam. ut videtur pendulae obtusae; bracteae
glabrae ciliolatae.

Piper subeburneum Trelease & Standley, sp. nov.

Moist or wet, mixed, mountain forest, 900-2,400 meters; endemic;
Quezaltenango; San Marcos (type from northwestern slopes of
Volcan de Tajumulco, barrancos six miles southwest of Tajumulco,
Steyermark 36702; in Herbarium of Chicago Natural History Mu-
seum).

A glabrous shrub 1.5-2.5 meters high, the branches with short internodes;
petioles rather stout, 1-2 cm. long, vaginate only near the base; leaf blades deep
green or blackish green when dried, rather thick and firm, dull, minutely pellucid-
punctate, lance-oblong or often narrowly so, mostly 13-19 cm. long and 4.5-5.5
or rarely 7 cm. wide, rather abruptly short-acuminate with a subacute tip, sub-
equal but often slightly oblique at the base and rounded or very obtuse, deep
green above, the nerves and veins usually prominulous or sometimes not at all
elevated, much paler beneath and glaucous or glaucescent, penninerved, the
nerves 6-9 on each side, divergent at an angle of 45 degrees or often much broader,
arcuate, ascending, the veins prominulous or obscure, very laxly reticulate;
spikes unknown.

Frutex glaber; folia modica 1-2 cm. longe petiolata, petiolo basi tantum
vaginante, lanceolato-oblonga vel anguste lanceolato-oblonga in sicco vix fusces-
centia minute pellucido-punctata, subabrupte breviterque acuminata, basi subae-
quali interdum aliquanto obliqua rotundata vel obtusissima, supra intense viridia,
subtus multo pallidiora glauca vel glaucescentia, penninervia, nervis utroque
latere 6-9 arcuato-adscendentibus, venis prominulis vel obscuris laxissime reticu-
latis; spicae ignotae.

326 FIELDIANA: BOTANY, VOLUME 24

A rather common shrub of the mountains of the Occidente, repre-
sented by nine collections. The species is noteworthy, for the lower
surface of the leaves has a very pale, glaucous or glaucescent color,
which often has a slightly yellowish tinge.

Piper subcitrifolium C. DC. Bot. Gaz. 70: 186. 1920. Cordon-
cillo.

Wet to rather dry forest and thickets, 1,200 meters or lower;
Santa Rosa (type from Cerro Redondo, J. D. Smith 3827); Escuintla;
Guatemala; Sacatepe"quez; Suchitepequez; Retalhuleu; Quezalte-
nango; San Marcos. Salvador.

An erect, sparsely branched shrub about 2 meters high, the branches glabrous
or merely granular, the internodes often short; petioles short, rather stout, generally
less than 1 cm. long, minutely puberulent, vaginate only at the base; leaf blades
elliptic or oblong-elliptic, epunctate or inconspicuously pellucid-punctate, with
translucent veins, green or somewhat blackish when dried, mostly 11-16 cm. long
and 5.5-7.5 cm. wide, abruptly acuminate or long-acuminate, acute or obtuse
at the base, glabrous above, usually somewhat lustrous, slightly paler beneath,
minutely puberulent or strigillose along the nerves, elsewhere glabrous or nearly
so, penninerved, the nerves usually 3 on each side, sometimes as many as 5,
arcuate-ascending, the upper ones arising above the middle of the blade, the
veins scarcely prominulous, inconspicuous, laxly reticulate; peduncles stout,
usually shorter than the petioles, glabrate; spikes pale green or whitish, mostly
8-10 cm. long and 3 mm. thick, obtuse; bracts truncate-triangular at the apex,
hispidulous on the margins; stamens 4; stigmas 3, sessile.

A common plant of the Pacific foothills and plains.

Piper tacananum Trelease & Standley, sp. nov.

Moist or wet thickets or dense mixed mountain forest, 1,800-
2,500 meters; endemic; Solola (type from Volcan de San Pedro,
northern slopes above village of San Pedro, Steyermark 47250; in
Herbarium of Chicago Natural History Museum) ; San Marcos (Rio
Tacand above San Antonio).

A slender shrub 1.5-3 meters high, the branches sparsely or densely pilose or
villous-pilose with weak spreading hairs; petioles rather stout, 1 cm. long or shorter,
vaginate only near the base, sparsely or densely pilose; leaf blades drying dark
green or fuscescent, dull or very slightly lustrous, opaque or obscurely pellucid-
punctate, oblong-ovate to oblong-lanceolate, mostly 12-16 cm. long and 5-7.5
cm. wide, narrowly very long-acuminate, at the base cordate or shallowly cordate,
at least in the lower leaves, or merely subcordate in the upper leaves, sometimes
villosulous above at the very base but glabrous elsewhere, somewhat paler beneath
or brownish, densely villous-pilosulous beneath over the whole surface or at least
on the nerves, palmately 5-nerved, the lateral nerves very irregular and usually
evanescent above, the principal veins divaricate from the costa at almost a right
angle, straight or slightly arcuate, laxly reticulate; peduncles slender, opposite

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 327

he leaves, densely villous-pilose, 2 cm. long or somewhat longer; spikes white,
nutant, very slender, about 9 cm. long and 2 mm. thick; bracts glabrous, ciliate;
anthers exserted; fruit glabrous or nearly so.

Frutex gracilis erectus, ramulis sparse denseve pilosis vel villoso-pilosis; folia
breviter petiolata, petiolo basi tantum vaginante sparse denseve piloso, in sicco
fuscescentia, opaca vel obscure punctata, oblongo-ovata vel oblongo-lanceolata,
anguste longissime acuminata, basi subaequali vel inaequali cordata vel breviter
cordata, supra fere omnino glabra, subtus saltern ad nervos dense villoso-pilosula,
palmatim 5-nervia, venis laxe reticulatis; pedunculi dense villoso-pilosi, spicis
nutantibus gracilibus elongatis; bracteae glabrae ciliolatae.

Piper tacticanum Trelease & Standley, sp. nov.

Dense, moist or wet, mixed, mountain forest, 1,350-1,650 meters;
endemic; Alta Verapaz (type from mountains east of Tactic, on the
road to Tamahu, Standley 61197; in Herbarium of Chicago Natural
History Museum) ; Quezaltenango (Finca Pirineos below Santa Maria
de Jesus).

A shrub 1-1.5 meters high, the branches slender, glabrous or sparsely and
minutely puberulent, strongly nodose, with short internodes; petioles rather
slender, 6-8 mm. long or sometimes as much as 15 mm., minutely and sparsely
strigillose or glabrate, not winged; leaf blades small, thin, blackish when dried,
minutely pellucid-punctate, lance-ovate, mostly 11-13 cm. long and 4.5-5 cm.
wide, rather abruptly and narrowly very long-acuminate, somewhat unequal and
oblique at the base, obtuse or sometimes subacute on each side, glabrous above,
dull, brownish beneath when dried, minutely strigillose on the nerves and veins
and sometimes between them but to the naked eye appearing glabrous, penni-
nerved, the nerves usually 3 on each side, very slender and scarcely elevated, arcu-
ate-ascending, the veins obscure, very laxly reticulate; peduncles opposite the
leaves, slender, about 1 cm. long, glabrous; spikes slender, erect-spreading, about
6.5 cm. long and 3 mm. thick, short-mucronate at the apex; bracts densely pale-
pubescent on the margins.

Frutex, ramis gracilibus glabris vel glabratis; folia breviter petiolata, petiolo
non alato vulgo 6-8 mm. longo minute sparseque strigilloso vel fere glabro, tenuia
in sicco fuscescentia minute pellucido-punctata, lanceolato-ovata, subabrupte
anguste longissime acuminata, basi inaequali atque obliqua utroque latere obtusa
vel subacuta, supra glabra, subtus in sicco brunnescentia ad nervos venasque et
interdum inter venas minute strigillosa, penninervia, nervis utroque latere vulgo
3 arcuato-adscendentibus; pedunculi oppositifolii ca. 1 cm. longi, spicis erecto-
patentibus gracilibus ca. 6.5 cm. longis atque 3 mm. crassis.

Piper tajumulcoanum Trelease & Standley, sp. nov.

Known only from the type, San Marcos, southern slopes of
Volcan de Tajumulco, above Finca El Porvenir on Todos Santos
Chiquitos, 1,300-1,500 meters, Steyermark 37235 (in Herbarium of
Chicago Natural History Museum).

328 FIELDIANA: BOTANY, VOLUME 24

A very slender shrub 1-1.5 meters high, glabrous throughout or nearly so,
the internodes short or elongate; petioles very short, rather stout, 4 mm. long or
shorter, sparsely short-hirtellous or glabrous; leaf blades very thin, drying dull
green, minutely pellucid-punctate, lanceolate or ovate-lanceolate, 8.5-12.5 cm.
long, 2.5-4.5 cm. wide, very narrowly long-attenuate, oblique and conspicuously
unequal at the base, shallowly and narrowly cordate, the two auricles broadly
rounded, glabrous, paler beneath, palmately 3-nerved or obscurely 5-nerved, the
nerves very slender, the veins inconspicuous, divergent from the costa at a wide
angle; peduncles opposite the leaves, almost filiform, almost 2 cm. long, glabrous;
spikes small, probably pendulous, pale green, about 2 cm. long, in fruit 3 mm.
in diameter, the bracts densely hirtellous on the margins; fruit globose, 1.5 mm.
long, glabrous or obscurely granular, the style very short and thick.

Frutex gracillimus fere omnino glaber; folia brevissime petiolata tenuiter
membranacea in sicco non fuscescentia, petiolo breviter hirtello vel glabrato,
minute pellucido-punctata, lanceolata vel ovato-lanceolata, anguste longissime
attenuato-acuminata, basi inaequali breviter angusteque cordata, glabra, subtus
paullo pallidiora, palmatim 3-nervia vel obscure 5-nervia; pedunculi oppositifolii
fere filiformes 2 cm. longi, spicis subaequilongis in statu fructifero 3 mm. diam.
obtusis; fructus globosus glaber vel obscure granulosus, stylo brevi.

In general appearance much like P. Standleyi Trelease, in which
the branches and leaves are abundantly pubescent.

Piper tecutlanum Trelease & Standley, sp. nov.

Moist or wet, mixed, mountain forest, sometimes on white-sand
slopes, 1,500-2,500 meters; endemic; El Progreso (Sierra de las
Minas) ; Quezaltenango (region below San Martin Chile Verde) ; San
Marcos (Volcan de Tajumulco; type collected between town of
Tajumulco and Tecutla, Steyermark 36753; in Herbarium of Chicago
Natural History Museum).

A sparsely branched, erect shrub 2.5-4.5 meters high, branches rather slender,
densely villous-pilose with soft spreading brownish multi cellular hairs; petioles
stout, 1-3 cm. long, densely villous-pilose with spreading brownish hairs, not
winged; leaf blades thin or rather thick, more or less fuscescent when dry, strongly
and closely bullate, at least when mature, densely and minutely pellucid-punctate,
broadly elliptic or ovate-elliptic, mostly 13-21 cm. long and 8-14 cm. wide,
abruptly acute or acuminate, unequal and somewhat oblique at the base, cordate
or subcordate on one or both sides, sometimes merely rounded on both sides,
villous-pilosulous above along the costa and nerves, elsewhere glabrous, dull,
the nerves and veins much impressed, slightly paler and often brownish beneath,
densely pilose on the nerves and veins with long soft spreading hairs, penninerved,
the nerves 5-7 on each side, slender, ascending at an angle of about 45 degrees,
slightly arcuate or almost straight, the veins very closely reticulate, the surface
deeply areolate between the veins; peduncles stout, about 1.5 cm. long, densely
villous-pilose, opposite the leaves; spikes in fruit as much as 13 cm. long and 9 mm.
thick, pendent, acutish or short-mucronate at the apex; bracts triangular-peltate
at the apex, densely sordid-pubescent; fruits glabrous.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 329

Frutex, ramis dense pilis patentibus brunnescentibus villoso-pilosis; petioli
crassi 1-3 cm. longi dense villoso-pilosi; lamina tenuis vel crassiuscula in sicco
fuscescens dense minute pellucido-punctata, dense arete bullata, late elliptica
vel ovato-elliptica, magna, abrupte acuta vel acuminata, basi inaequali atque
obliqua utroque latere cordata vel subcordata vel interdum altero latere rotundata,
supra ad costam nervosque villoso-pilosula, nervis venisque manifeste impressis,
subtus ad nervos venasque dense pilosa, penninervia, nervis utroque latere 5-6;
pedunculi oppositifolii 1.5 cm. longi, spicis in statu fructifero 13 cm. longis 9 mm.

Piper telanum Trelease in Standl. Field Mus. Bot. 10: 161.
1931. Cordoncillo.

Moist or wet, mixed, lowland forest, sometimes in Manicaria
swamps, 300 meters or lower; Pete'n (near Chinaja); Alta Verapaz
(region of Cubilgiiitz); Izabal; Huehuetenango (near Ixcan). South-
ern Mexico; Honduras (the type from Tela).

Plants herbaceous or frutescent, 1-2.5 meters high, erect, sparsely branched,
glabrous throughout; petioles rather stout, about 5 mm. long, somewhat dilated
and sheathing at the base; leaf blades narrowly lanceolate or oblong-lanceolate,
mostly 13-16 cm. long and 2.5-5.5 cm. wide, attenuate-acuminate, very acute
and slightly unequal at the base, finely pellucid-punctate, dull and usually blackish
when dried, dark green on the upper surface, somewhat paler and usually brownish
beneath, penninerved, the nerves about 6 on each side, arcuate-ascending, the
veins laxly reticulate, almost obsolete or at least inconspicuous; spikes opposite
the leaves, the peduncles rather stout, 7 mm. long or shorter, the spikes very dense,
obtusely pointed at the apex, mostly 12-23 mm. long, 3 mm. thick in anthesis but
in fruit as much as 6 mm.; bracts densely pubescent on the margins; fruits sub-
globose, about 2 mm. in diameter, glabrous, granular, abruptly short-rostrate at
the apex.

A well-marked species, related to the Costa Rican P. candelari-
anum C. DC., which has uniformly broader leaves and longer flower
spikes. It has been reported from Guatemala as P. lanceolatum
Ruiz & Pavon.

Piper tuberculatum Jacq. Icon. PI. Rar. 2: 2. pi. 210. 1786.
P. tuberculatum var. obtusifolium C. DC. in Donn. Smith, Bot. Gaz.
19: 8. 1894 (type from Rio Ocosito, Quezaltenango, J. D. Smith
2592). Cordoncillo; Cordoncillo negro.

Moist or wet thickets or forest, often in rather dry, lowland forests
or thickets, or in rocky places, 1,000 meters or lower, or perhaps
at somewhat higher elevations; Pete'n; Alta Verapaz; Baja Verapaz;
Izabal; Zacapa; Jalapa; Jutiapa; Santa Rosa; Escuintla; Guatemala;
Suchitepequez; Retalhuleu; Quezaltenango; San Marcos; Huehue-
tenango; Quiche". Southern and western Mexico; British Honduras
to El Salvador and Panama; South America.

330

FIELDIANA: BOTANY, VOLUME 24

Usually a shrub of 1.5-3 meters, densely branched, the branches slender,
finely and densely puberulent; leaves firm-membranaceous, short-petiolate, oblong
or ovate-oblong, mostly 7-14 cm. long and 2.5-7 cm. wide, obtuse or rounded at
the apex or sometimes acute, cordate at the base and very unequal, one of the
lobes 5-12 mm. longer than the other, the lobes short and broadly rounded,

FIG. 45. Piper tuberculatum. Habit of portion of flowering branch; X 3 A-

equally penninerved, with about 8 pairs of lateral nerves, deep green and some-
what lustrous above, somewhat paler and densely punctate beneath, glabrous or
nearly so on both surfaces; spikes opposite the leaves, short-pedunculate, mostly
4-8 cm. long, 2.5 mm. thick, obtuse, flowers in horizontal verticils. (Fig. 45.)

This is one of the most common and widely distributed of Central
American Piper species, and it is also one of those most easily recog-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 331

nized. While there is nothing particularly distinctive about the
leaves, at least nothing that can be described easily, once seen they
can not be confused with any other species. The plant is some-
what weedy, thriving in rather dry places, and in some areas, such
j as the lowland forests toward San Jose* el Puerto, becoming the
dominant shrub in thickets and forests. It is a superior hedge plant,
iand is much cultivated for this purpose in the Pacific bocacosta,
i especially about dwellings and along the borders of cafetales. The
shrubs grow readily from cuttings, and their foliage becomes very
dense if they are pruned severely. This pruning is generally done
with wide sweeps of a machete. The specific name alludes to the
fact that the branchlets often bear tubercle-like lenticels. Called
"Spanish elder" in British Honduras.

Piper Tuerckheimii C. DC. ex Bonn. Smith, Enum. PL Guat.
2: 96. 1891, nomen; in Bonn. Smith, Bot. Gaz. 19: 258. 1894.

Moist or wet, dense, mixed forest, 1,150 meters or lower; Alta
Verapaz (type from Pansamala, Turckheim 1038); Izabal. British
Honduras; Atlantic lowlands of Honduras.

An erect shrub 1.5 meters high or less, sparsely branched, or the younger
plants often sprawling over the ground, the branches slender, strongly nodose,
densely sordid-hirtellous, with short or elongate internodes; leaves peltate well
above the base, rather thick, usually drooping, on stout hirtellous petioles 4.5 cm.
long or shorter, oblong-ovate, mostly 16-25 cm. long and 6.5-10 cm. wide, narrowly
very long-acuminate, subcordate or rounded at the base, usually blackening when
dried, glabrous above, smooth, the nerves and veins not elevated, villosulous or
hirtellous beneath, especially on the nerves and veins, slightly rough to the touch,
palmately few-nerved at the base, penninerved above the base with usually 3
nerves on each side, the veins inconspicuous, very openly reticulate; peduncles
solitary opposite the leaves, usually much shorter than the petioles, hirtellous;
spikes 5 cm. long or shorter, with a naked tail-like tip, the bracts triangular-
peltate, hirtellous on the margins; stamens 4; fruit subglobose, hirtellous at the
apex.

One of the most distinct species of the genus, easily recognized
among Central American ones by its large, conspicuously peltate
leaves, abundantly pubescent beneath, at least on the nerves.

Piper umbellatum L. Sp. PI. 30. 1753. Heckeria umbellata
Kunth, Linnaea 13: 569. 1839. Pothomorphe umbellata Miq. Comm.
Phyt. 36. 1840. Santa Maria; Jute; Obet, Obbel (Coban, Quecchi).

Moist or wet forest and thickets, often in second growth, 1,500
meters or lower; Alta Verapaz; Chiquimula; Jalapa; Jutiapa;
Escuintla; Guatemala; Sacatepe"quez; Chimaltenango; Suchitepe-

332 FIELDIANA: BOTANY, VOLUME 24

quez; Retalhuleu; Quezaltenango; San Marcos. Southern Mexico;
British Honduras to Salvador and Panama; West Indies; South
America.

Plants erect, usually 1-1.5 meters high, sparsely branched, herbaceous almost
throughout but often somewhat woody below, the young branches densely villous-
pilose; petioles 20 cm. long or shorter, vaginate for part of their length; leaf blades
thin and flaccid, drying green, ovate-orbicular, mostly 20-30 cm. long and of
equal or even greater breadth, acute or abruptly short-acute, deeply and usually
narrowly cordate at the base, with large rounded basal lobes, green on the upper
surface, glabrous to densely villosulous, somewhat paler beneath, sparsely or
densely pubescent or villosulous, pellucid-punctate, palmately nerved, with about
13 nerves; spikes pale green or whitish, sometimes cream-colored, mostly 9-15 cm.
long and about 4 mm. thick, short-pedunculate, umbellate at the ends of the short
axillary peduncles; stamens 2; stigmas 3, sessile, recurved; fruits little more than
0.5 mm. long.

This species and P. peltatum as noted under the latter, are much
alike, except that one has peltate and the other epeltate leaves.
In Guatemala P. umbellatum is usually much more densely pubescent
than P. peltatum, but there is much variation in pubescence among
the Guatemalan specimens. Both P. umbellatum and P. peltatum
are noteworthy for the strong odor of the crushed leaves and stems,
an odor reminiscent of sarsaparilla and probably arising from the
translucent glands that are apparent when the dry leaves are viewed
against a strong light. The leaves also have a distinct flavor, and
on that account are often used for flavoring meat stews and other
dishes. There is a popular belief in some parts of Central America
that if the juice of the foliage is rubbed on the skin, it will prevent
attacks of redbugs or chiggers, or even of the small ticks. Wisdom
records that in the Jocotan (Chiquimula) region, a broth of jutes
or snails and Santa Maria leaves is drunk by nursing women to in-
crease the flow of milk.

Piper uspantanense C. DC. in Bonn. Smith, Bot. Gaz. 19:
6. 1894. Cordoncillo.

Moist or wet, mixed, mountain forest, 1,000-1,650 meters; so
far as known, endemic; Alta Verapaz; Quiche* (type from San
Miguel Uspantan, Heyde & Lux 3827); Quezaltenango; San Marcos.

A rather slender shrub of 1.5-3 meters, the young branches densely hirtellous,
the internodes short or elongate; petioles stout, 10 mm. long or shorter, hirtellous,
vaginate only at the base; leaf blades thin, usually grayish green when dried, very
inconspicuously if at all punctate, elliptic-lanceolate to lanceolate or oblong-
lanceolate, mostly 14-20 cm. long and 5.5-7.5 cm. wide, narrowly long-acuminate,
slightly or strongly unequal at the base, usually acute or subacute on both sides
but rarely obtuse, glabrous and smooth above, the nerves scarcely elevated,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 333

somewhat paler beneath, usually densely and softly pilosulous beneath over the
whole surface and on the nerves but sometimes glabrate in age, penninerved,
the nerves 3-4 on either side, arising at or below the middle of the blade, arcuate-
ascending, little elevated, the veins inconspicuous but often pale, very laxly
reticulate; peduncles rather slender, slightly exceeding the petioles or often shorter,
hispidulous; spikes very slender, mucronate at the apex, erect, mostly about 7-8
cm. long and 2 mm. thick; bracts densely hirsutulous on the margins; stamens 4;
ovary glabrous; stigmas 3.

Piper vaccinum Standl. & Steyerm., sp. nov.

Known only from the type, British Honduras, on hillside, Vaca,
El Cayo District, P. H. Gentle 2264 (in Herbarium of Chicago
Natural History Museum; duplicates in Herbarium of University
of Michigan).

A slender shrub, the branches finely puberulent or in age glabrate, the inter-
nodes mostly elongate; leaves on slender petioles 1-1.5 cm. long, thin, green when
dried, minutely pellucid-punctate, the petioles puberulent; leaf blades elliptic-
ovate, mostly 9-11 cm. long and 4.5-5.5 cm. wide, rather abruptly and narrowly
long-acuminate, obtuse or rounded and slightly oblique at the base, almost glabrous
above, puberulent along the nerves, somewhat paler beneath, densely and finely
puberulent along the nerves, sparsely and minutely puberulent or glabrate between
them, 5-nerved, the veins few, laxly reticulate; peduncles slender, 6-7 mm. long,
densely puberulent; spikes very slender, spreading, 5-7.5 cm. long, 2 mm. thick,
very densely flowered; bracts densely ciliate.

Frutex gracilis, ramis minute denseque puberulis vel serius glabratis, internodiis
plerumque elongatis; folia graciliter petiolata tenuia in sicco viridia dense minute
pellucido-punctata, elliptico-ovata, subabrupte angusteque longiacuminata, basi
obtusa vel rotundata atque subobliqua, supra fere glabra ad nervos puberula,
subtus aliquanto pallidiora ad nervos dense puberula, 5-nervia; pedunculi 6-7 mm.
longi dense puberuli, spicis gracilibus ca. 5-7.5 cm. longis 2 mm. crassis, bracteis
dense ciliatis.

Piper variabile C. DC. ex Donn. Smith, Enum. PI. Guat. 2:
66. 1891, nomen; in Donn. Smith, Bot. Gaz. 19: 259. 1894. P.
purulhanum C. DC. 70: 169. 1920 (type from Purulha, Baja Verapaz,
Turckheim 11.1705). P. middlesexense Trelease ex Standl. Field
Mus. Bot. 12: 103. 1936, without Latin description (type from
Middlesex, British Honduras, W. A. Schipp 286). Cordoncillo.

Moist or wet thickets or forest, sometimes in rather dry places,
1,600 meters or lower; Pete"n; Alta Verapaz (type collected near
Coban, Turckheim 434); Baja Verapaz; Izabal; Escuintla; Guate-
mala; Retalhuleu; Quezaltenango; San Marcos. Veracruz; British
Honduras.

An erect shrub or small tree, usually 1-3 meters high, sparsely branched,
glabrous throughout or nearly so; petioles rather stout, 1.5-5 cm. long or some-

334 FIELDIANA: BOTANY, VOLUME 24

times 10 cm. or more, vaginate only at the base; leaf blades epunctate or nearly
so, rather thick and firm, usually not at all blackish when dried, broadly ovate-
cordate to ovate-orbicular, mostly 13-24 cm. long and 7-18 cm. wide, abruptly
acute or acuminate, shallowly or deeply cordate at the base, or in the upper leaves
truncate or rounded, with broadly rounded basal lobes, usually palmately 7-nerved
at the base, the nerves slender but prominent, the costa emitting 1-2 strong
nerves on each side slightly above the base of the blade, near the middle, or even
higher, the veins barely prominulous, very laxly reticulate, not conspicuous;
peduncles opposite the leaves, rather stout, mostly. 1-1.5 cm. long; spikes whitish,
long and slender, pendent or recurved, commonly 9-18 cm. long, 3-4 mm. thick,
very obtuse; bracts glabrous; stamens 3; fruits small, glabrous.

The material referred here is somewhat variable, as the specific
name might suggest, but there is no obvious basis for dividing it.
It is quite possible that the proper name for this plant is P. Schiede-
anum Steud., based on Mexican plants.

Piper vergelense Trelease & Standley, sp. nov.

Dense, wet, mixed, mountain forest, or often on forested plains,
120-1,200 meters; endemic; Santa Rosa; Escuintla; Retalhuleu;
San Marcos (type from Finca Vergel, near Rodeo, Standley 68950;
in Herbarium of Chicago Natural History Museum).

A slender shrub 1.5-3 meters high, glabrous throughout, the internodes short
or often much elongate; leaves medium-sized, membranaceous, drying green, on
stout petioles 2-3 mm. long, narrowly oblong or oblong-lanceolate, mostly 8-15 cm.
long and 2.5-5 cm. wide, somewhat abruptly long-acuminate, slightly unequal at
the base, rounded or obtuse on both sides, palmately 3-nerved, with two very
irregular and obscure additional marginal nerves, epunctate, deep green above,
paler beneath, the veins prominulous, inconspicuous, very laxly reticulate; young
and immature spikes slender, pendent, about 4 cm. long, 2 mm. thick, the peduncles
almost filiform, about 1.5 cm. long; bracts large, glabrous.

Frutex gracilis omnino glaber; folia modica membranacea brevissime petiolata
anguste oblonga vel oblongo-lanceolata abrupte longiacuminata, basi paullo
inaequalia, utrinque obtusa vel rotundata, palmatim 3-nervia, venis prominulis
inconspicuis laxissime reticulatis; spicae juveniles gracillimae pendulae, pedunculis
fere filiformibus ca. 1.5 cm. longis; bracteae glabrae.

Piper virginicum Trelease & Standley, sp. nov.

Known only from the type, Izabal, in forest along creek, between
Virginia and Lago de Izabal, base of Montana del Mico, at or little
above sea level, Steyermark 38812 (in Herbarium of Chicago Natural
History Museum).

A rather slender shrub 1.5-2.5 meters high, glabrous throughout; petioles
stout, 8-15 mm. long, vaginate only at the base; leaf blades thick and stiff, pale
green when dried, opaque or nearly so, oblong-lanceolate or ovate-lanceolate,
mostly 15-18 cm. long and 5.5-7 cm. wide, narrowly and gradually long-acuminate,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 335

equal at the base, symmetric or sometimes oblique, rounded, dull or scarcely
lustrous, slightly paler beneath, the veins and nerves prominent on both surfaces,
the blades penninerved, the nerves usually 6 on each side, ascending at a rather
wide angle, strongly arcuate, the veins closely reticulate; spikes (only very young
ones seen) on stout peduncles 6-7 mm. long, erect, the longest ones seen 4.5 mm.
long and 2 mm. thick, but doubtless much longer when fully developed; bracts
pale green, glabrous, minutely ciliate.

Frutex erectus, internodiis superioribus brevibus, omnino glaber; folia 8-15
mm. longe petiolata in sicco pallide viridia subcoriacea rigida subtus pallidiora
fere omnino opaca, oblongo-lanceolata vel ovato-lanceolata, magna, sensim
longiacuminata, basi aequalia symmetrica vel obliqua rotundata, penninervia,
nervis utroque latere vulgo 6 arcuatis, utrinque prominentibus, venis utrinque
prominentibus arete reticulatis.

Piper xanthostachyum C. DC. Anal. Inst. Fis. Geogr. Costa
Rica 9: 169. 1897. P. flavirameum C. DC. Bot. Gaz. 70: 181. 1920.
P. bryogetum C. DC. op. cit. 175. 1920.

Usually on tree trunks in moist or wet, dense, mixed forest, 1,400-
2,850 meters; Alta Verapaz; Zacapa; Quezaltenango; San Marcos.
Chiapas; Costa Rica.

Plants almost always epiphytic, rarely terrestrial and then low shrubs, usually
creeping or scandent on tree trunks, glabrous throughout or nearly so, the branches
conspicuously nodose, with short or elongate nodes, commonly glabrous but
sometimes sparsely hirtellous; stipules often conspicuous on young branches,
narrow, as much as 3 cm. long or even longer, caducous; petioles stout, usually
very short and often scarcely exceeding the basal lobes of the leaf; leaf blades
subcoriaceous, drying greenish or rarely blackish, epunctate or nearly so, lance-
oblong or ovate-oblong, mostly 11-24 cm. long and 4.5-10 cm. wide, narrowly
long-acuminate, very oblique and unequal at the base, rounded on one side and
cordate on the other or more or less cordate on both sides, penninerved, with
usually 3 pairs of nerves, or sometimes with as many as 5 nerves on one side,
the nerves usually all arising below the middle of the blade, strongly elevated
beneath, ascending at a narrow angle, slightly arcuate, the veins rather closely
reticulate, the leaves usually lustrous on both surfaces, little paler beneath but
sometimes brownish, the veins often very prominent beneath, the nerves frequently
conspicuously impressed on the upper surface; peduncles stout, mostly 1-1.5 cm.
long, much exceeding the petioles; spikes reddish brown or greenish, mostly 5-9
cm. long, 4 mm. thick or more, very obtuse; bracts hirtellous on the margins;
stamens 4; stigmas 4; ovary glabrous.

The Guatemalan material we place here was referred by Trelease
to eleven unpublished species. The species, as here treated, is
variable in shape, size, and texture of the leaves, but it is believed
that some or most of these differences may be explained on the
basis of the stage of their development, the leaves of young sterile
plants being thin while those of mature plants are more or less
coriaceous. It is, of course, possible that we have erred in uniting

336 FIELDIANA: BOTANY, VOLUME 24

the three Costa Rican species and combining with them the several
Guatemalan forms, but we believe that ultimately at least they will
have to be united, unless better characters are found than are now
apparent for separating them.

Piper yucatanense C. DC. Linnaea 37: 334. 1871-73.

Moist or wet forest, little above sea level; Pete"n. Yucatan
Peninsula of Mexico; British Honduras.

A slender shrub of 1-3 meters, glabrous throughout; leaves on very short,
slender petioles, oblong-ovate, mostly 10-16 cm. long and 3-6 cm. wide, narrowly
cuspidate-attenuate, obliquely rounded at the base, rather thin, usually blackening
when dried, epunctate, palmately 6-7-nerved, the inner lateral nerves extending
to the apex of the blade, very slender, the veins very slender, little elevated, laxly
reticulate; inflorescences opposite the leaves, on slender peduncles 10-13 mm. long,
the flowers racemose, the racemes lax, 2-3.5 cm. long, the pedicels divaricate,
2-2.5 mm. long; stamens 5, minute, the anthers globose; fruit glabrous, ovoid-
oblong, the stigmas 3-5, minute.

This species is referable to the genus Arctottonia proposed by
Trelease, but the transfer of this name has not been made, and need
not be. Schipp describes the inflorescences as cream-colored.

Piper yzabalanum C. DC. ex Donn. Smith, Enum. PL Guat.
2: 66. 1891, nomen; in Donn. Smith, Bot. Gaz. 19: 7. 1894. Tzakisba
(Huehuetenango) .

Moist or wet, mixed, mountain or lowland forest, ascending from
sea level to rarely as high as 2,000 meters; Pete"n; Alta Verapaz;
Izabal (type collected at the mouth of Rio Polochic, J. D. Smith
1712); El Progreso; Huehuetenango. Veracruz and probably else-
where in southern Mexico.

A shrub 1-3 meters high, sparsely branched, glabrous throughout; petioles
stout, 2.5-4 cm. long, vaginate throughout; leaf blades broadly ovate to oval
or elliptic, mostly 18-21 cm. long and 8-12 cm. wide, abruptly acute or short-
acuminate, broadly rounded at the base and equal or only slightly asymmetric,
most often shallowly and narrowly cordate but often not at all cordate, some-
what lustrous on both surfaces when dried, grayish green or often somewhat black-
ish, penninerved, with 5-7 nerves on each side, these arcuate, ascending at an angle
of 45 degrees or more, slender, prominent beneath, the veins inconspicuous, laxly
reticulate, the blades densely and very finely pellucid-punctate; spikes on stout
peduncles about 1 cm. long, erect, whitish or greenish white, 4.5-8 cm. long,
4-5 mm. thick, short-mucronate at the apex, the bracts spatulate, triangular and
inflexed at the apex, glabrous; stamens 4; fruit glabrous, the stigmas 3.

This species may not be different from P. chinantlense Mart. &
Gal. of Oaxaca and other localities in southern Mexico. We refrain
from using that name only because we have not seen type material

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 337

of the species. Trelease has indicated in the herbarium two new
species from Guatemala which we consider referable to P. yzaba-
lanum. In Veracruz this species is known as "cordoncillo" and
"acuyo cimarron."

Piper zacapanum Trelease & Standley, sp. nov.

Known only from the type, Zacapa, cloud forest in ravine
bordering Quebrada Alejandria, summit of Sierra de las Minas, 2,500
meters, Steyermark 29905 (in Herbarium of Chicago Natural History
Museum).

A shrub, the branches stout, tortuous, strongly nodose, with short internodes,
densely hispidulous with short, spreading or reflexed, sordid hairs; petioles stout,
4-7 mm. long, densely hispidulous, vaginate only at the base; leaf blades thick
and stiff, strongly rugose, grayish when dried, densely pellucid-punctate, oblong-
elliptic or lance-oblong, mostly 7-10 cm. long and 3-4 cm. wide, acute or acuminate,
conspicuously unequal at the base, subcordate on one side, rounded or obtuse on
the other, glabrous above or nearly so, sometimes with a few scattered hairs
along the costa, smooth to the touch, the veins and nerves strongly impressed,
almost concolorous, penninerved, the nerves usually 3 on each side, arcuate-
ascending at an angle of about 45 degrees, elevated and rather stout, the veins
prominent and closely reticulate, appressed-hispidulous on the nerves and veins,
glabrous or glabrate between them and coarsely granular; spikes unknown.

Frutex ramosus, ramis tortuosis crassiusculis, internodiis brevibus pilis
brevibus patentibus vel reflexis dense hispidulis; folia crasse petiolata, petiolo
dense hispidulo 4-7 mm. longo, in sicco grisea, dense pellucido-punctata, non lucida,
oblongo-elliptica vel lanceolato-oblonga, parva, acuta vel acuminata, basi valde
inaequalia, uno latere subcordata, altero obtusa vel rotundata, supra glabra vel
glabrata, insigniter bullata, nervis venisque impressis, subtus concoloria ad nervos
venasque adpresso-hispidula, penninervia, nervis utroque latere vulgo 3, venis
elevatis arete reticulatis; spicae ignotae.

CHLORANTHACEAE

Usually trees, sometimes shrubs or even herbs; leaves opposite, mostly dentate,
penninerved, the petioles connected at the base by a transverse line or connate
to form a sheath; stipules small, borne on the margins of the sheaths or at the
base of the petioles; inflorescences terminal or pseudo-axillary, the flowers spicate
or the pistillate variously paniculate or capitate; flowers unisexual, or the staminate
and pistillate coherent by their bases and falsely perfect; staminate flower without
a perianth, the stamens 1 or 3, connate to form a 3-lobate mass; filaments very
short, the anthers 2-celled, the cells adnate at the base, parallel or divergent,
longitudinally dehiscent, the lateral anthers sometimes 1-celled; perianth of the
pistillate flower adnate to the ovary, usually minutely 3-dentate at the apex;
ovary inferior or naked, 1-celled; stigma sessile, or rarely borne on a short style,
truncate, clavate, or short-linear, entire; ovule one, orthotropous, pendulous from
the apex of the cell; fruit a small drupe, ovoid or globose, the exocarp fleshy, the
endocarp crustaceous or indurate; seed with a membranaceous testa, the endosperm

338 FIELDIANA: BOTANY, VOLUME 24

abundant, fleshy; embryo minute, remote from the hilum, the cotyledons divaricate
or scarcely distinguishable.

A small family of three genera, in tropical America and Asia,
the Pacific islands, and New Zealand. A single genus is found in
America.

HEDYOSMUM Swartz

Trees or shrubs, often aromatic, the branches articulate at the nodes and
very brittle; leaves usually serrate, the petioles enlarged and connate at the base,
forming a sheath, the small stipules borne on the margin of the sheath; inflores-
cences axillary or pseudoterminal; flowers monoecious or dioecious, each inflores-
cence unisexual; staminate flowers spicate, ebracteate; anther one, subsessile,
linear or oblong, the connective short-appendaged or subpeltate above the cells;
pistillate flowers capitate or scattered, the perianth tube adnate to the ovary,
the limb very short, 3-dentate; style very short, stigmatose at the apex, or the
stigma sessile, obtuse; drupes small, globose or ovoid, the exocarp very juicy.

About twenty species, in tropical America. Three other Central
American ones are known in Nicaragua, Costa Rica, and Panama.

Hedyosmum mexicanum Cordemoy, Adansonia 3: 307. 1862-
63. H. Artocarpus Solms in DC. Prodr. 16: pt. 1: 485. 1869. Palo
de agua, Ocze (Huehuetenango) ; Te azteco; Te maya; Te de monte;
Mazorco; Onj or One (Coban, Quecchi); Macetero (fide Aguilar);
Sandio (Jalapa).

Wet mixed mountain forest, or often in pine forest, 1,200-2,900
meters; Alta Verapaz; Baja Verapaz; Izabal; Zacapa; Jalapa; Guate-
mala; Chimaltenango; Solola; Suchitepequez; Quiche"; Huehuete-
nango; Quezaltenango; San Marcos. Southern Mexico; Honduras;
Costa Rica.

Very variable in size, often blooming when only a shrub of 2 meters, but more
commonly a tree of 8-12 meters or more, with a thick trunk and dense crown;
leaves short-petiolate, the sheaths often 2.5 cm. long, funnelform; leaf blades
narrowly lanceolate or oblong-lanceolate, mostly 12-20 cm. long and 3-6 cm.
wide, acuminate, acute to attenuate at the base, finely or usually coarsely crenate-
serrate; stipules small, multilaciniate; staminate spikes cylindric and ament-like,
2-8 cm. long, 5-8 mm. thick; pistillate flowers very numerous in a dense, ovoid
or subglobose head, the fruiting head 2-3 cm. long and 1.5-2 cm. thick, or in the
fresh state larger, very fleshy and juicy, whitish at maturity. (Fig. 46.)

This species has always been called H. Artocarpus, H. mexicanum
having been placed by Solms (in DC. Prodr. 16, pt. 1: 485. 1869)
among the doubtful species, because it was based upon "fragmen-
tary" material. A photograph of the type collection of H. mexicanum
in the Delessert Herbarium, presumably the one seen by Solms, is

FIG. 46. Hedyosmum mexicanum. A. Habit of staminate branch; X %.
B. Sheath at base of leaves; X 1. C. Staminate flowers on portion of inflores-
cence of staminate branch; X 2. D. Habit of pistillate branch; X %

339

340 FIELDI AN A: BOTANY, VOLUME 24

scarcely to be described as fragmentary, and agrees well with the
very numerous sheets of H. Artocarpus. Since only one species of
Hedyosmum has been found in recent years in Mexico, it is safe to
assume that H. Artocarpus and H. mexicanum are synonymous.
In the Coban region and especially in the mountain forests along
the barranco of the Rio Samala in Quezaltenango, this is one of the
commonest and most characteristic trees, at times almost dominating
the forest. It is strange to find low shrubs sometimes loaded with
flowers and fruits, while most of the plants are tall trees. The brittle
branches are very easily detached. The leaves, which have a dis-
tinctive aromatic odor, turn black as they wither. The fruits at
maturity are very succulent, whitish, and edible, with a sweetish
flavor. In the Occidente a beverage, a substitute for coffee, is
prepared by placing two or three leaves in a cup of hot water. No
use, apparently, is made of the wood. It is of interest to note that
specimens of this tree were collected in Mexico by Sesse" and Mocino
150 years ago, and referred by them, strangely enough, to the genus
Pandanus!

LACISTEMACEAE

Shrubs or small trees; leaves alternate, short-petiolate, penninerved, without
stipules; flowers perfect, solitary within the imbricate bracts of a slender ament-
like spike, 2-bracteolate, the spikes sessile and often fasciculate in the leaf axils;
bractlets inserted at the base of the pedicel, very small, linear-subulate; perianth
none or of six or fewer unequal, linear to setaceous or spatulate scales; disk fleshy,
explanate and shallowly sinuate-lobate, sometimes concave or cupular, often
cleft on one side; stamen one, attached to the disk near its center, the filament
slightly complanate or thick; anther of two distinct, usually divaricate-stipitate,
ovoid, longitudinally dehiscent cells; ovary superior in the center of the disk,
sessile or short-stipitate, 1-celled, with three parietal placentae; stigmas three,
erect; ovules 1-2 on each placenta, anatropous, pendulous; fruit capsular, ovoid
or subglobose, usually triquetrous, 3-valvate, the valves coriaceous, the placentae
medial upon them; seeds by abortion usually one in each capsule, obo void-oblong,
pendulous, the outer coat usually fleshy; embryo straight, the cotyledons broad
and foliaceous.

The family consists of a single genus, with about twenty species,
all in tropical America. A single species is found in North America.

LACISTEMA Swartz

Lacistema aggregatum (Berg.) Rusby, Bull. N. Y. Bot. Card.
4: 447. 1907. Piper aggregatum Berg. Act. Helv. 7: 131. pi. 10.
1777. L. myricoides Swartz, Prodr. Veg. Ind. Occ. 12. 1788.

FIG. 47. Lacistema aggregatum. A. Habit of portion of branch; X M-
B. Tip of inflorescence; X 8. C. Longitudinal section through young fruit;
X 5. D. Flower with bract and bracteoles; X 20. E. Pistil and stamen; X 20.
F. Pistil and stamen from the side; X 20. G. Fruiting branch with fruits; X 1.

341

342 FIELDIANA: BOTANY, VOLUME 24

Wet thickets, 300 meters or less; Pete*n; Alta Verapaz; Izabal;
Huehuetenango. Southern Mexico; British Honduras to Panama;
West Indies; South America.

A glabrous shrub or tree, sometimes 11 meters high, with a trunk 10-20 cm.
in diameter; leaves on short petioles, elliptic or elliptic-oblong, 7-15 cm. long,
abruptly acuminate, acute or obtuse at the base, thick-membranaceous, entire;
flower spikes about 1 cm. long, sessile and clustered in the leaf axils, shorter than
the petioles; capsule ovoid, reddish, 6-7 mm. long. (Fig. 47.)

The wood is brown, light in weight, tough, and fibrous, somewhat
suggestive of willow (Salix). So far as known, no use is made of it
in Central America. The tree has no outstanding characters by which
it may be recognized easily, unless it be the clusters of small, catkin-
like, bracted inflorescences, which are rather distinctive when their
appearance is once known. It is called "palo mulato" in British
Honduras.

SALICACEAE. Willow Family

Trees or shrubs with bitter bark; leaves alternate, with stipules, entire or
usually serrate; flowers minute, dioecious, in aments, often appearing before the
leaves, the individual flowers solitary in the axils of bracts; staminate flowers
consisting of 1-many stamens inserted on the receptacle and subtended by a gland-
like or cup-shaped disk; anthers 2-celled, the cells longitudinally dehiscent;
pistillate aments sometimes raceme-like, the flower consisting of a 1-celled ovary
subtended by a minute disk; placentae 2-4, parietal, the ovules usually numerous,
anatropous; stigmas 2-4, simple or 2-4-cleft; seeds small or minute, bearing a
dense coma of long, soft, silky, usually white hairs; endosperm none, the coty-
ledons plano-convex.

The family consists of two genera and about 250 species, mostly
in the north temperate and arctic zones. Only one genus is native
in tropical America.

Disk of the flower cup-like, entire or lobate; leaves broad; aments, at least the
staminate ones, pendulous Populus.

Disk composed of 1-2 glands, these distinct or scarcely connate at the base; leaves
linear or lanceolate; aments usually erect Salix.

POPULUS L.

Trees, the bud scales usually resinous; petioles often compressed, the blades
usually ovate or broader, penninerved and 3-nerved from the base, usually dentate,
sometimes lobate; stipules narrow, fugacious; aments lax, the flowers often short-
pedicellate; disk fleshy or membranaceous, shortly or broadly cyathiform, often
oblique, entire, dentate, or irregularly lobate; stamens 4-30, the filaments filiform;
ovary sessile within the disk, with usually 2-3 placentae; style short, the branches
as many as the placentae, entire or 2-fid; ovules usually very numerous; capsule
2-3-valvate, small; seeds minute, usually with a long dense white coma of soft
hairs.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 343

About thirty species, natives of the northern hemisphere. Several
species are native in Mexico, but none can be expected to extend to
Central America.

Populus alba L. Sp. PI. 1034. 1753. Alamo; Alamo bianco; Coca
(Occidente).

Planted commonly in the central and especially the western
highlands, in parks and along streets and roads. Native of Europe
and Asia, naturalized in some parts of the United States, and perhaps
in Mexico.

A large or medium-sized tree with smooth, light gray bark; leaves slender-
petiolate, broadly ovate to suborbicular, 5-10 cm. long, acute, truncate or cordate
at the base, glabrate and dark green above, beneath often densely white-tomentose,
at least when young, irregularly dentate and often shallowly lobate; staminate
aments pendulous, 5-10 cm. long.

While this species may be seen in large numbers in some of the
central regions, as about Chimaltenango, it is by far most common,
and often a conspicuous feature of the landscape, in the Occidente,
about Quezaltenango and San Marcos, where there are many large
trees. The finest planting is the long avenue called La Alameda,
in the city of Quezaltenango. In passing it may be noted that the
word alameda is technically a grove of alamos, i.e. Populus alba, but
in vernacular Spanish, at least in Central America, it has come to
be applied to any ornamental grove or avenue. The white poplar
has been used to some extent by the government of Guatemala
for planting along roadsides. In the Occidente the trees are rather
untidy in the middle of the winter, half or wholly bare, the leaves
gradually shriveling and littering the ground. Pistillate trees of this
and other species of Populus are a nuisance as shade trees when
the fruits are ripening, for then the "cotton" of the seeds is wafted
by the wind in all directions, becoming attached to all sorts of objects,
entering dwellings, and even being drawn into one's nose, eyes, and
mouth. This nuisance may be avoided by planting only cuttings
from staminate trees.

SALIXL. Willow

Reference: C. Schneider, A conspectus of the Mexican, West
Indian, Central and South American species of Salix, Bot. Gaz. 65:
1-41. 1918.

Shrubs or trees with single-scaled buds, the scales with an inner adherent
membrane; leaves short-petiolate, mostly linear or lanceolate, persistent or

344 FIELDIANA: BOTANY, VOLUME 24

deciduous; stipules minute or large and broad, persistent or caducous; bracts
the aments entire; staminate aments dense, erect or spreading, rarely drooping;
stamens 1-10, usually 2, the filaments distinct or sometimes united; pistillate
aments commonly erect or spreading; ovary sessile or short-stipitate, the style
short or filiform; stigmas 2, entire or 2-cleft; capsule usually 2-valvate.

About 200 species, almost all in the north temperate and arctic
zones, only a few extending to the tropics or to the southern hemi-
sphere. Only the following species are known in Central America.

Leaves densely sericeous, mostly 1.5-3 cm. long; stamens 2 S. taxifolia.

Leaves glabrous or sparsely pubescent, mostly more than 3 cm. long.

Leaves very glaucous beneath S. laevigata.

Leaves not glaucous beneath.

Stamens 3-7; native trees; leaves linear or lance-linear S. chilensis.

Stamens 2; introduced cultivated trees or shrubs.

Leaves lanceolate; branches not recurved and pendulous S. alba.

Leaves lance-linear; branches recurved and pendulous S. babylonica.

Salix alba L. Sp. PL 1021. 1753.

Collected in cultivation and perhaps naturalized in fence-rows
in Quezaltenango, near the city of Quezaltenango and San Mateo,
2,280-2,500 meters. Native of Europe, naturalized in eastern
United States and Canada.

A large shrub or a tree with rough gray bark; leaves short-petiolate, lanceolate
or oblong-lanceolate, mostly 5-10 cm. long and 8-15 mm. wide, acute or acuminate,
narrowed to the base, serrulate, when young often sparsely sericeous but soon
glabrate or glabrous, rather pale green beneath but not glaucous in the Guatemalan
form; stipules ovate-lanceolate, deciduous; aments on short leafy branches;
stamens 2; pistillate aments 3.5-6 cm. long; stigmas almost sessile; capsule short-
pedicellate or sessile, ovoid, acute, glabrous.

At the time of collection, it was not noted that the trees were
unlike the native willows of Guatemala, and so no special search
was made for other trees, which may be of wider distribution in the
Occidente. It would be of interest to know how this species, un-
known elsewhere in Central America or Mexico, reached Quezal-
tenango.

Salix babylonica L. Sp. PL 1017. 1753. Sauce lloron.

Not common in Guatemala, but occasionally planted in the
central and western highlands, especially about Quezaltenango and
in Chimaltenango. Native of Asia.

A medium-sized tree with rough gray bark, the twigs very slender and elongate,
drooping; leaves short-petiolate, lance-linear, 8-15 cm. long, 6-12 mm. wide, long-
acuminate, serrulate, glabrous at maturity, slightly paler beneath but not glaucous;

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 345

ments appearing on short leafy lateral branches; stamens 2; style almost obsolete;
capsule ovoid, sessile, glabrous.

The finest planting of the weeping willow in Guatemala is along
the eastern entrance to Quezaltenango, where there is a double
avenue of large and well-grown trees. There are also trees about
San Juan Ostuncalco (Quezaltenango) and many handsome ones
about Tecpam (Chimaltenango), especially near the Molino Helvetia.
Scattered individuals are found in other parts of the highlands.

Salix Bonplandiana HBK. Nov. Gen. & Sp. 2: 20. pi 101.
1817. Sauce.

Chiefly in thickets along streams, 1,350-2,800 meters; Alta
Verapaz; Baja Verapaz; Jutiapa; Sacatepe"quez; Chimaltenango;
Solola; Huehuetenango; Quezaltenango; San Marcos. Arizona and
New Mexico; Mexico.

A shrub or small tree, sometimes 12 meters high but usually lower, the trunk
sometimes 40 cm. in diameter, the bark brown, irregularly fissured; branchlets
slender, red-brown, often somewhat drooping; leaves slender-petiolate, narrowly
lanceolate, 5-15 cm. long, mostly 1.5-2.5 cm. wide, serrulate, long-acuminate or
attenuate, obtuse or acute at the base, bright green above, very glaucous beneath,
glabrous or nearly so; capsule glabrous.

The species is abundant in many places in the Occidente. Worthy
of special note in connection with the willows is a large thicket
many acres in extent, beside the river at Chiantla (Huehuetenango),
where all three of the native Guatemalan species are found together
in a very dense growth. Usually in Guatemala the species are
found singly. S. Bonplandiana has been much planted in San Marcos
along the Carretera Internacional. The shrubs were planted in
dense rows, principally for ornament, it is presumed, but perhaps
partly to prevent erosion. The soil is poor and the rainfall insufficient,
so that the bushes have not thrived, although they have continued
to grow slowly. In some parts of San Marcos there are hedges,
which are of only fair appearance from an ornamental standpoint.
S. Bonplandiana has been reported from Guatemala under the name
S. laevigata Bebb. Dr. Raup has determined all the Guatemalan
specimens of S. Bonplandiana as S. laevigata.

Salix chilensis Molina, Sagg. Storia Nat. Chil. 169. 1782.
S. Humboldtiana Willd. Sp. PL 4: 657. 1805. S. stipulacea Mart.
& Gal. Bull. Acad. Brux. 10, pt. 1: 343. 1843. S. Humboldtiana var.
stipulacea C. Schneid. Bot. Gaz. 65: 7. 1918. Sauce; C'os (Poconchi);
Saccos (Quiche", Totonicapan, fide Tejada) ; Chicaj (Huehuetenango,
fide Tejada).

FIG. 48. Salix chilensis. A. Habit of portion of staminate branch and leaves
of staminate plant; X %. B. Staminate flower; X 8. C. Portion of leaf margin;
X 2. D. Pistillate flower; X 5. E. Stamen; X 9.

346

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 347

Along streams or in swamps and marshes, abundant in many
regions, ascending from sea level to about 1,900 meters; Pete*n; Alta
Verapaz; Baja Verapaz; Izabal; Zacapa; El Progreso; Jalapa;
Jutiapa; Santa Rosa; Escuintla; Guatemala; Sacatepe*quez; Retal-
huleu; Huehuetenango; Quezaltenango; San Marcos; probably in
all the departments, except perhaps Totonicapan. Mexico and
British Honduras to Panama, southward to Argentina.

A shrub or usually a medium-sized or rather small tree, sometimes 18 meters
high, the branches brown or yellow-brown, slender and often drooping; leaves
short-petiolate, linear or lance-linear, 6-15 cm. long, mostly 3-10 mm. wide,
long-attenuate, acute at the base, serrulate, bright green, glabrous; capsules
short-stipitate, ovoid, glabrous. (Fig. 48.)

According to Schneider, all the Central American material is
referable to var. stipulacea, the typical form being South American,
but the variety is based upon rather feeble characters. It is of
interest that this is probably the only species of Salix in South
America, where it has a wide range. A few others have been de-
scribed, but all probably are mere forms of S. chilensis. This willow,
closely related to the black willow (S. nigra Marsh.) of the United
States and northern Mexico, is very common in many parts of Guate-
mala and is certainly native in the country. Its range, however, is
curious, for few are the native trees that abound in both the hot
plains and the cool highlands. S. chilensis is particularly abundant
along the Motagua Valley, forming forests in its lower stretches in
the banana country, and it is sometimes planted as a shade tree
in parks and elsewhere. There is some question as to whether the
tree is really native in some regions of southern Central America.
In Costa Rica, for instance, it is claimed that all the trees are pistil-
late, but this is not the case in Guatemala. The bark is light brown
with coarse scales, the inner bark medium brown; the sapwood is
white, and the heartwood pinkish or dark brown. Little use is
made of the wood locally, but the slender flexible tough branches are
used in Guatemala for making baskets and wicker furniture. The
tree keeps its leaves for all or most of the year and is always green
and handsome. The individual trees vary greatly in appearance.
When they have plenty of space in which to grow, they have broad
and ample crowns; when crowded they grow very tall and narrow,
but are not columnar. Worthy of particular note is the columnar
form of S. chilensis, which in habit is like the Lombardy poplar
(Populus italica Moench), i.e. with a tall and massive, pole-like
trunk and suberect branches that form a narrow, column-like crown.
This species is common in central Mexico, and has been introduced

348 , FIELDIANA: BOTANY, VOLUME 24

into Guatemala, where it is seen only in cultivation, and infrequently.
There are numerous trees of this form at Jalapa, about Chichicas-
tenango, in the Coban region, and doubtless elsewhere. Such a
well-marked form should have a name by which to distinguish it,
but none seems to have been proposed. Tahuesco, an aldea of
Suchitepequez, is said to derive its name from two Nahuatl words,
tlalli, "land" or "region," and huexotl, "willow."

Salix taxifolia HBK. Nov. Gen. & Sp. 2: 18. 1817. S. micro-
phylla Schlecht. & Cham. Linnaea 6: 354. 1831. S. taxifolia var.
microphylla C. Schneid. Bot. Gaz. 65: 24. 1918. Sauce; Ispac bianco
(Huehuetenango, fide Skutch).

Thickets along streams, 1,200-2,000 meters; Chiquimula (near
Concepcion de las Minas); Guatemala; Huehuetenango. South-
western United States and Mexico.

A shrub or small tree, usually 2-3 meters high; leaves linear or lance-linear,
in the Guatemalan form mostly 1-1.5 cm. long, almost sessile, densely silvery-
sericeous on both surfaces, entire or practically so; aments small, oblong, dense,
the pistillate ones few-flowered; capsule sessile or nearly so, densely sericeous.

This, the rarest of the Guatemalan species, is local in distribution.
It is easy of recognition by its very small, silvery leaves. The
Central American specimens are all referable to var. microphylla,
which differs but little from the typical form.

MYRICACEAE. Bayberry Family

Shrubs or small trees, aromatic; leaves alternate, coriaceous, simple, entire
or dentate, rarely lobate, resin-dotted, usually without stipules; flowers small,
monoecious or dioecious, in oblong or globose aments, the flowers solitary in the
axils of bracts; perianth none; staminate flower with 2-16 (usually 4-8) stamens
inserted on a receptacle, the filaments short, distinct or slightly united; anthers
ovoid, 2-celled, the cells longitudinally dehiscent; pistillate flower of a solitary
1-celled ovary, subtended by 2-8 bractlets; ovule solitary, orthotropous; style
very short, the 2 stigmas linear; fruit a small, oblong or globose drupe or nut,
the exocarp often waxy; seed erect, the cotyledons plano-convex.

The family consists of a single genus with the characters of the
family.

MYRICAL. Bayberry

Reference: Auguste Chevalier, Me"m. Soc. Sci. Nat. Cherbourg
32: 85-340. 1901.

The genus consists of about forty species, widely distributed in
tropical, temperate, and arctic regions. Two other species occur
in Costa Rica.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 349

Leaves acute to long-acuminate, sinuate-dentate or subentire M. cerifera.

Leaves mostly obtuse or very obtuse.

Leaves obovate, mostly 2-4 cm. long, entire or sinuate-dentate . . . . M. Pringlei.

Leaves oblanceolate-oblong, mostly 5-11 cm. long, conspicuously appressed-
serrate for almost their whole length M. Lindeniana.

Myrica cerifera L. Sp. PI. 1024. 1753. M. mexicana Willd.
Enum. PI. 2: 1011. 1809. M. xalapensis HBK. Nov. Gen. & Sp. 2:
10. 1817. Arrayan; Cera vegetal; Gua-ut (Coban, Kekchi); Tea-bark,
Tea-box (British Honduras).

Common and widely distributed, in a wide variety of habitats,
chiefly in pine forest, but often in lowland savannas, coastal thickets,
or in bogs, ascending from sea level to about 2,500 meters; Alta
Verapaz; Baja Verapaz; Izabal; Zacapa; Chiquimula; Jalapa; Guate-
mala; Escuintla (Rio Guacalate); Sacatepe*quez; Chimaltenango;
Quezaltenango; Quiche"; Huehuetenango. Southeastern United
States to Mexico, British Honduras, and Costa Rica; West Indies.

Usually a shrub of 2-4 meters but sometimes in Guatemala as much as 9
meters tall, sparsely or densely branched; leaves short-petiolate, oblong-oblanceo-
late to linear-oblanceolate, coriaceous, mostly 5-10 cm. long and 1.5-2 cm. wide,
usually acute to long-acuminate, rarely obtuse, long-attenuate to the base, sub-
entire or more often sinuate-dentate with only a few remote teeth, at least in leaves
of older branches, those of young branches sometimes coarsely serrate, glabrate
but often villous-pilose beneath along the costa, densely dotted beneath with
golden wax glands; staminate aments sessile, mostly 1 cm. long or less, the pistil-
late aments larger, laxly flowered, usually with only a few fruits; drupes globose,
3 mm. or less in diameter, covered with a thick coat of white wax. (Fig. 49.)

The Mexican and Central American species of Myrica are difficult
of delimitation, so far as the ample material available to us is con-
cerned. The only recent monograph of the American species is that
of Chevalier, which is poor so far as these species are concerned,
probably because he had little material for their study. The plant
or plants of Mexico and Central America generally have been
referred to M. mexicana, but close comparison of the many specimens
of M. cerifera, which as usually understood is rather variable, suggests
no characters by which M. mexicana and M. cerifera may be sepa-
rated. The senior author has previously referred the coastal material
from Central America to M . cerifera, but when the whole series of
material is examined, the differences between the two species are
found to be intangible. It is remarkable to find a shrub with such
a wide distribution, at least altitudinally, in Central America. The
habitats likewise are diverse, and one would not expect a bog plant
to be the same species as a plant found in the well-drained pine

FIG. 49. Myrica cerifera. A. Habit of portion of pistillate plant; X Vs.
B. Habit of portion of staminate plant; X 2 A- C. Fruit; X 8. D. Staminate
flower; X 19. E. Pistillate flower ; X 18. F. Vertical section through ovary; X 18.

350

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 351

forests. In Alta Verapaz the species is found in either habitat, in
adjoining areas.

The name "chac olol" is reported for this species from Chiapas
by Seler. Large amounts of wax are extracted from the fruits in
Guatemala and other parts of Central America, a use of the plant
well known also in the United States. The crude wax is greenish,
but can be bleached. It is used in Guatemala for making candles
that burn with a very pleasing aroma, and are used principally
in the churches or on altars in homes. These candles are a common
article in the markets, particularly those of Coban and Quezal-
tenango, and many thousands of them are burned in the processions
of Holy Week in Coban. The wax is firm and hard and on that
account the candles are especially suited to hot climates. A wax
sold in the Quezaltenango market as "brea" or "brea de Campeche,"
for use in candle-making, may be made from Myrica fruits, but this
is uncertain. Merchants said only that it was brought in by the
Indians. Myrica wax is said to be used commercially in Guatemala
for making shoe polish and in Huehuetenango for making soap.
Candles similar to the bayberry ones, and prized for the same
purposes, often are made in Guatemala from beeswax, but their
odor when burning is quite different from that of burning Myrica
wax.

Myrica Lindeniana C. DC. in DC. Prodr. 16, pt. 2: 150. 1864.
Cera vegetal.

Mostly in moist or wet, mixed forest, 1,600-2,750 meters;
Chiquimula; Huehuetenango; San Marcos. Guerrero to Veracruz,
Oaxaca, and Chiapas, the type from Chiapas.

A large shrub or a tree, sometimes 12 meters tall, with a trunk 20 cm. in
diameter, the tips of the branches usually pilose or puberulent; leaves rather large,
short-petiolate, chiefly oblanceolate-oblong, mostly 7-12 cm. long and 2-3.5 cm.
wide, commonly obtuse or often acute, cuneate-attenuate to the base, rather closely
and evenly serrate almost from base to apex, glabrate, densely yellow-punctate
beneath; aments and fruits similar to those of M. cerifera.

This plant seems to be reasonably distinct from M. cerifera, and
recognizable when a considerable number of specimens is available
for comparison. In Trees and Shrubs of Mexico it was treated as
a synonym of M. mexicana, since at that time no authentic material
was at hand for comparison. It may be, on the other hand, that it
can not be maintained as a distinct species. The serration of the
leaves is much more regular and conspicuous than in those of M.
cerifera.

352 FIELD IANA: BOTANY, VOLUME 24

Myrica Pringlei Greenm. Proc. Amer. Acad. 41: 236. 1905.

Jalapa, Cerro Alcoba, east of Jalapa, 1,300-1,700 meters, open
slopes among oak trees, Steyermark 32585. Hidalgo, Puebla, and
Oaxaca.

A densely branched shrub 1-2.5 meters high; leaves small and broad, mostly
obovate, 1.5-4 cm. long, 1-1.5 cm. wide, rounded or very obtuse at the apex, entire
or remotely and obscurely serrate, glabrous or nearly so, densely gland-dotted
beneath; aments and fruits like those of M. cerifera.

Known in Guatemala from the single collection cited, growing in
a locality far removed from the Mexican stations known for the
species. It is quite possible that further collections will show that
the plant is a distinct species, or perhaps only an aberrant form of
M. cerifera.

JUGLANDACEAE. Walnut Family

References: Wayne E. Manning, The morphology of the flowers
of the Juglandaceae: I. The inflorescence, Amer. Journ. Bot. 25:
407-419. 1938; II. The pistillate flowers and fruit, op. cit. 27:
839-852. 1940.

Trees or rarely shrubs; buds naked or scaly, often several superposed; leaves
usually deciduous, alternate, rarely opposite or whorled, estipulate, pinnately
compound; leaflets entire or serrate, glandular-dotted beneath; flowers monoecious
or rarely dioecious, staminate or pistillate flowers or both in elongate drooping
or erect catkins or spikes; staminate and pistillate inflorescences sometimes
separate, sometimes combined into an androgynous panicle, the central spike
wholly or partly pistillate, the lateral elongate or the short branches staminate;
separate staminate inflorescence a cluster of 3-8 catkins or a solitary catkin;
separate pistillate inflorescence a catkin, a few-flowered spike, or a solitary flower;
inflorescences terminal, or lateral on old wood, or the staminate sometimes lateral
at the base of new growth; staminate calyx commonly 4-lobed or lacking, the sub-
tending entire or 3-lobed bract and 2 bracteoles usually appearing as part of the
perianth; petals none; stamens 3-many; pistillate calyx usually 4-lobed or absent,
the subtending 3-lobed or entire bract and 2 or 3 bracteoles fused only with the
pedicel and base of the ovary or with the whole ovary, a ring of minute inner
bracteoles very rarely present; ovary inferior, 1-celled above, 2- to 4-celled below,
with one erect orthotropous ovule in the center at the top of the primary partial
partition; style one with usually 2 stigmatic branches; fruit a nut enclosed in a
husk, or a nutlet with 2 or 3 wings; seed solitary, large, 2-4-lobed, without endo-
sperm; cotyledons 4-lobed, oily, at germination remaining in the nut or appearing
above ground.

Seven genera are known and about sixty species, in temperate
regions of both hemispheres, in the tropics confined to mountain
regions. Four genera are represented in America, one of them, Carya,

STANDEE Y AND STEYERMARK: FLORA OF GUATEMALA 353

the hickory nuts, ranging from southern Canada to Mexico. Only
the following genera are found in Central America.

Leaflets conspicuously lepidote beneath; fruit subtended by a very large, lobate

bract Engelhardtia.

Leaflets not lepidote beneath or sometimes very sparsely and obscurely so when

very young; fruit not subtended by a large bract.

Leaves, at least most of them, opposite; staminate inflorescences erect.AZ/aroa.
Leaves alternate; staminate inflorescences pendulous Juglans.

ALFAROA Standl.
By WAYNE E. MANNING

Trees or large shrubs; leaves mostly opposite, pinnate, without a terminal
leaflet, the leaflets numerous, mostly alternate but often opposite, membranaceous,
serrate or entire, lepidote beneath on the youngest leaves but only remotely so
on the older ones; flowers monoecious or partially dioecious, the inflorescence
terminal, consisting usually of an androgynous open or spike-like panicle, the
main portion pistillate, with 30-50 flowers, bearing near the base usually 2-4
lateral short or elongate branches, these staminate, the staminate catkins some-
times forming a separate terminal panicle on different branches or on different
trees; bracts of the staminate flowers minute, 3-lobate, beneath the calyx, the
2 bracteoles and 2-4 variable sepals together appearing as an irregular 4-6 lobate
calyx, the lobes oblong, obtuse; stamens 6-10, inserted in a single series around
a naked center or rarely around a rudimentary ovary, the filaments almost obsolete;
anthers 2-celled, glabrous, dehiscent by longitudinal slits; pistillate flowers sub-
tended by a minute 3-lobate free bract shorter than the ovary; perianth deeply
4-lobate, the lobes oblong-linear, unequal, obtuse, erect, persistent upon the apex
of the fruit; style shorter than the perianth lobes, bifurcate, the stigmas sub-
globose; fruit oval or obovoid, small, the skin ("pericarp" or "exocarp") almost
dry, thin, indehiscent, adhering closely to the nut proper ("endocarp"); nut thin-
walled, smooth, loculicidally dehiscent at time of germination of the seed, falsely
8-celled in the lower half, 4-celled above the middle, 1-celled at the very apex,
the partitions nearly complete, with lamellae projecting from them into the
loculus; seeds 8-lobate to the base, each cotyledon 4-lobate.

The genus consists of a single species that has been named for
Anastasio Alfaro, a Director of the Museo Nacional de Costa Rica,
a distinguished naturalist, and a highly esteemed friend of the
senior author of this Flora.

Alfaroa costaricensis Standl. Journ. Wash. Acad. Sci. 17: 78.
1927.

Dense, moist or wet, mixed forest, 1,200-2,100 meters; Suchite-
pequez; Quezaltenango; Huehuetenango. Mountains of Costa Rica
and Panama.

A large shrub or a tree, in Guatemala 6-9 meters high or probably also taller,
the crown of rather few, spreading branches, the bark almost smooth, pale brown-

354 FIELDIANA: BOTANY, VOLUME 24

ish; branchlets and leaf rachis usually densely hirsute with long stiff spreading
hairs, sometimes merely hirtellous or in age glabrate; leaves almost all opposite,
those of a pair often unequal, or one of the leaves sometimes suppressed; leaflets
mostly 10-20, highly variable, often almost all opposite, oblong to narrowly lance-
oblong, mostly 10-18 cm. long and 1.5-4 cm. wide, the lowest leaflets generally
reduced, acute to long-acuminate or rarely obtuse, sessile or nearly so, obtuse
to truncate at the base and usually very oblique, conspicuously appressed-serrate
or almost entire, membranaceous or thicker, glabrous or nearly so along
the costa, usually glaucous or glaucescent beneath, usually hirtellous or hir-
sute along the nerves but sometimes glabrate; flower spikes stout, 3-5 cm.
long, short-pedunculate, the rachis densely hirtellous and glandular; staminate
flowers 4 mm. broad, the perianth glandular; pistillate flowers green, 5-6 mm.
long; ovary sparsely hirtellous and densely covered with golden glands; stigmas
red; fruiting spikes 12-18 cm. long or more, each bearing numerous oval or obovoid
fruits, about 2.5 cm. long and 2 cm. thick, densely velutinous-hirsute and covered
with sessile glands; nut smooth, broadly rounded at base and apex, the wall
("endocarp") less than 1 mm. thick.

The status of the Guatemalan tree referred here is somewhat
uncertain because so far only sterile collections have been made.
These are rather numerous (about ten), and the tree is plentiful in
some regions. Its habitat in Guatemala is somewhat unlike that of
the Costa Rican tree, which has been found abundantly in flower
and fruit. The tree is particularly plentiful in the mountain forest
along the old road between Finca Pirineos and Patzulin, Quezal-
tenango, and many tall trees are scattered through the forest. Pro-
tracted search failed to discover any nuts on the ground, and fallen
flowers also were lacking. It is possible that the Guatemalan tree
is actually a species of Engelhardtia (distinct fromE". guatemalensis),
but the leaves are so closely like those of Costa Rican Alfaroa that
the Guatemalan tree probably belongs at least in that genus. It is
to be expected that the Guatemalan tree represents a distinct
species, but although the leaves do not agree always in all respects
with leaves of Costa Rican material, it has been impossible to suggest
any definite character by which two species can be distinguished.
Foliage of Costa Rican trees is highly variable in pubescence, serra-
tion, and other details.

ENGELHARDTIA Lesch.

Large trees; leaves large, pinnate, the leaflets numerous, dentate or entire,
usually lepidote or resinous-punctate beneath; staminate aments solitary or
geminate or sometimes several on a short peduncle, lateral, pendulous; pistillate
spikes lateral or terminal, usually elongate and many-flowered, erect or in fruit
recurved; staminate perianth concrete with a linear or dilated bract, the margin
irregularly 3-6-lobate; stamens 3-15, biseriate, a linear ovary rudiment generally
present; pistillate bracts short-cupular at the base and adnate to the ovary, 3-5-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 355

lobate, the middle lobe longest; perianth adnate to the ovary, the limb free, short,
4-fid, the lobes subvalvate; style branches 2-4, stout, papillose-fimbriate; bract
in fruit much enlarged, coriaceous, venose, its base embracing the nut and adnate
to it; nut small, the endocarp crustaceous or indurate, variously intruded within
and forming 2-4 imperfect cells.

About a dozen species, in southeastern Asia and the islands of
the southern Pacific; in America three species are known: one in
Chiapas (E. mexicana Standl.), another, E. pterocarpa (Oerst.)
Standl., in the mountains of Costa Rica, and the following:

Engelhardtia guatemalensis Standl. Field Mus. Bot. 22: 12.
1940. Palo Colorado (Quezaltenango).

Moist or wet, mixed forest, 1,300-2,300 meters; Alta Verapaz
(type collected near Tactic, Standley 70477); El Progreso; Guate-
mala(?); Huehuetenango; Quezaltenango; San Marcos; endemic.

A small to large tree, the branchlets slender, subterete, with numerous elevated
lenticels, densely brown-lepidote when young, glabrate in age, sometimes also
short-hirsute, especially at the nodes; leaves all or mostly opposite, large, petiolate,
the rachis slender, lepidote, sometimes hirtellous; leaflets usually 14-16, narrowly
oblong-lanceolate, 9-20 cm. long or more, 2.5-5 cm. wide, narrowly long-attenuate,
acute at the base and almost symmetric, sessile or very shortly petiolulate, usually
entire, at least in adult leaves, green above, minutely hispidulous along the costa
or almost wholly glabrous, pale and often glaucescent beneath, rather densely
and minutely appressed-lepidote.

This species is known only from sterile material, but numerous
specimens of it are at hand. These specimens exhibit so much varia-
tion that it is quite possible that they represent two distinct species.
When fruiting material of Engelhardtia and the tree here referred to
Alfaroa has been collected in ample quantity, it is possible that some
surprising results may be obtained. E. guatemalensis is abundant
in Alta Verapaz, especially about Tactic, where it was first observed.
The trees may be recognized from a long distance because the young
foliage is crimson. In its coloring as well as in the form of the leaves
and leaflets the tree closely resembles the dreaded "Amcne*" (RhtLS
striata), and the resemblance is so great that the senior author
ventured to collect material only after a close inspection of the
branches. The leaves are opposite in Engelhardtia and alternate,
of course, in the Rhus. In spite of its abundance in this region-
one where every plant has a name no name could be obtained for
this conspicuous tree. Neither has it been possible to find flowering
or fruiting trees in any part of Guatemala where they grow. The
nut in this genus is much like a small hickory nut (Gary a), but is
subtended by a very large, hand-shaped bract.

356 FIELDIANA: BOTANY, VOLUME 24

JUGLANS L.

By WAYNE E. MANNING

Trees, rarely shrubby; branchlets with chambered pith; buds scaly; leaves
alternate, deciduous; leaflets serrate or entire; flowers monoecious; staminate
flowers in sessile, lateral, solitary, though frequently superposed, drooping catkins
on last year's twigs; staminate calyx 1-4 lobed, the two subtending bracteoles
(each sometimes divided) apparently in the same whorl with the calyx so the
perianth appears 6- to 8-lobed in addition to the bract; bract entire; stamens
7-100; pistillate flowers in few- to many-flowered terminal spikes; pistillate
calyx usually 4-lobed; the bract, the two bracteoles (the tips of the latter frequently
split into several teeth), and the calyx adherent to the ovary in apparently two
rings; inner bracteoles present at least in J. nigra; ovary inferior; style branches
(stigmas) carinal, the inner surface of each plumose and stigmatic; fruit a nut
enclosed in an indehiscent (or in J. regia irregularly dehiscent) rather fleshy husk,
the latter composed of the ripened involucre and calyx; nut woody, typically
rough-ridged or sculptured; a pair of secondary wall cavities present in the outer
wall of the nut at each end of the secondary (dorsal) partition or in that location,
and traversing the wall of the nut; tertiary wall cavities, between the secondary
cavities and the primary partition, frequently present, and at certain levels
fusing with the secondary wall cavities; primary wall cavities, at each end of the
primary partition, present in a few species; nut finally separating into two valves
at germination, the large fleshy cotyledons remaining within the shell.

About twenty species in North, Central and South America and
in the Old World from southeastern Europe to eastern Asia. Five
or six species of Juglans occur in Mexico.

Leaflets entire or nearly so, 7-9 (rarely to 13); staminate flowers subsessile, the
floral receptacle elongate; husk of fruit finally splitting irregularly; nut merely
rugose, with a pair of strong longitudinal ridges or lips on each side of the

line of dehiscence J. regia.

Leaflets serrate, 15-25; staminate flowers clearly stalked, the floral receptacle

round; husk not splitting, the nut without lips.

Rachis, fruit, and lower surface of leaflets glabrate; leaflets short-stalked,
tapering at base or sometimes subtruncate above the stalk; fruit strongly

verrucose J. guatemalensis.

Rachis, fruit, and lower surface of leaflets definitely pubescent; leaflets truncate
at base, essentially sessile; fruit not strongly verrucose J. Steyermarkii.

Juglans guatemalensis Manning, sp. nov. NogaL

Mountain sides, river bottoms, along water courses, cafetales,
500-1,500 jneters; endemic; Alta Verapaz; Baja Verapaz; Quiche"
(Finca San Francisco Cotzal, alt. 1,250 meters, December 6, 1934,
Alexander F. Skutch 1866; type in Herbarium of Arnold Arboretum;
isotype in Herbarium of Chicago Natural History Museum), Hue-
huetenango; occasionally planted (Guatemala).

A medium-sized or large tree, sometimes 55 meters tall with a trunk 2 meters
in diameter, the bark dark and deeply furrowed; twig dark brown, appearing

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 357

glabrate, but minutely glandular-puberulent, the lenticels prominent; leaves
alternate, large, chiefly 45-65 cm. long and 30-35 cm. wide, the terminal leaflet
usually present; leaflets large, distant, chiefly 17-21, each 14-17 cm. long and
5-6 cm. wide, decidedly stalked, the stalk 2-4 mm. long; leaflets finely to strongly
serrate, oblong-elliptic, obliquely truncate or rounded at the base above the stalk,
commonly cuneate on the lower side, long-acuminate at apex, the lowest leaflets
frequently small; rachis glabrous or glabrate, but with a strong lens appearing
densely puberulent with minute stalked glands; upper surface glabrate; lower
surface of leaflets glabrate, with only a few stalked glands and a few minute
fascicled hairs on midrib and a few on larger lateral nerves, young leaflets with
whitish pointed hairs on the smallest veins; very young leaflets drying green, with
almost no fascicled hairs beneath; staminate catkins 22-30 cm. long, each bract
short-hairy, small, 1 mm. long, obtuse, essentially on the catkin-rachis at the
base of the flower-stalk throughout the catkin; flower-stalk well developed, 4-5
mm. long; the bracteole-sepal ring frequently with 8 organs; stamens 61-102;
pistillate flower and immature fruit unknown; mature fruit large, subglobose-
pyriform, 4.3-4.5 cm. thick and 4.7-4.8 cm. long to probably 5.5 cm. thick and
5.8 cm. long, essentially glabrous, shining greenish-brown, strongly and conspic-
uously white-punctate-verrucose (warty), with the punctations open and lenticel-
like, not wrinkled in drying, the husk thick; nut reddish-brown, subglobose, slightly
flattened, 3.4-4.5 cm. thick and 3.2-4.2 cm. long, strongly ridged, the ridges broad,
flat-topped, interrupted, rather rounded in cross-section, the furrows comparatively
shallow except at the depressed base of the nut, where the ridges are sharp and the
furrows deep; primary wall cavities well developed usually at all levels, frequently
united with tertiary wall cavities, and at the very base of the nut uniting also with
secondary wall cavities; secondary wall cavity ridges strongly projecting into the
loculus at the one-celled level and at least in one collection at the 2-celled level;
secondary septa rather low.

Folia inaequaliter pinnatim foliata; foliolis magnis 17-21 valde 2-4 mm.
longe petiolatis oblongo-ellipticis basi oblique truncatis 14-17 cm. longis 5-6 cm.
latis, tenuiter vel argute serratis, utrinque glabratis; rhachi glabra vel glabrata
sed minute glandulari-puberulenti; foliolis novellis in si ceo viridibus glabratis;
amends masculis 22-30 cm. longis, bracteis minutis 1 mm. longis; fructibus sub-
globoso-pyriformibus 4.7-5.8 cm. longis 4.3-5.5 cm. diam., punctato-verrucosis
praecipue glabris lucidis; mice subglobosa 3.2-4.2 cm. longa 3.4-4.5 cm. diam.
valde costata, costis latis planis, sulcis tenuibus.

The collections of Popenoe and of Standley are represented only
by nuts, and there is some uncertainty about the identification;
the collection of Steyermark has the internal cavities absent at the
end of the primary partition in the nut. This may be the species
reported by Re'ne' GueYin (as nogal, Juglans nigra), in Catalogue
des products presentes par la Republique de Guatemala a I' exposition
universelle de Paris, 1900, from Coban, Alta Verapaz; Cuilapa,
Santa Rosa; Chimaltenango, Zacapa, and Chiquimula. He reports
the wood of the tree as used for manufacturing fancy furniture; its
fruits are edible and the husk of the fruit is used for tanning
materials.

358 FIELDIANA: BOTANY, VOLUME 24

Juglans guatemalensis has been confused with J. pyriformis
Liebmann, collected by him near Coscomatepec, Vera Cruz, Mexico;
that species has about 24 narrow sessile leaflets which are truncate
at base, a glandular pubescent rachis, a rougher husk on the fruit
and a slightly more roughly ridged nut. J. guatemalensis resembles
most closely the walnuts of central and western Mexico; the cross
section of the nut of the type resembles closely that of J. peruviana
described from "Peru" by Dode. This species may be the one
cultivated in El Salvador; a similar collection has been made in
Honduras.

Juglans regia L. Sp. PI. 1415. 1753. Palo de nuez; nogal.
Huehuetenango (planted) and probably elsewhere cultivated.

Broad-headed tree to 100 feet, though in cultivation usually only 40-60 feet
tall, with light gray bark, remaining smooth a long time; twigs glabrous; leaflets
5-9, rarely to 13, mostly obovate, entire, though on young plants frequently
somewhat serrate, essentially glabrous; rachis glabrous; staminate catkins 5-10 cm.
long; staminate flowers with elongate receptacles; fruit subglobose, glabrous, green,
4-5 cm. in diameter; nut ovoid to globose, thick to thin-shelled, wrinkled or
shallowly pitted externally, with two longitudinal ribs (lips) on each side of line
of dehiscence; nut splitting early; septa thin.

Persian or English Walnut, native of southeastern Europe to
Himalaya Mountains, the most frequently planted commercial
walnut, especially in warmer areas.

The one specimen collected does not seem to be typical J. regia
and may represent a hybrid between this species and a native one.
The specimen is sterile.

Juglans Steyermarkii Manning, sp. nov. Nogal.

Canons, between 1,200 and 1,300 meters; endemic; Huehue-
tenango (Paso del Boqueron, Rio Trapichillo, below La Libertad,
alt. 1,200-1,300 meters, type collected August 21, 1942, Steyer-
mark 51140, in Herbarium of Chicago Natural History Museum;
isotype in W. E. Manning Herbarium).

Tree about 17 meters tall, the bark gray but furrowed; twigs densely glandular
pubescent; terminal bud elongate, densely grayish-tomentose; leaf scar only
slightly notched; leaves alternate, large, up to 60 cm. long and 30 cm. wide, the
terminal leaflet usually absent; leaflets large, distant, mostly 16-18, each 14-17 cm.
long by 5-8 cm. wide, essentially sessile or very short-stalked, ovate or ovate-
lanceolate, typically finely but rather remotely serrate with the serrations short
and turned outward, and with most of the small veins going directly to the serra-
tion instead of to the sinus; base of leaflet usually truncate to broadly rounded,
the apex acute or abruptly acuminate, the lowest leaflet somewhat reduced;

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 359

rachis conspicuously pubescent with reddish glandular hairs; upper surface of
leaflets pubescent with minute glandular hairs, the lower surface pubescent with
reddish glandular hairs on larger and smaller veins and especially on the midrib
and with solitary or paired whitish pointed hairs on the surface and smaller
veins; flowers unknown; nearly mature fruit wrinkled, subglobose-pyriform,
truncate at base, 4 cm. diameter and 4.5 cm. high, strongly pubescent with reddish
glandular hairs; punctations numerous, drying brown, very inconspicuous; husk
not very thick; nut apparently deeply grooved, with rounded ridges, but nut
immature; cavities present in the wall at each end of the primary partitions.

Arbor ca. 17-metralis; foliis equaliter pinnatim foliatis; foliolis magnis 16-18
ovatis vel ovato-lanceolatis praecipue sessilibus obtusis vel abrupte acuminatis
basi truncatis 14-17 cm. longis 5-8 cm. latis, tenuiter remoteque serratis, subtus
pilis prominentibus rubescentibus glandularibus et albidis solitariis vel binis
pubescentibus; rhachi dense rubello-glandulari pubescenti; fructibus subgloboso-
pyriformibus, 4 cm. diam., 4.5 cm. longis, dense glandulari-pubescentibus, parce
brunneo-punctatis, in sicco corrugatis; nuce subglobosa costata, costis rotundatis
tenuibus.

Nuts edible and of value where sufficiently common, but the
meat small; wood probably useful, as in all of the black walnut
group.

BETULACEAE. Birch Family
Reference: Hubert Winkler, Pflanzenreich IV. 61: 1-149. 1904.

Trees or shrubs, the buds with few or numerous scales; leaves alternate, penni-
nerved, mostly serrate or dentate, petiolate; stipules free, commonly deciduous;
flowers monoecious, the staminate in aments, the pistillate also in aments or some-
times capitate, terminal or lateral, usually appearing when the trees are leafless;
perianth simple, membranaceous, the segments free or united, sometimes absent;
stamens 2-12, the filaments free; ovary hypogynous, 2-celled, usually laterally
compressed; styles 2, free, filiform; ovules 2, pendulous, anatropous; fruit a nut,
usually laterally compressed, often winged, sometimes crowned by the remains
of the perianth, often subtended or inclosed in the persistent bracts and bractlets;
seed by abortion usually solitary, without endosperm, the testa membranaceous;
cotyledons fleshy, smooth, rugose, or ruminate.

Six genera are recognized, chiefly in temperate regions of both
hemispheres, some of the genera extending to mountain regions of
the tropics. No other genera are represented in Central America.
The monograph of Winkler cited above is unsatisfactory so far as
tropical American members of the family are concerned.

Pistillate aments cone-like in fruit, with woody bracts; staminate flowers 2-3 in
each bract; no perianth in the pistillate flower Alnus.

Pistillate aments not cone-like, the bracts thin; staminate flowers one in each
bract; perianth present in the pistillate flower.

Fruit enclosed in a bladder-like closed sac Ostrya.

Fruit merely subtended by a flat leaf-like 3-lobate bract Carpinus.

360 FIELDIANA: BOTANY, VOLUME 24

Corylus Avellana L. The European hazel or filbert (Spanish
"avellana") is well known for its edible nuts, which are imported
into Guatemala in small quantities. A few plants of this species
were seen growing in a plantation near Coban. They were said to
have been there for some years but consisted of soft shoots only
a few inches high and were said never to grow any taller.

ALNUSHill. Alder

References: M. L. Fernald, Proc. Amer. Acad. 40: 24-28. 1904;
H. H. Bartlett, Proc. Amer. Acad. 44: 609-612. 1909.

Trees or shrubs; leaves alternate, petiolate, serrate or dentate, penninerved,
deciduous; stipules caducous; flowers dioecious; staminate spikes from terminal
buds formed the preceding year; pistillate spikes solitary or racemose, arising in
the axils of the leaves of the previous year or in the axils of the new leaves; stami-
nate flowers in 3-flowered clusters, these arranged in cylindric, scaly-bracteate
spikes; perianth segments 4 or fewer, connate at the base or free; stamens 4,
opposite the perianth segments; anthers ovate, the cells distinct, parallel; pistillate
flowers in erect, cylindric or oblong, scaly spikes, without a perianth; ovary 2-celled,
the styles short, stigmatose at the apex; ovule one in each cell, pendulous, anatro-
pous; fruit a nutlet, compressed, usually winged, by abortion 1-celled.

About 25 species, in America, central Asia, Algeria, and Europe.
No others are known from Central America.

Leaves densely tomentose beneath over almost the whole surface, rarely glabrate

in age. Leaves acute or acuminate, not glandular beneath A. ferruginea.

Leaves glabrate beneath, at least in age, the long hairs, if any, confined to the
nerves.

Leaf blades all or mostly rounded or very obtuse at the apex A. firmifolia.

Leaf blades all acute or acuminate.

Leaves with rather dense, yellowish wax glands on the lower surface.

A. jorullensis.
Leaves nearly or quite without wax glands on the lower surface. . .A. arguta.

Alnus arguta (Schlecht.) Spach, Ann. Sci. Nat. II. 15: 205.
1841. Betula arguta Schlecht. Linnaea 7: 139. 1832. A. ovalifolia
Bartlett, Proc. Amer. Acad. 44: 611. 1909 (type. from San Lucas,
Sacatepe"quez, J. D. Smith 2188). Aliso; Ilamo;Lemop (Guatemala).

Common in the mountains, often forming almost pure, dense,
extensive stands, but more often associated with oaks and often
with pines, sometimes in Juniperus forest, 1,350-3,000 meters; Alta
Verapaz; El Progreso; Guatemala; Sacatepe"quez; Chimaltenango;
Solola; Totonicapan; Quiche"; Huehuetenango; Quezaltenango; San
Marcos. Southern Mexico; El Salvador.

A tree, sometimes 30 meters high but ordinarily less than half as large, the
bark pale, thin, smooth, the branchlets glabrous or nearly so; leaves slender-petio-

FIG. 50. Alnus arguta. A. Habit of portion of fruiting and flowering branch;
X H- B. Fruiting strobile; X 1. C. Staminate flower with bract; X 8. D.
Staminate flower; X 7. E. Pistillate flower with bract; X 8. F. Nutlet; X 8.
G. Scale of a mature strobile with nutlets in position; X 8.

361

362 FIELDIANA: BOTANY, VOLUME 24

late, oblong-ovate to broadly ovate, 3-9 cm. wide, mostly acute or acuminate,
obtuse or rounded at the base, duplicate-serrate, often shallowly lobate, glabrous
above or nearly so, beneath usually pilose or villous along the nerves, in age usually
glabrate, pale or often ferruginous; staminate aments 4-10 cm. long; pistillate
strobiles sessile or pedunculate, mostly 2-3 cm. long, sometimes somewhat
shorter. (Fig. 50.)

The term "aliso" is the Spanish name for the alder. In geo-
graphic names it appears locally in Alisos and Los Alisos, caserios
of San Marcos and Huehuetenango, but the name would be appro-
priate for almost any settlement of the western mountains. Alders
are most abundant in the Occidente, and on the inland mountain
slopes of San Marcos they form extensive and often pure stands,
especially on the white sand slopes typical of that region. The
stands are either dense or sparse, and in early morning, seeing the
expanses of pure white sand between the trees, one has the impression
that snow has fallen during the night, a not unreasonable phenomenon
considering the frosty temperatures that commonly prevail. The
flora of these Alnus forests is meager and uninteresting. Toward
the end of the dry season the trees shed all or most of their leaves,
and soon afterward they become covered with pale, bright green,
young foliage, a color of vegetation most unusual in Central America,
where the prevailing greens are dark and somber. Alders are con-
fined mostly to the drier mountains, and are scarce or quite absent
on the slopes facing the Pacific, where there is abundant moisture
throughout the year. A few trees do grow below Santa Maria de
Jesus, Quezaltenango, but they do not seem to thrive. Probably
the wind carries the winged nutlets down from the abundant stands
on the higher slopes of the Volcan de Santa Maria. On the white
sand slopes of San Marcos, alders are perhaps the first trees to estab-
lish themselves on abandoned land, or that on which volcanic ash
has newly fallen. The trees here bloom in January and February.
The bark turns red when cut. It is used to obtain a brown dye
much used for cotton and other textiles. The wood seems to be
little used except for firewood. Many of the trees on the Volcan
de Santa Maria were bearing "witches' brooms," probably the result
of insect attacks.

Alnus ferruginea HBK. Nov. Gen. & Sp. 2: 21. 1817. A.
acuminata var. ferruginea Regel in DC. Prodr. 16. pt. 2: 184. 1864.
Lenop (Guatemala); A. guatemalensis Gandoger, Bull. Soc. Bot.
France 66: 289. 1920 (type from Coban, Turckheim probably no.
11.1013). Hamam (Coban, Quecchi).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 363

Along streams or in wet thickets, often in pine forest, sometimes
forming extensive, almost pure stands on mountain slopes, 1,250-
2,400 meters; Alta Verapaz; Guatemala; Chimaltenango; Quiche";
Huehuetenango; Quezaltenango; San Marcos. Probably in Chiapas;
Andes of South America.

A small or medium-sized tree, sometimes very large, with thin, dark brown,
smooth bark; leaves petiolate, elliptic or broadly ovate, 7-15 cm. long, acute or
acuminate, obtuse or rounded at the base, duplicate-serrate, green above and
glabrate or puberulent; staminate aments 5-15 cm. long; pistillate strobiles short-
pedunculate or subsessile, 1.5-3 cm. long or sometimes slightly larger, 10-14 mm.
thick.

This plant is easy of recognition by the abundant pubescence of
the leaves, but may prove to be no more than a pubescent variety
of the widely distributed A. jorullensis. It is particularly plentiful
in the mountains of Huehuetenango, where it may be found almost
anywhere along streams or on hillsides. It grows in abundance
also in the region of Coban, where it is mostly confined to stream
banks, often with Carpinus and Acer.

Alnus firmifolia Fernald, Proc. Amer. Acad. 43: 61. 1907.
Aliso; Alls; L'm'ump (Palojunoj, Quezaltenango); Ilamo; Analmat
(Volcan de Agua).

Abundant in mixed or coniferous forest on the higher mountains,
mostly at 2,500-3,600 meters, rarely at lower elevations (in Huehue-
tenango as low as 1,900 meters); Guatemala (Volcan de Pacaya);
Sacatepe'quez (Volcan de Agua); Huehuetenango; Totonicapan;
Quezaltenango; San Marcos. Southern Mexico.

A shrub or a tree, sometimes 18 meters high with a trunk almost a meter in
diameter, often flowering when only a low shrub, especially on the highest moun-
tain slopes, the bark thick and corky, silvery gray, smooth, the branchlets glabrous
or often sparsely villous; leaves petiolate, usually thick and subcoriaceous, variable
in form, elliptic-oblong to elliptic or oval, mostly 5-10 cm. long, generally obtuse
to rounded at the apex but some of the leaves frequently acute or rarely acuminate,
obtuse or acute at the base, glabrous and lustrous above, beneath often glaucescent
and almost always pale, rarely brownish, at first villous-pilose, at least on the
nerves, in age often almost glabrous, the lateral nerves very prominent and con-
spicuous; staminate aments 3-4 cm. long; pistillate strobiles oval-cylindric, mostly
10-17 mm. long, about 8 mm. thick.

This is one of the most abundant and characteristic trees of the
high mountains of central and western Guatemala, where it is
practically confined to the uppermost slopes, above those frequented
by A. arguta and A. ferruginea. There is usually an abrupt transition
from one species to the other, as one ascends the trails, the change

364 FIELDIANA: BOTANY, VOLUME 24

in species being easy to recognize because of the difference in color
of the trunks. The bark of A. firmifolia is unique, suggesting some-
what that of the corky-barked firs (Abies) of the Rocky Mountains.
The bark has transverse constrictions, as if it were bound tightly
with twine. On the Volcan de Santa Maria the Indians made no
distinction as to the two species so abundantly represented there,
yet when the bark difference was pointed out, they recognized it
immediately. This is the only Alnus species, apparently, of the
higher parts of the Cuchumatanes. Skutch remarks that this and
Pinus Montezumae var. rudis are the only common trees in the
Cuchumatanes above 3,150 meters, but he overlooked Juniperus
Standleyi, which surely is a tree, and abundant. The leaves of A.
firmifolia, at least on some specimens, are suggestive of those
of Crataegus. They usually are devoid of wax glands on the lower
surface, but sometimes bear widely scattered, small glands that
turn blackish with age. The wood of Alnus is rich in tannin. In
Mexico, and quite probably in Guatemala, it is used for tanning
leather, to which it imparts a red color. The junior author remembers
numerous occasions when his native guides selected alder trees for
marking the new trails through the mountain forests. The red
marks of the bark cut by their machetes were as effective as red
paint. The wood is rather light and soft but firm, straight-grained,
rather fine-textured, easy to work, tough and strong, finishes
smoothly, is not durable in contact with the ground.

Alnus jorullensis HBK. Nov. Gen. & Sp. 2: 27. 1817. Aliso.

Moist or wet mixed forest, often associated with oaks, 1,800-3,700
meters; Sacatepe"quez; Chimaltenango; Solola; Quezaltenango; San
Marcos; Huehuetenango. Mexico, the type from Volcan de Jorullo;
Andes of South America, from Venezuela to Bolivia.

A small or sometimes large tree with thin brown smooth bark, the branchlets
usually glabrous; leaves oblong to ovate or oblong-obovate, mostly 7-12 cm. long,
acute or acuminate, or a few of the leaves obtuse, acute to rounded at the base,
irregularly serrate, green and glabrous above, usually lustrous, pale beneath,
more or less pilose at first but in age glabrate, dotted beneath with yellow wax
glands, these usually conspicuous and often very dense; staminate aments 3-6 cm.
long; strobiles sessile or subsessile, generally about 12 mm. long, sometimes
slightly larger.

This species seems to be of only sporadic occurrence, and only
a few specimens of it have been collected, in contrast with very
numerous collections of the other local species.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 365

CARPINUS L.

Large or medium-sized trees; leaves petiolate, serrate, the stipules caducous;
staminate flowers in cylindric pendulous aments, bracteate; perianth none; stamens
4-12, inserted on the pilose torus, the filaments bifid; anthers inserted dorsally
below the middle, 1-celled, pilose at the apex; pistillate flowers in a large-bracted
spike or ament, the flowers solitary within the bract; styles short, the 2 stigmas
erect, linear-subulate; nut ovoid, subcompressed, by abortion 1-celled, the pericarp
somewhat ligneous; seed solitary, pendulous.

About twenty species, in the northern hemisphere. A single
species is found in America.

Carpinus caroliniana Walt. Fl. Carol. 236. 1788.

Ranging from southeastern Canada to central Mexico. Repre-
sented in Central America by the following variety:

Carpinus caroliniana var. tropicalis Bonn. Smith, Bot. Gaz.
15: 28. 1890. C. tropicalis Lundell, Lloydia 2: 79. 1939. Duraznillo
(fide Aguilar).

Usually along streams, in moist or wet forest, 1,300-2,800 meters;
Alta Verapaz (type from Chicoyonits, J. D. Smith 1667); Baja
Verapaz; Zacapa (Sierra de las Minas); Jalapa; Quiche"; Quezal-
tenango (south slope of Volcan de Santa Maria); San Marcos;
Huehuetenango. Chiapas; Honduras.

A large or medium-sized tree, often 12 meters high or more, with smooth dark
bark, the young branchlets densely pilose with ascending or subappressed hairs;
leaves membranaceous, on petioles 1 cm. long or less, ovate to ovate-oblong, mostly
5-7 cm. long, acute or acuminate, rounded or subcordate at the base, appressed-
pilose on the nerves, slightly paler beneath, with 11-15 pairs of conspicuous straight
lateral nerves, unequally duplicate-serrate; staminate aments 1.5-2.5 cm. long, 5
mm. thick, the bracts broadly ovate, acute; stamens 6-10; pistillate inflorescences
7 cm. long or less, the bracts foliaceous, 5-7-costate, trilobate, 2-3 cm. long, the
lobes unequal, dentate or subentire; nut ovoid, nerved, pubescent. (Fig. 51.)

Although raised to specific rank by Lundell, this differs but little
from the blue beech as it is found in eastern United States, except
in its more abundant pubescence. The trunk is more or less fluted,
although the fluting seems not to be so conspicuous in Guatemalan
trees as in those of eastern United States. So far as known, the wood
is not used in Guatemala. In Mexico it is reported to be utilized
for firewood and charcoal.

OSTRYA Scopoli. Hop hornbeam

Trees, pubescent, the leaves short-petiolate, membranaceous, penninerved,
plicate along the nerves in vernation; stipules caducous; flowers monoecious,

FIG. 51. Carpinus caroliniana var. tropicalis. A. Habit of portion of flower-
ing branch; X 2 A- B- Habit of portion of fruiting branch; X 2 A- C. Staminate
flower with scale; X 6. D. Stamen, X 24. E. Pistillate flower with scale; X 6.
F. Pistillate flower with bract ;X 5. G. Nut with involucre; X 1. H. Fruit; X 8.

366

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 367

the staminate in cylindric aments with scale-like bracts; perianth none; stamens
3-14, inserted on the pilose torus, the filaments slender, bifid at the apex; anthers
dorsifixed near the base, pilose at the apex; pistillate flowers disposed in dense
spikes, the bracts utricle-like, open at the apex in anthesis, closed in age and accres-
cent, membranaceous and inflated; ovary 2-celled, the style short, the 2 stigmas
erect, linear-subulate; ovules 2 in each cell, pendulous, anatropous; nutlet small,
by abortion 1-celled, costate, the pericarp subligneous; seed solitary, pendulous,
the cotyledons fleshy, plano-convex, obovate.

Probably three species, in Asia and North America, only one
species reaching Central America.

Ostrya virginiana Mill. Diet. ed. 8. 1768.

Widely distributed in eastern North America from southeastern
Canada to Florida, and southward into Mexico. Represented in
Central America by the following variety:

Ostrya virginiana var. guatemalensis (Winkl.) Macbride,
Field Mus. Bot. 4: 193. 1929. 0. italica subsp. virginiana var.
guatemalensis Winkl. Pflanzenreich IV. 61: 22. 1904. 0. guate-
malensis Rose, Contr. U. S. Nat. Herb. 8: 292. 1905. Duraznillo
(the name most generally used in Guatemala); Aliso bianco; Aliso
Colorado (Huehuetenango) ; Mescal; Tatiscoba (Volcan de Agua);
Gamuso (Zacapa).

Dry or moist, mountain forests, 1,000-3,000 meters; Baja Vera-
paz; Zacapa; Chiquimula; Jalapa; Santa Rosa; Guatemala; Saca-
tepe*quez; Chimaltenango; Solola; Suchitepequez; Quiche"; Huehue-
tenango; Quezaltenango; San Marcos. Southern Mexico; Honduras.

A small to rather large tree, mostly 6-18 meters tall, with thin, light brown
bark; leaves thin, short-petiolate, ovate to lance-oblong, mostly 6-10 cm. long,
acute to long-acuminate, rounded or subcordate at the base, duplicate-serrate,
usually densely soft-pilose beneath and often also on the upper surface; staminate
aments slender and elongate, pendent; fruiting aments resembling strobiles of
hops (Humulus Lupulus), 5-7 cm. long, the soft enlarged bracts pale green or
finally brownish, conspicuously nerved, inflated. (Fig. 52.)

Called "guapaque" in southern Mexico, and "canillo de venado"
in Honduras. The wood is very hard, fine-grained, tough, and fairly
durable. In some regions it is utilized for railroad ties, tool handles,
and fuel. The bark is said to be used in Mexico for tanning and
dyeing. The tree is abundant in many parts of the central mountains,
where it sometimes forms rather extensive and almost pure stands,
as in Guatemala and Sacatepe"quez. The foliage is almost exactly
like that of Carpinus, but the trunks and bark are somewhat different,

FIG. 52. Ostrya virginiana var. guatemalensis. A. Habit of portion of flower-
ing branch; X H- B. Habit of portion of fruiting branch; X H- C. Staminate
flower with scale; X 5. D. Stamen; X 20. E. Pistillate flower with scale; X 3.
F. Pistillate flower with bract laid open; X 5. G. Fruit and involucre; X 1.

368

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 369

and the two genera apparently do not grow together in Guatemala.
In fruit it is easy to separate the two trees.

FAG ACE AE. Beech Family

Trees or sometimes shrubs, the buds with imbricate scales; leaves alternate,
usually petiolate, persistent or deciduous, penninerved, entire or dentate to
deeply pinnatifid; stipules present, generally deciduous; flowers monoecious,
usually axillary on young branchlets, the perianth 4-7-lobate; staminate spikes
(catkins) slender, usually elongate, pendent, and flexuous, each bract enclosing
a single flower; stamens as many or twice as many as the perianth lobes, rarely
more numerous, the filaments usually slender; anthers 2-celled, the cells erect,
parallel, sessile or short-stipitate at the apex of the filaments, dehiscent by a longi-
tudinal slit; pistillate flowers solitary or in 3's, forming spikes or short clusters,
sometimes borne at the base of the pistillate spikes; ovary inferior, 3-celled or rarely
6-celled at the base, with 3 styles, each cell 2-ovulate, but all the ovules except
one abortive; fruit a nut, solitary or 2-3, partly or wholly enclosed by an in-
volucre or cupule; seed large, without endosperm, the cotyledons thick and carnose.

Six genera, with about 600 species (or perhaps more), widely
dispersed in both hemispheres, mostly in temperate and subtropical
regions, rarely in truly tropical areas, in the tropics mostly confined
to the mountains. Only one genus is native in Central America.
To this family belong the chestnut (Castanea; "Castano;" "castana"),
whose edible nuts doubtless reach Guatemala at times. The
European chestnut, Castanea sativa Mill., is in cultivation and fruit-
ing in Costa Rica, and may well be planted in Guatemala, although
we have no record of its occurrence there. A few species of the same
genus are native in the United States, one of them, C. dentata
(Marsh.) Borkh., formerly of great importance as a source of lumber
and nuts, but now very much depleted because of the attacks of
a fungus disease. Also belonging to this family is the beech (Fagus;
"haya"), one species of which is native in the eastern half of temper-
ate North America. Another species, Fagus mexicana Martinez, has
been described recently from Mexico.

QUERCUSL. Oak

References: William Trelease, The American oaks, Mem. Nat.
Acad. Sci. 20: 1-255. pis. 1-420. 1925. Cornelius H. Muller, The
Central American species of Quercus, U. S. Dept., Agr. Misc. Publ.
no. 477: 1-92. pis. 1-124. 1942.

Large or medium-sized trees or sometimes shrubs; buds crowded toward the
ends of the usually fluted twigs; leaves alternate, almost always distinctly petiolate,
entire, dentate, or pinnatifid, persistent or deciduous, the teeth or lobes often aris-

370 FIELDIANA: BOTANY, VOLUME 24

tate-tipped; stipules associated with the buds rather than the leaves, subulate
to ligulate, usually caducous; staminate flowers small, in elongate flexuous pendent
catkins, apetalous; calyx of 5 lobes united to form a more or less cupular perianth,
this enclosing 5-10 free stamens with short anthers and slender filaments; pistillate
flowers borne in a reduced catkin, the stiff woody rachis either short or long and
1-several-flowered, the calyx of 6 sepals adherent to the bases of the styles and
fused into a tube; pistil of 3 carpels forming a single 3-celled ovary and 3 free
styles, these stigmatic ventrally toward the dilated apex; fruit an acorn (Spanish,
bellota), 1-celled and 1-seeded, the 5 other ovules abortive and adherent to
the developed seed; seed enclosed in a shell and seated in a cup, this formed of
numerous small imbricate scales, the cup enveloping the whole nut or covering
only a part of it, or only its base.

The only large genus of the family, the number of species very
uncertain because of different treatments by various authors. Rehder
estimates the total number of species at "more than 200," while
Trelease recognized about 370 from America alone. Muller recog-
nized 46 as occurring in Central America. The majority of these
are found in Guatemala, but numerous other species grow south-
ward in Costa Rica and Panama. The oaks of Guatemala and
Costa Rica have been collected rather thoroughly, but those of the
intervening countries are little known.

The present account of Guatemalan oaks is taken directly from
the excellent and recent account by Muller, who has determined the
greater part of our collections. We have made rather extensive
collections in Guatemala, and as these have been taken from what
were assumed to be average and characteristic trees in each region
where we have worked, they should be representative of the oak
flora as a whole. Most of the species thus far found in the country
are represented by a substantial number of collections. Many
of our specimens are sterile, probably because they have been
taken chiefly during the dry season. At this time of the year it is
difficult to find acorns, even old ones on the ground beneath the
trees.

As a source of wood and lumber the genus Quercus is one of the
most important of all groups of trees. The timber is noted for its
strength, durability, and beauty, and is used everywhere for in-
numerable purposes, ranging from fuel to railroad ties, construction
of buildings and ships, interior trim, flooring, and all grades of
furniture. The woods of different species vary as to their physical
qualities; some of them are very hard and tough, others are lighter
in weight, softer, and less tough. Cork of commerce is obtained
from the bark of the cork oaks grown for this product in Spain,
Portugal, and North Africa.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 371

In Guatemala oaks and pines are the two most characteristic
and important trees. They often are mixed in pine-oak forests,
but frequently the oaks form almost pure stands, which formerly
must have covered almost unbrokenly the middle elevations of the
drier mountains. Even at the present time, when the oak forests
have so long been a source of fuel and lumber, there are extensive
stands covering great areas of such departments as Guatemala,
Quiche", and Huehuetenango. These regions afford the best grazing
areas of the mountains. It is an impressive sight to stand on some
low elevation near Huehuetenango and observe the oak forests
extending in every direction. Most of the trees are rather small,
it is true, and often reduced to thicket-forming shrubs, but on all
sides, as far as the eye can see, there are oak forests or thickets,
extending from the plains up to the highest summits of the great
wall of rock that constitutes the westward face of the Cuchumatanes.
Few or none of the Guatemalan oaks assume the brilliant colors
peculiar to many North American oaks in autumn, but there is a
great deal of bright yellow and some dull reds and purples, and the
brown coloring of the dead but persistent leaves is characteristic.

Oaks often are found where there are no pine trees, and they
frequently occur in mixed forest associations. About Coban the
customary place of the oaks is usurped in places by Liquidambar
trees, but not far from Coban there are extensive stands of oak
forest, and the dry mountains of Baja Verapaz probably have the
greatest display of large oak trees to be found anywhere in Guate-
mala. In the dense mixed wet forests of the Pacific slopes of the
main chain of volcanoes, oaks occur in great numbers, principally
Quercus Skinneri. The drier mountains of Quezaltenango and San
Marcos, at middle or rather high altitudes, also afford a great display
of oak forest, but this, as about the capital, has been greatly reduced
by cultivation and the demands for fuel and lumber.

Oak wood is the favorite fuel everywhere in Guatemala. The
wood is carried great distances on men's backs or on animals.
It always is sold so cheaply that the woodcutters can earn at best
a mere pittance, but firewood is one of the necessities all over Guate-
mala, and its scarcity imposes great hardship. Oak bark is rich in
tannin, and supplies the best of all materials for tanning tough
and durable leather. It is much used for this purpose in Guatemala,
and it is used there also to furnish a brown dye for textiles. The
insect galls found so abundantly and often conspicuously on the
trees are particularly rich in tannin, and in many countries have been

372 FIELDIANA: BOTANY, VOLUME 24

much used for making ink. They doubtless are or have been thus
utilized in Guatemala. Because of the tannin present, a decoction
of oak leaves or bark often is used as an astringent in domestic
medicine, commonly as a mouth wash for alleviating toothache.
Ashes of the wood are used commonly for lye in soap-making.

On the Cerro Quemado and elsewhere about Quezaltenango
the abundant fallen oak leaves are used commonly for fertilizing the
sterile fields of the dry mountain slopes. Large heaps of them
are seen rotting about the borders of fields, and in February they are
worked into the soil.

The seeds of all the oaks are edible, but in most species they are
very bitter. Those of some of the white oaks are relatively sweet
and of agreeable flavor. Acorns were an important food among some
of the North American Indians. We have no information as to such
a use anywhere in Central America, but there can be no doubt that
acorns have been eaten in Guatemala, at least in times of famine,
such as have occurred in very recent years, when the corn crop
fails.

The Pokonchi names for oaks are reported as "pi tan" and
"zinuh," and the name "tuhs" is reported from Sierra de las Minas.
Among Spanish-speaking people of Guatemala the names "encino"
and "roble" are applied to the trees, and no distinction is made in
the use of these names, nor are the different species, so far as we have
observed, given distinctive names. The term "encinaladas" is the
one most often applied to oak forests, at least in the central region.

All the Guatemalan oaks are evergreen, as that term is used in
the United States, or essentially so. The term is, however, a mis-
leading one. In the wetter forests the trees probably are covered
with green leaves at all seasons of the year, the change in foliage
taking place gradually. In nearly all the species, however, the
new leaves appear in the spring months of the North, and remain
on the tree until the following spring, or at least well into the verano.
They are dormant but may retain their green coloring, or change
gradually to brown. At any rate, when the new buds begin to swell
after the first showers, or with the increase in temperature, the old
leaves fall rapidly, and the trees often remain for some time quite
bare except for catkins. At this time the oak forests in the barrancos
of the central mountains have exactly the aspect of northern forests
in springtime. They are full of small twittering birds, and many
of the shrubs and herbs are in fresh bloom. We have not seen a
similar appearance of spring vegetation anywhere else in all Central

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 373

America, although it may be duplicated in the Alnus forests of the
white-sand mountains of the Occidente of Guatemala.

Bark rather soft, gray, and scaly; leaves, if dentate, only mucronate-tipped or
rounded, never spinose-tipped or aristate-tipped; stigmas abruptly dilated
on short styles; fruit annual; scales of the cup usually prominently thickened
at the base and loosely appressed at the apex; shell of the acorn glabrous on
the inner surface; abortive ovules basal. Subgenus Lepidobalanus. White
oaks.

Twigs of the season persistently and densely fulvous-tomentose.

Leaves oblanceolate, tapering to the cuneate or narrowly rounded base,
coarsely dentate above the middle; petioles 3-5 mm. long. . . .Q. oocarpa.

Leaves obovate to elliptic-oblanceolate, not tapering below, the base rather
broad, cuneate or truncate, subentire or with low teeth; petioles mostly
more than 10 mm. long Q. insignis.

Twigs of the season glabrate or sparsely pubescent or gray-tomentose.
Leaves with a persistent continuous short tomentum beneath, or the lower
surface canescent and covered with minute appressed stellate hairs.

Leaf blades cordate at the base, the nerves 11-15 on each side, the lower
surface covered with a cream-colored short spreading tomentum.

Q. purulhana.
Leaf blades cuneate to subobtuse at the base, with 6-8 pairs of nerves,

appressed-tomentulose beneath Q. oleoides.

Leaves glabrate or pubescent beneath, not with a short dense tomentum or

with appressed stellate hairs.
Leaves glabrate throughout.

Petioles 1-4 mm. long Q. pilaria.

Petioles 15-40 mm. long Q. corrugata.

Leaves villous or tomentose beneath, or at least with the costa strigose or
stellate-tomentose.

Fruit large, the acorns 2.5-3 cm. broad Q. oocarpa.

Fruit small, the acorns usually less than 2 cm. broad at maturity.
Leaves very thick and coriaceous, with conspicuously impressed veins
above, evidently reticulate-veined beneath and sparsely tomentose,
conspicuously cordate at the base Q. pilicaulis.

Leaves relatively thin, the veins not conspicuously impressed above
nor evidently reticulate beneath, or, if the leaves are thick and
impressed-veined above, then persistently and densely long-
tomentose beneath, or not cordate at the base.

Leaves with a waxy-glaucous bloom beneath.

Petioles usually 15-25 mm. long; leaves entire or with a few teeth

near the apex Q. polymorpha.

Petioles 7 mm. long or shorter, dentate throughout . Q. peduncularis.

Leaves without a waxy-glaucous bloom beneath, or this obscured
by the long lax tomentum.

Leaves thick, impressed-veined on the upper surface, usually laxly
long-tomentose beneath Q. peduncularis.

Leaves if thick not conspicuously impressed-veined above, glabrate
beneath or merely villous or sparsely tomentose, the pubes-
cence not obscuring the surface.

Leaves regularly and usually repandly 10-12-dentate on each
side, or merely erose-dentate.

Leaves rather thick and coriaceous, erose-crenate, drying
yellowish brown Q. segoviensis.

374 FIELDIANA: BOTANY, VOLUME 24

Leaves thin, usually with regular, rounded or acute teeth,

drying green Q. peduncularis var. sublanosa.

Leaves entire or irregularly few-dentate above the middle.
Leaves thick and leathery, glabrate except for the veins, the
margins evidently revolute, entire or finely dentate.

Q. pacayana.
Leaves rather thin, glabrate except for the strigose costa

and nerves, subentire Q. aaata.

Bark rather hard, black, and furrowed but scarcely scaly; leaves, if toothed,
aristate-tipped, never with rounded lobes; stigmas gradually (or rarely
abruptly) dilated, on long styles; fruit biennial or annual; scales of the cup
scarcely thickened at the base and usually tightly appressed at the apex;
shell of the acorn tomentose on the inner surface; abortive ovules usually
apical. Subgenus Erythrdbalanus. Black oaks.
Leaves entire, or the margins sometimes crispate but not at all dentate or

aristate.
Leaves rounded or obtuse at the apex, granular-bullate beneath, at first

tomentose but becoming glabrate in age.
Leaves granular-bullate beneath, at first tomentose, becoming more or

less glabrate Q. tristis.

Leaves smooth beneath, not at all bullate, glabrous or at most with tufts

of hairs beneath in the axils of the nerves Q. sapotaefolia.

Leaves acute at the apex and aristate, or acuminate.
Leaf blades low-bullate or granular-bullate beneath.

Leaves glabrate or somewhat pubescent beneath but not persistently

and laxly tomentose Q. tristis.

Leaves covered beneath with a persistent lax tomentum.

Twigs covered with a persistent velvety tomentum; leaves waxy-
glaucous as well as bullate when denuded of pubescence.

Q. brachy stocky s.

Twigs glabrate or furfuraceous but not velvety; leaves not waxy-
glaucous beneath.
Leaves rather sparsely short-tomentose or merely crisped-villpus

beneath, at most low-bullate Q. crispipilis.

Leaves rather densely thick-tomentose beneath or finally somewhat
glabrate, never sparsely tomentose or short-tomentose, granular-
bullate Q. crispipilis var. pannosifolia.

Leaf blades smooth beneath, not bullate.

Leaves linear-lanceolate, at least 5 times as long as wide, very long-
tapering at the apex, the nerves 18-24 pairs, divergent at an angle

of 80-85 degrees from the costa Q. flagellifera.

Leaves not linear-lanceolate, less than 5 times as long as wide, the
nerves mostly 15 or fewer pairs, usually divergent at a narrower angle.
Fruit biennial.

Leaves acuminate or else narrowly lanceolate.

Petioles 5 or rarely 10 mm. long, the blades entire; acorns 25-30

mm. long and 22-26 mm. broad Q. crispifolia.

Petioles 10 or usually 15-25 mm. long or, if shorter, the blades

dentate; acorns 17 mm. long and 14 mm. broad or smaller.

Leaves glabrous beneath except for axillary tufts of hairs, about

1.5-2 cm. wide Q. acatenangensis.

Leaves resinous-puberulent beneath, mostly 3-5 cm. wide.

Q. conspersa.

Leaves neither acuminate nor lanceolate, mostly oblong or ovate,
abruptly aristate at the apex, crisped-pubescent or lanate be-
neath.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 375

Leaves rather sparsely short-tomentose beneath, smooth or at

most low-bullate Q. crispipilis.

Leaves rather densely thick-tomentose beneath or finally glabrate,
never sparsely tomentose or short-tomentose, bullate.

Q. crispipilis var. pannosifolia.
Fruit annual.

Scales of the cup very loosely appressed Q. Benthami.

Scales of the cup rather closely appressed.

Twigs velutinous-tomentose with short hairs or tardily glabrate.

Q. hondurensis.

Twigs laxly stellate-tomentose with long hairs or almost wholly
glabrous.

Leaves long-acuminate Q. crispifolia.

Leaves short-acuminate or obtuse Q. borucasana.

Leaves toothed or the margins aristate.

Leaves very large, thin, typically obovate, coarsely dentate or merely long-
aristate above the middle, the lower surface persistently and densely

short-tomentose, but the veins glabrous Q. candicans.

Leaves medium-sized or, if large, not short-tomentose on the lower surface,

variously lanate to glabrous.
Leaves persistently, densely, and laxly lanate beneath, or partially glabrate,

bullate-granular; fruit biennial.

Twigs coarse, 3-5 mm. thick, persistently velvety-tomentose; leaves
waxy-glaucous beneath where denuded of pubescence.

Q. brachystachys.

Twigs 2 or rarely 3 mm. thick, loosely tomentose or glabrate or rather
persistently pubescent but not velvety, not waxy-glaucous beneath

when denuded Q. crispipilis var. pannosifolia.

Leaves glabrous or loosely tomentose and soon glabrate, or with axillary
tufts of hairs, or puberulent beneath, or, if bullate-granular, then the
fruit annual, otherwise the fruit either annual or biennial.
Leaves bullate-granular beneath, the veins strongly impressed on the

upper surface Q. tristis.

Leaves smooth and not at all bullate beneath, the veins slightly if at all

impressed on the upper surface.

Leaves very narrow, elongate, and with tapering tips, finely dentate
or merely aristate; petioles 1-3 mm. long Q. flagellifera.

Leaves broad or, if narrow, not as much as 5 times as long as wide and
not attenuate-tipped, coarsely dentate or, if merely aristate, then
not elongate; petioles much more than 5 mm. long or else the blades
coarsely dentate.

Fruit very large, the acorns more than 3 cm. broad; scales of the cup
corky-thickened at the base at maturity. Fruit biennial.

Q. Skinneri.

Fruit moderate-sized or small, the acorns usually less than 2 cm.
broad; scales of the cup thin and flat or if basally thickened
this resulting from swelling of buds in their axils and rarely
from their own thickening.

Acorns about 2 cm. broad, almost globose; shell of the acorn 3 mm.
thick or more, very hard; scales of the cup corky-thickened
at the base at maturity; fruit biennial Q. Skinneri.

Acorns less than 2 cm. broad, ovoid or depressed, never subglobose;
shell of the acorn 1.5 mm. thick or less; scales of the cup thin
or thickened at the base at maturity.

Fruit annual Q. anglohondurensis.

Fruit biennial.

376 FIELDIANA: BOTANY, VOLUME 24

Leaves evidently aristate-tipped at the acute apex, typically
yellow-glandular-puberulent beneath, or rarely glabrate
except for axillary tufts of hairs; margins of the cup
coarsely inrolled Q. conspersa.

Leaves not markedly aristate-tipped at the acute apex,
glabrous beneath or with axillary tufts of hairs; margins
of the cup not inrolled Q. acatenangensis.

Quercus aaata Muller, U. S. Dept. Agr. Misc. Publ. 477: 27.
pis. 23-26. 1942.

Moist mixed mountain forest, 1,800-2,200 meters; Jalapa (Volcan
Jumay); Guatemala (Volcan de Pacaya, above Las Calderas; type,
Standley 58466). Honduras; Costa Rica.

A large tree, sometimes 22 meters high with a trunk 2 meters in diameter,
the twigs slender, 1.5-2.5 mm. thick, strigose and stellate-pubescent with appressed
pubescence, becoming glabrous and gray or whitish; buds rounded-ovoid to
ellipsoid, 3-5 mm. long, glabrous, stramineous; leaves thin but firm, 6-16 cm.
long, 2-6 cm. wide, elliptic or oblanceolate to oblong-obovate or rhombic, much or
only slightly narrowed toward the apex, the tip narrowly or broadly rounded,
attenuate to the very narrowly cuneate to rounded base, subentire or dentate near
the apex, the margins plane or slightly revolute, deep green and dull above,
glabrous except for the costa, this strigose toward the base, lustrous or dull beneath,
strigose or fulvous-tomentose on the costa, otherwise glabrous, the lateral nerves
10-15 pairs, slightly elevated on the upper surface; petioles 2-7 mm. long, usually
about 4 mm.; fruits annual, solitary or geminate, pedunculate; cup about 2.5 cm.
broad, hemispheric or shallowly cup-shaped, the scales ovate with abruptly con-
tracted oblong tips, these laxly appressed, the scales much thickened and velutinous
at the base, the apices brown and strigose; acorn ovoid, 2-2.5 cm. long, scarcely
2 cm. broad, glabrate, one-third to one-half included in the cup.

Called "encino" in Honduras.

Quercus acatenangensis Trelease, Mem. Nat. Acad. Sci. 20:
163. pi. 320. 1924 (type from Guatemala, without definite locality,
Warscewicz 47, 48). Q. longifolia Liebm. Dansk. Vid. Selsk. Forh.
185. 1854, not Q. longifolia Raf ., 1838 (type same as of Q. acatenan-
gensis, which is a renaming of Q. longifolia Liebm.). Q. acutifolia
var. longifolia A. DC. in DC. Prodr. 16, pt. 2: 67. 1864. Q. xalapensis
var. longifolia Wenzig, Jahrb. Bot. Gart. Berlin 3: 210. 1884. Q.
Donnell-Smithii Trelease, Mem. Nat. Acad. Sci. 20: 162. pi. 318.
1924 (in part, but not the collection cited as the type, which is
Q. sapotaefolia) . Q. ambivenulosa Trelease, Journ. Wash. Acad. Sci.
23: 62. 1933 (type from Chichavac, Chimaltenango, A.F. Skutch 14).
Encino; Encino sunuj, Sunuj (Sacatepe"quez) ; Masket (Volcan de
Santa Maria); Duraznillo (Zacapa; probably an erroneous name).

Moist or dry, mountain plains and hillsides, in mixed, oak, or
pine-oak forest, sometimes associated with Cupressus and Abies,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 377

often on white-sand hillsides, 1,500-3,300 meters, most common
at the higher elevations; Baja Verapaz; Zacapa; Jalapa; Guatemala;
Sacatepe*quez; Chimaltenango; Solola; Quiche"; Huehuetenango;
Totonicapan; Quezaltenango; San Marcos. Chiapas.

A medium-sized or often very large tree, sometimes 30 meters high or more
with a trunk a meter in diameter, the twigs 1-2 mm. thick, glabrous, or sparsely
stellate-pubescent becoming glabrate and reddish brown, with rather inconspicuous
lenticels; buds about 3 mm. long, ovoid, acute, sparsely pubescent or glabrate,
light brown; leaves rather thick and hard or rarely thin, 3-5 or even 10-15 cm.
long, usually 2-3 but sometimes 1-4.5 cm. wide, typically lanceolate to elliptic
or narrowly ovate, acute or acuminate, not conspicuously aristate-tipped, cuneate
to rarely attenuate at the base, or in some forms rounded or subcordate, entire or
with few obscure teeth toward the apex, in juvenile forms coarsely serrate, the
margins minutely cartilaginous-revolute and somewhat crispate, the upper surface
more or less lustrous, glabrous or slightly pubescent along the costa, the lower
surface less lustrous or dull, glabrous or with rather conspicuous tufts of hairs
in the axils of the nerves, the surface not bullate, the lateral nerves 10-14 pairs,
more or less elevated on both surfaces; petioles 5-10 or sometimes 20 mm. long,
in juvenile forms sometimes very short; staminate catkins 3 cm. long, fulvous-
villous or gray-villous, loosely flowered, the anthers oblong, well exserted; pistillate
catkins 5-10 or even 20 mm. long, the glabrous peduncle usually 2-flowered;
fruits biennial, solitary or geminate, on a peduncle 5-20 mm. long; cup 10-20 mm.
broad, 7-12 mm. high, cup-shaped to turbinate and constricted at the base, the
scales triangular to lanceolate or ovate, the apices usually narrowed but rounded,
sericeous or glabrate; acorn 10-17 mm. long, 8-14 mm. broad, elliptic to ovoid,
sparsely puberulent or glabrate, light brown, one-half or only one-third included
in the cup.

This is one of the abundantly and widely distributed oaks of
Guatemala, and with a wider variety of habitats than most of the
species. It is particularly frequent at the higher elevations in the
mountains, and often grows in moist if not wet situations. The
species has been reported from Guatemala as Q. nitens Mart. & Gal.

Quercus anglohondurensis Muller, U. S. Dept. Agr. Misc.
Publ. 477: 76. pis. 114, 115. 1942.

Moist or wet, mountain forest, about 800 meters; Pete'n (Camp 36,
British Honduras boundary, W. A. Schipp 1249). British Honduras
(type collected in mountain pine ridge, along Rio Frio, San Agustin,
El Cayo District, C. L. Lundell 6615).

A tree of 15-27 meters, the trunk 45 cm. in diameter, the twigs 1-2.5 mm.
thick, laxly fulvous-tomentose at first, soon glabrate and dark reddish brown,
becoming gray; buds 2-3 mm. long, ovoid, glabrate, rather dark brown or grayish
brown and lustrous; leaves rather thin but firm, 7-10 or even 16 cm. long, 2-3.5
or even 5 cm. wide, lanceolate or elliptic-lanceolate, attenuate-acute, attenuate-
cuneate at the base or merely cuneate or sometimes rounded, low-dentate or merely

378 FIELDIANA: BOTANY, VOLUME 24

aristate from the margin, the teeth aristate-tipped, glabrate and lustrous on both
surfaces, or stellate-pubescent along the costa, especially beneath in the axils of
the nerves, the lateral nerves 10-12 pairs, rather prominent on both surfaces;
petioles 5-16 mm. long; pistillate catkins 5-10 mm. long, 1-2-flowered, the
peduncles glabrous; fruits annual, solitary or geminate, on a peduncle 4-8 mm.
long; cup 13-16 mm. broad, 10-11 mm. high, turbinate or deeply cup-shaped,
somewhat constricted at the base, the margin not inrolled, the scales ovate, the
narrow apex rounded, closely appressed, minutely fulvous-puberulent or the brown
margins glabrate; immature acorn 10-13 mm. long and about as broad, transiently
buff-puberulent, probably about half included in the cup at maturity.

This has been reported from the boundary region as Q. acutifolia
Ne'e, a Mexican species.

Quercus Benthami A. DC. in DC. Prodr. 16, pt. 2: 29. 1864.
Q. gemmata Trelease, Mem. Nat. Acad. Sci. 20: 152. pi. 299. 1924
(type from Volcan de San Salvador, El Salvador). Q. undulata Benth.
PI. Hartweg. 81. 1841. notQ. undulata Torr. 1828 (type from Santa
Maria, probably Volcan de Santa Maria, Quezaltenango, Hartweg
563).

Moist or wet, usually mixed, mountain forest, mostly at 1,500-
2,700 meters; Jutiapa (Volcan de Suchitan); Chimaltenango; Solola;
Quezaltenango; San Marcos. Chiapas.

A medium-sized or large tree, the twigs 1.5-3 mm. thick, densely velutinous-
tomentose with short hairs at first, soon glabrate, or sometimes persistently
fulvous-pubescent into the second year, dark reddish brown; buds 5-7 mm. long,
2-3 mm. thick, fusiform, light or dark brown, fulvous-tomentose or glabrate;
leaves moderately thin but firm and coriaceous, 5-8 or sometimes 10 cm. long, 2-3
or sometimes 5 cm. wide, acute to attenuate-acuminate but rarely aristate-tipped,
cuneate at the base or broadly rounded to narrowly acute, entire, the margins
narrowly but distinctly revolute, finely crispate or flat, glabrate on the upper sur-
face and somewhat lustrous, coarsely fulvous-tomentose beneath along the costa
and in the axils of the nerves, otherwise glabrate and lustrous, the lateral nerves
10-12 pairs, prominent on both surfaces; petioles 1-2 cm. long; fruits annual,
solitary or geminate, subsessile or on a peduncle 3-5 mm. long; cup 20-22 mm.
broad, hemispheric, rounded or somewhat constricted at the base, the margins not
inrolled, the scales oblong or narrowly ovate, their apices narrowed but truncate
or rounded, laxly appressed, finely fulvous-tomentose; acorn depressed-globose or
hemispheric, densely fulvous-sericeous or glabrate and light brown, one-half or
more included.

Quercus borucasana Trelease, Mem. Nat. Acad. Sci. 20: 161.
pi. 315. 1924.

Cloud forest, growing with Abies, 2,500-3,200 meters; Zacapa
(Volcan Gemelos, Sierra de las Minas); Huehuetenango (Cerro
Canana, Sierra de los Cuchumatanes). Costa Rica.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 379

A tree of 9-12 meters or sometimes larger, in Costa Rica as much as 30 meters
high, the twigs glabrate and dark reddish brown; buds 1.5-2 mm. long, ovoid or
round, obtuse; leaves thick, chartaceous, 3-7 cm. long, 1.25-2.5 cm. broad, elliptic
or oblong to narrowly lanceolate, acute and aristate-tipped, base cuneate, entire,
the margins slightly thickened but scarcely revolute, lustrous and glabrous above,
beneath glabrous or nearly so, the lateral nerves 10-12 pairs; fruits annual, small,
solitary or paired on glabrous peduncles 2-5 mm. long; cup 10-15 mm. broad,
about 4-5 mm. deep, cup-shaped or more shallow, rounded or slightly constricted
at the base, the scales ovate, broadly rounded at the narrowed apex, closely
appressed, puberulent to glabrate except at the brown glabrous apex and margins;
acorns 10-15 mm. long, 9-13 mm. broad, ovoid to subrotund, fulvous-sericeous-
puberulent, light brown where abraded, about one-fifth or less included.

The two Guatemalan specimens had been determined previously
by Dr. Muller as atypical Q. eugeniaefolia, but he has recently shown
them to be conspecific with Q. borucasana (Madrono 10: 136. 1950).

Quercus brachystachys Benth. PI. Hartweg. 91. 1842. Roble;
Encino; Masket (Quezaltenango) ; Pat&n (Volcan de Agua, Sacate-
pe'quez); Col (Huehuetenango).

Moist to dry, mountain forest, often or usually associated with
pines and often forming extensive forests, 1,500-2,600 meters; Jalapa;
Guatemala (type from Cuesta de Mixco near San Lucas, Hartweg
618); Sacatepe"quez; Chimaltenango; Quiche"; Huehuetenango; Que-
zaltenango. Chiapas.

A medium-sized or large tree, the branchlets 3-5 mm. thick, covered with a
persistent, gray or brownish, velvety tomentum, reddish brown where denuded;
buds 3 mm. long or more, ovoid; leaves very thick and coriaceous, 4-15 cm. long,
2-10 cm. wide, obovate or sometimes oblong or ovate, acuminate to broadly obtuse
but aristate-tipped, cordate to truncate at the base, repandly and rather coarsely
dentate, the teeth aristate-tipped or reduced to aristae, the margins somewhat
revolute, more or less lustrous above, glabrous except on the stellate-tomentose
costa and nerves, persistently fulvous-tomentose beneath, the surface where
denuded bullate-granular and somewhat waxy-glaucous, the lateral nerves usually
8-10 pairs, strongly impressed on the upper surface, elevated beneath; petioles
mostly 12-22 mm. long; staminate catkins about 7 cm. long, tomentose, rather
laxly flowered, the anthers glabrous, mucronate, well exserted; pistillate catkins
1-2 or rarely 5 cm. long, 2-4-flowered or sometimes 10-flowered, the peduncle
tomentose; fruits biennial, medium-sized, solitary, geminate or ternate on a
peduncle 1-2 cm. long and 3-4 mm. thick; cup about 15 mm. broad and 10 mm.
high, often constricted at the base, the margin not inrolled, the scales ovate,
obtuse, thin, closely appressed, puberulent or with glabrate margins; acorn 15-18
mm. long, 10-12 mm. broad, ovoid or narrowly ovoid, glabrous, light brown,
about half included.

This probably is the species reported from Guatemala by Hemsley
as Q. crassifolia Humb. & Bonpl., a Mexican species. It is one of
the most easily recognized of Guatemalan species because of its large

380 FIELDIANA: BOTANY, VOLUME 24

and broad, very thick, handsome leaves, covered beneath with a
lax or often very dense, soft felt of brownish color. These leaves
often form dense heaps under the trees in late winter or early spring.
This species is an important and sometimes the principal element of
the mountain oak forests of the central and western regions of
Guatemala.

Quercus candicans Ne'e, Anal. Cienc. Nat. 3: 277. 1801. Q.
calophylla Schlecht. & Cham. Linnaea 5: 79. 1830. Q. chimaltenan-
gensis f. gemmata Muller, Amer. Midi. Nat. 18: 855. 1937 (type from
Nebaj, Quiche", A.F. Skutch 1654).

Rather dry, openly forested mountain slopes or in quebradas or
barrancos, about 1,700-2,000 meters; Jalapa; Solola; Quiche"; Hue-
huetenango. Central and southern Mexico.

A large tree, sometimes 23 meters high with a trunk 45 cm. in diameter;
branchlets 4-5 mm. thick, glabrate or rather persistently and shortly stellate-
tomentose, dark reddish brown or grayish; buds 5-6 mm. long, acute, the scales
sometimes ciliate; leaves rather thin but firm and hard, 10-23 cm. long, 4-14 cm.
wide, generally obovate, sometimes elliptic-oblong but broadest above the middle,
acuminate and aristate-tipped or rarely rounded at the apex, subcordate or truncate
at the base, the margins undulate or coarsely dentate, the teeth with aristate tips,
usually glabrate above, somewhat lustrous, densely, persistently, and closely buff-
tomentose beneath with matted stellate hairs, the costa and nerves glabrous, the
lateral nerves 8-14 pairs; petioles 15-20 or sometimes 40 mm. long; staminate
catkins 5-6 cm. long, villous, laxly flowered, the anthers apiculate, somewhat
exserted; pistillate catkins 1.5-2 cm. long, 2-3-flowered; fruit biennial, solitary or
geminate on a stout peduncle 8-12 mm. long; cup 16-20 mm. broad, 10-12 mm.
high, hemispheric or deeper, the margin sometimes inrolled, the scales ovate to
lanceolate, thin, closely or rather laxly appressed, rounded at the apex, puberulent;
acorn 15-18 mm. long, 11-14 mm. broad, ovoid, at first sericeous-puberulent,
becoming glabrate and light brown, about one-third included in the cup.

Quercus conspersa Benth. PI. Hartweg. 92. 1842 (type from
mountains of Las Casillas, Hartweg 617; probably in Santa Rosa).
Q. acutifolia var. conspersa A. DC. in DC. Prodr. 16, pt. 2: 66.
1864. Q. correpta Trelease, Mem. Nat. Acad. Sci. 20: 153. pi. 300.
1924 (excluding the detached fruit, which is Q. sapotaefolia; type
from Guatemala, Warscewicz 25, the locality unknown). Q. con-
spersa f . ovatifolia Trelease, op. cit. 192. pi. 389 (type from Guatemala,
Warscewicz 28). Q. conspersa f. caudata Trelease, loc. cit. (type col-
lected between Guatemala and Coban, F. C. Lehmann 1320) . Encino;
Roble; Sical (Huehuetenango) ; Huite (Zacapa); Bans (Huehuete-
nango).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 381

Usually in pine-oak or oak forest, on moist or dry mountain hill-
sides, 1,000-2,700 meters; Baja Verapaz; Zacapa; Jalapa; Santa
Rosa; Guatemala; Sacatepe"quez; Chimaltenango; Solold; Quiche*;
Huehuetenango; Quezaltenango; San Marcos. Southern Mexico;
Honduras.

A large or medium-sized tree, the twigs 1.5-2.5 or rarely 4 mm. in diameter,
minutely stellate-pubescent becoming glabrate and reddish brown, with a few
inconspicuous lenticels, becoming gray the second season; buds 3-4 mm. long,
narrowly ovoid, acute, glabrate or the apex fulvous-tomentose; leaves rather thick
and very hard, 6-20 cm. long, 2-10 cm. wide, usually lanceolate and long-acute
but varying from oblong to ovate or obovate and attenuate-acute to merely acute,
cuneate to unequally rounded or rarely cordate at the base, entire to coarsely
dentate, the teeth aristate-tipped or reduced to aristae, antrorse or sometimes
widely spreading, the margins rather coarsely cartilaginous but scarcely revolute,
the upper surface glabrous and somewhat lustrous, the lower surface glabrate or
typically fulvous-puberulent and stellate-barbate in the axils of the nerves, the
lateral nerves 9-15 pairs, both the nerves and veins prominent on both surfaces;
petioles mostly 8-30 mm. long, puberulent or glabrate; staminate catkins 6-8 cm.
long, the rachis densely tomentose or glabrate, rather loosely flowered, the anthers
ellipsoid, apiculate, moderately exserted; pistillate catkins 5-20 mm. long, 2-6-
flowered; fruits biennial, solitary or geminate, on a peduncle 3-10 mm. long;
cup 15-18 or rarely 22 mm. broad, 7-10 or rarely 13 mm. high, cup-shaped to
goblet-shaped, the base rounded or somewhat constricted, the margin coarsely
inrolled and sometimes inflated, the scales ovate to triangular, the apices rounded
and often much narrowed, rather closely appressed, fulvous-pubescent or in age
gray-pubescent or glabrate; acorn about 16, rarely 20 mm. long, 13 or rarely
20 mm. broad, ovoid to subglobose, rounded at the apex, minutely puberulent,
becoming glabrate and light brown, about one-third included in the cup.

This has been reported from Guatemala as Q. Sartorii Liebm., a
Mexican species. It is one of the most abundant oaks of Guatemala,
especially in Huehuetenango.

Quercus corrugata Hook. Icon. PL 5: pis. 403, 404- 1842
(type from Cerro del Tambor, Chimaltenango[?] Skinner 5). Q.
cyclobalanoides Trelease, Proc. Amer. Phil. Soc. 54: 11. pi. 3. 1915
(type from Chiapas). Q. Reevesii Trelease, Mem. Nat. Acad. Sci.
20: 45. pi. 22. 1924 (type from mountains west of Volcan de Santa
Maria, Quezaltenango, E. Reeves). Q. corrugata var. granulifera
Trelease, loc. cit. pi. 24 (type from Guatemala, without definite
locality, Warscewicz 11). Q. corrugata var. ipalensis Trelease, loc.
cit. (type from Volcan de Ipala, Chiquimula, around the lake,
H. Pittier 1869). Chicharro (San Marcos).

Moist or dry mountain forest, 950-1,700 meters; Chiquimula;
Alta Verapaz; Suchitepequez; Retalhuleu; Quezaltenango; San

382 FIELDIANA: BOTANY, VOLUME 24

Marcos. Chiapas; British Honduras; El Salvador; Honduras;
Panama.

A medium-sized or large tree, sometimes 20 meters high or more, the twigs
slender or coarse, 1.5-5 mm. thick, glabrous or at first strigose but soon glabrate,
brown at first, becoming gray; buds 3 mm. long, round-ovoid, glabrous, grayish
brown; leaves thick and rather hard, 5 to usually 15 or even 25 cm. long, 2-5 or
as much as 7 cm. wide, lanceolate to oblanceolate or sometimes elliptic to obovate,
acute to attenuate-acuminate, with a narrowly rounded or acute tip, cuneate to
rounded at the base, coarsely dentate with the teeth directed forward and mucro-
nate, entire toward the base, the margin narrowly revolute or plane, somewhat
lustrous and glabrous on both surfaces, the old leaves somewhat bullate-granular
above or smooth, the lateral nerves 12-18 pairs, prominulous above; petioles 15-40
mm. long or rarely shorter, glabrous or puberulent; staminate catkins 5-6 cm.
long, sparsely villous, laxly flowered, the anthers much exserted; pistillate catkins
5 mm. long, 1-2-flowered; fruits annual, solitary on a peduncle 5-10 mm. long,
rather large; cup 3-4 or even 6 cm. broad, shallowly cup-shaped or hemispheric,
very thick, the scales broadly ovate with narrowed apices, much thickened basally,
appressed, closely tomentose; acorn subglobose to ovoid or oblong, 3-5 cm. in
diameter, typically corrugate longitudinally but often smooth, dark brown, sparsely
villous at first, soon glabrate, about half included in the cup.

Called "roble" in El Salvador; "encino bianco" (Honduras). Two
collections from Huehuetenango are noteworthy in having the leaves
stellate-puberulent beneath. These have been determined by Dr.
Muller with the remark that they are "atypical but not significantly
so."

Quercus crispifolia Trelease, Mem. Nat. Acad. Sci. 20: 147.
pi. 286. 1924 (type from Tapachula, Chiapas). Q. amphioxys Tre-
lease, op. cit. 141. pi. 268 (type from Volcan de San Salvador, El
Salvador). Q. incrassata Trelease, op. cit. 147. pi. 287 (type col-
lected near Tapachula, Chiapas). Encino; Roble; Roble amarillo.

Moist or wet, mixed, mountain forest, 1,300-2,700 meters; Alta
Verapaz; Chiquimula; Jalapa; San Marcos. Chiapas; El Salvador.

A large or medium-sized tree, the twigs 1-3 mm. thick, laxly fulvous-stellate-
tomentose at first, soon glabrate and grayish or reddish brown, with prominent
or inconspicuous lenticels; buds 5-6 mm. long, oblong-fusiform, acute, light
brown, glabrous, the scales ciliate; leaves thin but hard and rather coriaceous, 10
to usually 15-20 or rarely 25 cm. long, 3 or 5 or even 7.5 cm. wide, oblanceolate
to linear-lanceolate, acuminate to attenuate or flagellate, inconspicuously aristate-
tipped, attenuate below, the base cuneate to narrowly rounded or rarely subcordate,
entire, the upper surface somewhat lustrous, glabrous or stellate-pubescent at the
base of the costa, the lower surface similar, usually somewhat more conspicuously
stellate-pubescent along the base of the costa, the lateral nerves 15-20 on each
side; petioles 5 to rarely 10 mm. long, dark red at the base, laxly fulvous-tomentose
becoming glabrate; fruits biennial (?), the cup unknown; acorn broadly ovoid,

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 383

25-30 mm. long, 22-26 mm. broad, minutely sericeous-puberulent becoming gla-
brate and brown, included only at the base.

Quercus crispipilis Trelease, Mem. Nat. Acad. Sci. 20: 184.
pi. 370. 1924. Q. cerifera Trelease, op. cit. 184. pi. 370. 1924 (type
from Chiapas). Q. cinnamomea Trelease, op. cit. 184. pi. 371 (type
collected between Quiche" and Totonicapan, 0. F. Cook 27). Ma-ach
(Huehuetenango) .

Rather dry plains and hillsides in the mountains, 1,400-2,700
meters; Chimaltenango; Solola; Quiche"; Huehuetenango (type from
Guaxacana, Seler 2667); Totonicapan; Quezaltenango. Chiapas.

A large or medium-sized tree, the twigs 1.5-3 mm. thick, dark reddish brown,
with a sparse or dense, stellate, buff tomentum, the lenticels few, scarcely promi-
nent, the branchlets becoming grayish; buds scarcely 2 mm. long, round-ovoid,
glabrous, the scales sometimes ciliate, brown; leaves thin but firm and chartaceous,
3-8 or even 12 cm. long, 1-3 cm. wide, oblong-elliptic to oblong-lanceolate or
rarely oblanceolate, acute or usually rounded at the apex, aristate-tipped, rounded
or subcordate at the base, rarely subcuneate, entire, the margins minutely revolute
and often undulate-crispate, the upper surface rather dull, glabrous or stellate-
pubescent about the base of the costa, the lower surface dull or somewhat lustrous,
inconspicuously low-bullate or smooth, with a more or less dense, buff, stellate
tomentum, tardily glabrate or persistently hairy, the lateral nerves 12-15 or
even 20 pairs; fruits biennial, solitary, geminate, or several on a peduncle, 2-7
or rarely 10 mm. long; cup about 12 mm. broad, 10 mm. high, turbinate, constricted
at the base, the margin inrolled, the scales ovate-lanceolate, rounded at the apex,
rather closely appressed, grayish-puberulent; acorn about 12 mm. long and 8 mm.
broad, glabrous or slightly sericeous, light brown, about one-half or two-thirds
included in the cup.

This seems to be one of the common oaks of Huehuetenango,
where oaks and pines usually are the dominant trees of the land-
scape, at least in the drier regions of that department.

Quercus crispipilis var. pannosifolia Muller, U. S. Dept. Agr.
Misc. Publ. 477. 79. 1942. Q. brachystachys f. caerulea Trelease,
Mem. Nat. Acad. Sci. 20: 130. pi. 240. 1924 (type from barranco
below Quezaltenango, Trelease 29). Q. brachystachys f. venulosa
Trelease, loc. cit. (type from barranco below Quezaltenango, Trelease
28). Q. Skutchii Trelease, Journ. Wash. Acad. Sci. 23: 61. 1933
(type from Chichavac, Chimaltenango, A. F. Skutch 44). Q. chi-
chavacana Trelease, op. cit. 62 (type from Chichavac, Chimaltenango,
Skutch 57). Q. chimaltenangana Trelease, loc. cit. (type from Chi-
chavac, Chimaltenango, Skutch 62). Q. aristigera Trelease, op. cit.
63 (type from Chichavac, Chimaltenango, Skutch 86). Q. chicha-
vacana f. undulata Muller, Amer. Midi. Nat. 18: 852. 1937 (type

384 FIELDIANA: BOTANY, VOLUME 24

from Chichavac, Chimaltenango, Skutch 689). Q. chichavacana L
sublobata Muller, loc. cit. (type from Chichavac, Skutch 712). Q.
chichavacana f. oblanceolata Muller, loc. cit. (type from Chichavac,
Skutch 681). Q. pannosifolia Muller, op. cit. 855 (type from Chi-
chavac, Skutch 698). Encino; Masket (Volcan de Santa Maria,
Quezaltenango).

Usually on dry plains or hillsides in oak or pine-oak forest, some-
times with Juniperus, 1,400-2,900 meters; Guatemala; Chimalte-
nango; Quich^ ; Huehuetenango; Quezaltenango. Chiapas.

A large or medium-sized tree, similar in most respects to the typical form of
the species; leaves thick and firm, 3-10 or even 14 cm. long, 1.5-4 or even 6 cm.
wide, oblong-elliptic to obovate or oblanceolate, rarely lanceolate, usually broadest
above the middle, rounded to acute or rarely acuminate at the apex, aristate-
tipped, cuneate to rounded, truncate, or cordate at the base, entire or coarsely
several-dentate or sublobate, the lobes broadly rounded but mucronate, the margins
often coarsely crispate, the upper surface somewhat lustrous, sparsely stellate-
pubescent becoming glabrate or persistently pubescent about the base of the
costa, the lower surface dull, densely velutinous-stellate-tomentose with buff or
brownish, curled hairs becoming subglabrate or persistently flocculent to velutinous,
the surface low-bullate; petioles 4-12 mm. long; fruits solitary or geminate on a
glabrous peduncle 3-10 mm. long; cup 12-20 mm. broad, 10-12 mm. high, hemi-
spheric to cup-shaped or turbinate, the scales oblong or ovate, truncate, rather
laxly appressed; acorn ovoid to subellipsoid, 15-18 mm. long, 10-14 mm. broad,
glabrous, light brown, half or more included in the cup.

Quercus flagellifera Trelease, Mem. Nat. Acad. Sci. 20: 162.
pi. 319. 1924. Encino.

Moist or wet, usually mixed, mountain forest, 1,250-2,500
meters; endemic, but to be expected in Chiapas; Alta Verapaz (type
from Secoyonte", near Finca Sepacuite', 0. F. Cook & R. F. Griggs
607); Chiquimula (Volcan de Quezaltepeque) ; El Progreso (Sierra
de las Minas); Solola; Suchitepequez; San Marcos (Volcan de
Tacana); Huehuetenango (Sierra de los Cuchumatanes).

A tree of 15-20 meters, the twigs slender, 1-2 mm. thick, stellate- villous at
first becoming glabrate and reddish brown; leaves rather thin but becoming
coriaceous, mostly 8-16 cm. long and 1.5-2.5 cm. wide, linear-lanceolate to linear-
oblanceolate, narrowly long-attenuate, often aristate-tipped, gradually attenuate
to the cuneate or very narrowly rounded base, entire or rarely with a few aristate
teeth toward the apex, the margins rather conspicuously but very narrowly
revolute, somewhat lustrous on the upper surface, glabrous or inconspicuously
pubescent, especially near the base of the costa, lustrous beneath, glabrous or the
costa sometimes sparsely stellate-pubescent, the lateral nerves mostly 18-24 pairs,
divergent at a wide angle, plane or prominulous on the upper surface; petiole about
2 mm. long, tardily glabrate; acorns unknown.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 385

Quercus hondurensis Trelease, Mem. Nat. Acad. Sci. 20: 140.
pi. 266. 1924. Q. comayaguana Trelease in Standl. Journ. Arnold
Arb. 11: 25. 1930 (type from Honduras). Q. guayabalana Trelease
in Standl. Field Mus. Bot. 8: 6. 1930 (type from Honduras).

In pine-oak forest on mountain slopes, 800-1,500 meters; Zacapa
(Sierra de las Minas); Huehuetenango (Sierra de los Cuchumatanes).
British Honduras (mountain pine ridge, San Agustin); El Salvador;
Honduras.

A large or medium-sized tree, the twigs 2-4 mm. thick, persistently velutinous-
tomentose with a yellow or buff tomentum, becoming dark reddish brown; buds
5 mm. long, broadly fusiform, apparently glabrous; leaves thick and coriaceous,
8-12 or even 18 cm. long, 3-6 cm. wide, lance-elliptic to oblong or oblanceolate,
obtuse to usually acute or acuminate or rarely rounded at the apex, sometimes
aristate-tipped, cordate or rounded at the base, entire or slightly undulate and
usually crispate, the margins minutely or coarsely revolute, somewhat lustrous
on the upper surface, minutely stellate-pubescent at first, soon glabrate, the
costa persistently tomentose, the lower surface similar, not bullate, the lateral
nerves 8 to usually 10-12 pairs, slightly impressed above, very prominent beneath;
petioles 5-10 mm. long, tomentose like the twigs; staminate catkins 5-10 cm. long,
villous, sparsely flowered, the anthers well exserted; fruits annual, solitary or
geminate, the peduncle 5-10 mm. long, yellow-tomentose or glabrate; young cups
rather small, half-round or constricted at the base, the scales ovate, thin, buff-
tomentose, the narrowed apices rounded, closely appressed; young acorns sericeous,
becoming glabrate and light brown, becoming exserted when half grown.

Called "encino" and "encino de cerro" in Honduras; "roble
negro," "roble amarillo," "roble mamilca" (El Salvador).

Quercus insignis Mart. & Gal. Bull. Acad. Brux. 10, pt. 1:
219. 1843. Q. Schippii Standl. Carnegie Inst. Wash. Publ. 461: 53.
1936 (type from Camp 36, boundary between Pete"n and British
Honduras, W. A. Schipp 1248).

Moist or wet, mountain forest, about 850 meters; Pete"n. South-
ern Mexico; British Honduras; Honduras; Costa Rica.

A tree about 30 meters high with a trunk a meter in diameter, the twigs
4-6 mm. thick, densely yellow-tomentose at first, becoming glaucous or gray;
buds 10-15 mm. long, ovoid, glabrous; leaves rather thick and hard, large, 10-28
cm. long, 4-10 cm. wide, elliptic to oblong-elliptic or narrowly obovate, acute,
with a rounded tip, cuneate to rounded or truncate at the base, subentire to undu-
late or rarely dentate, the upper surface somewhat lustrous, glabrous, or puberulent
along the nerves, the costa tomentose, the lower surface dull, fulvous-villous-
tomentose or glabrate, the lateral nerves 18-20 pairs, elevated on the upper surface
and prominent beneath; petioles 8-25 mm. long, densely fulvous-tomentose;
fruits annual, solitary, subsessile, very large at maturity; cup 4-8 cm. broad,
saucer-shaped or deeper, often contracted at the base, the scales very coarse,
oblong or attenuate, much thickened and very broad at the base, the narrow apices

386 FIELDIANA: BOTANY, VOLUME 24

laxly appressed or spreading, densely fulvous-tomentose or in age glabrate; acorn
4-7 cm. broad, depressed-hemispheric, longitudinally striate, very thick-shelled,
about half included in the cup.

This, like some of the other local species, is noteworthy for its
very large acorns, of a size never found in temperate North America.

Quercus oleoides Schlecht. & Cham. Linnaea 5: 79. 1830.
Q. oleoides var. australis Trelease, Mem. Nat. Acad. Sci. 20: 114.
pis. 192, 193. 1924. Encino negro; Roblecito.

Moist or dry plains or hillsides, often in lowland pine forest, 300
meters or lower; Pete"n; Alta Verapaz; Izabal; Zacapa; Chiquimula.
Southern Mexico; British Honduras; Honduras; Costa Rica (Guana-
caste).

Usually a small tree with a short trunk and a dense, somewhat depressed,
spreading crown; branchlets densely and shortly stellate-tomentose, becoming
glabrate or persistently puberulent, gray; buds about 2 mm. long, dark reddish
brown, glabrate; leaves persistent, thick and stiff, 4-8 cm. long and 2-4 cm. wide,
or sometimes 10 cm. long and 6 cm. wide, obovate to oblanceolate or rarely elliptic
or rhombic, broadly rounded to acute at the apex, cuneate or subobtuse at the
base, entire or sometimes with a few mucronate teeth, especially toward the apex,
stellate-puberulent on the upper surface, becoming glabrate or persistently pubes-
cent, somewhat lustrous, densely and minutely stellate-pubescent beneath with an
appressed tomentum, the veins often glabrate, the lateral nerves 6-8 pairs; petioles
4-6 mm. long, or sometimes 2-10 mm.; staminate catkins about 3 cm. long, the
rachis puberulent, rather closely flowered, the anthers puberulent, scarcely ex-
serted; pistillate catkins 5-25 mm. long, 1-8-flowered; fruits annual, solitary,
geminate, or several on a peduncle 1-6 cm. long; cup turbinate or hemispheric,
10-15 mm. broad, the scales flat or somewhat carinate, not evidently thickened
basally except on very young cups, subacute, gray-puberulent; acorn 18-25 mm.
long, 13-18 mm. broad, ovoid or ellipsoid, glabrous, light brown, about one-third
or sometimes only one-fourth included in the cup.

Called "encino prieto" and "hojaviushi" in Oaxaca; "encino"
(Honduras); "roble" (Chiapas). A collection from the Department
of Guatemala doubtless was taken from a tree planted in Guatemala,
probably in Finca La Aurora. This species is related to the common
live oak (Quercus virginiana Mill.) of the Gulf region of the United
States. It is a lowland tree in Central America, seldom if ever form-
ing pure stands but occurring as isolated individuals in forests of
pine or mixed trees.

Quercus oocarpa Liebm. Dansk. Vid. Selsk. Forth. 184. 1854
(type from Guatemala, Warscewicz 50a, the locality unknown).
Q. Warscewiczii Liebm. op. cit. 187 (based on the same collection
as Q. oocarpa). Q. Yunckeri Trelease in Yuncker, Field Mus. Bot.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 387

17: 358. 1938 (type from Honduras). Roble; Encino; Ji (Huehue-
tenango).

Wet to dry, mixed, oak, or pine-oak forest, sometimes in cloud
forest but also on open rocky mountain slopes, 800-2,400 meters;
Alta Verapaz; Baja Verapaz; Zacapa; Chiquimula; Guatemala;
Huehuetenango. Honduras; Costa Rica; Panama.

A medium-sized or large tree, sometimes 25 meters high or more, the twigs
rather stout, 3-5 mm. thick, at first densely fulvous-tomentose, becoming glabrate
and gray or light brown, with a few scarcely evident lenticels; buds oblong, acute,
4 mm. long, glabrous; leaves thin and papery or rather firm, 10-30 cm. long, 3-14
cm. wide, oblanceolate to obovate or narrowly elliptic, attenuate-acute, narrowly
rounded or cuneate at the base, undulately or acutely low-dentate except near
the entire base, the upper surface somewhat lustrous, glabrous or glabrate, when
young often minutely puberulent, the costa often persistently fulvous-tomentose,
the lower surface dull, persistently and sparsely villous, especially on the nerves
and veins, the lateral nerves 14-22 pairs; petioles 3-5 mm. long, densely or sparsely
fulvous-tomentose; fruits annual, rather large, solitary or geminate on a peduncle
5-18 mm. long; cup 3-4 cm. broad, openly goblet-shaped or cup-shaped, the
scales triangular-ovate to oblong, narrowly obtuse, densely short-pubescent, laxly
appressed; acorn depressed-subglobose or ellipsoid, 2.5 cm. long, 3 cm. broad,
one-half to only one-fourth included in the cup.

This species, and perhaps some of the others also, often bear
on their branches great numbers of solid, hard, and heavy galls.
These vary greatly in size, some of them being as much as 15 cm.
in diameter and at times probably weighing several pounds. A
study of the oak galls of Guatemala would be of considerable en-
tomological interest, for there seems to be a great variety in their
forms.

Quercus pacayana Muller, U. S. Dept. Agr. Misc. Publ. 477:
30. pi. 30. 1942.

Moist or wet, mixed, mountain forest, 1,800-2,600 meters;
endemic; Zacapa (region of Cerro de Monos, Sierra de las Minas);
Guatemala (upper slopes of Volcan de Pacaya, above Las Calderas;
type, Standley 58467).

A large tree, the twigs 1.5-3 mm. thick, glabrate, pale brown becoming
gray; buds rounded-ovoid, 3 mm. long, glabrous, light brown; leaves thick and
rigid, 4-8 cm. long, 2-3.5 cm. wide, drying somewhat yellowish, elliptic to usually
narrowly obovate, broadly rounded at the apex, subcordate or usually rounded at
the base, subentire or crenately low-dentate, especially above the middle, the
margins narrowly revolute, dull on the upper surface, glabrous or sparsely strigose
on the costa, glabrous beneath except on the strigose and fulvous-tomentose costa,
the lateral nerves 10-12 pairs, somewhat impressed on the upper surface; petioles
4-12 mm. long, usually about 6 mm., dark reddish brown; acorns unknown.

388 FIELDIANA: BOTANY, VOLUME 24

Quercus peduncularis Ne'e, Anal. Cienc. Nat. 3: 270. 1801.
Q. callosa Benth. PI. Hartweg. 91. 1842 (type Hartweg 616; Guate-
mala, "in montibus Las Casillas, Ingenio de Ayarza, Petaxa, Mixco,
etc."). Q. arachnoidea Trelease, Mem. Nat. Acad. Sci. 20: 59. pi. 55.
1924 (type from Volcan de San Salvador, El Salvador). Q. Barbeyana
Trelease, op. cit. 67. pi. 76 (type collected between Ipala and Ama-
tillo, Chiquimula, F. C.Lehmann 1711). Q. barbanthera Trelease, op.
cit. 68. pi. 81 (type from "road to Petapa," Guatemala, Skinner in
1845). Q. barbanthera var. calva Trelease, loc. cit. (type from Gar-
rucha, Chimaltenango, Heyde &Lux 3152, in part). Q. peduncularis
subsp. callosa A. Camus, Monogr. Quercus 2: 599. 1939. Q. aguana
Trelease in Yuncker, Field Mus. Bot. 9: 281. 1940. Encino; Roble;
Col (Huehuetenango).

Moist or dry plains and hillsides, often forming almost pure
stands of considerable extent, or more often associated with other
species to form mixed oak forests, or very often associated with pines,
1,000-3,000 meters; Baja Verapaz; Zacapa; Chiquimula; Jalapa;
Jutiapa; Santa Rosa; Escuintla; Guatemala; Sacatepe"quez; Chimal-
tenango; Solola; Quiche" ; Huehuetenango; Quezaltenango; San
Marcos. Southern Mexico; El Salvador; Honduras.

Usually a medium-sized tree with short trunk and spreading crown, sometimes
18 meters high, often only a shrub, the trunk sometimes almost a meter in diameter;
twigs 1.5-5 mm. in diameter, brown, with numerous pale lenticels, glabrous or at
first usually densely stellate-tomentose; buds 2-5 mm. long, ovoid or lanceolate,
obtuse or acute, at first sparsely tomentose, glabrate and reddish brown; leaves
thick and coriaceous, 6-16 cm. long and 3-10 cm. wide, usually about 12 cm. long
and 7 cm. wide, obovate to oblanceolate or elliptic, broadly rounded to obtuse or
abruptly acute at the apex, narrowed to the cordate or rarely rounded or even
cuneate base, undulately low-dentate or acutely serrate, subentire only near the
base, the upper surface stellate-tomentose becoming glabrate or persistently
pubescent along the costa, somewhat lustrous, the lower surface densely or sparsely
tomentose, persistently pubescent or glabrate, dull when denuded, the surface
depressed-bullate, green or waxy-glaucous, the lateral nerves mostly 10-12 pairs,
impressed on the upper surface, very prominent beneath; petioles 3-5 mm. long,
dark reddish brown, stellate-pubescent or glabrate; staminate catkins 4-7 cm.
long, at length loosely flowered, the rachis yellow-villous, the anthers glabrous,
moderately exserted; pistillate catkins 1-5 cm. long, with 2-4 or sometimes more
numerous flowers scattered toward the end of the yellow-villous peduncle; fruits
annual, solitary or several, subsessile or usually pedunculate; cup 15-18 mm.
broad, shallowly saucer-shaped to usually hemispheric, the scales ovate to lanceo-
late, more or less thickened basally, yellowish-pubescent or grayish-puberulent,
the apices thin, appressed, narrowly rounded, glabrate; acorn about 15 mm.
long and 12 mm. broad, ovoid, densely pubescent at first, becoming glabrate
except about the apex, light to very dark brown, one-third to one-fourth included
in the cup.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 389

Called "roble negro" in El Salvador. This is probably the most
widely distributed of all the oaks of Guatemala, if one may judge
from our own collections, which are very ample and were taken more
or less at random. It is one of the abundant trees of the mountains
of Guatemala, Chimaltenango, Quiche", and Huehuetenango, and
probably the dominant one in some regions. Naturally it is highly
variable, and some of the extreme forms look decidedly different
from the mass of material, but there is no reason for supposing that
they should be separated specifically or otherwise. The trees are
heavily infested with insect galls, as are many of the other Guate-
malan oaks. The galls, appearing on the leaves and younger branches,
are mostly globose and with a very dense covering of cotton-like,
very long and soft, bright red or pink hairs.

Quercus peduncularis var. sublanosa (Trelease) Muller, U. S.
Dept. Agr. Misc. Publ. 477; 34. pis. 38, 39. 1942. Q. achoteana
Trelease in Yuncker, Field Mus. Bot. 17: 356. 1938 (type from
Honduras). Q. achoteana var. sublanosa Trelease, loc. cit. (type
from Honduras).

Pine-oak or oak forest in the mountains, in dry or moist situa-
tions, 1,100-2,000 meters; Baja Verapaz; Zacapa; Jalapa; Jutiapa;
Sacatepe"quez; Chimaltenango; Huehuetenango. British Honduras;
El Salvador; Honduras; Nicaragua.

A medium-sized or large tree, in most respects similar to the typical form
of the species but less variable in leaf shape; leaves rather thin, mostly 10-18 cm.
long and 5-10 cm. wide, obovate, broadly rounded to rather abruptly acuminate
at the apex, much narrowed to the rounded or cordate base, regularly undulate-
dentate or serrate with mucronate teeth, the upper surface glabrate or the base
of the costa puberulent, the lower surface glabrate or persistently stellate-puberu-
lent or villous, the surface flat-bullate, the lateral nerves 12-16 pairs; petioles 3-7
mm. long; pistillate catkins 2-4-flowered, the peduncle tomentose or glabrate;
fruit usually evidently pedunculate, the cup about 18 mm. broad, hemispheric,
the scales lanceolate, thickened basally, puberulent or tomentose; acorn ovoid,
about 18 mm. long, glabrous, one-third to one-half included in the cup.

Called "roble" or "roble belloto" in El Salvador.

Quercus pilaria Trelease, Mem. Nat. Acad. Sci. 20: 44. pi. 19.
1924 (type from Tapachula, Chiapas). Q. Yousei Trelease in
Yuncker, Field Mus. Bot. 17: 358. 1938 (type from Honduras).
Encino; Chicharro (Quezaltenango).

Moist or wet, usually mixed, mountain forest, 500-2,400 meters;
Alta Verapaz; Escuintla; Guatemala; Solola; Suchitepequez; Que-
zaltenango; San Marcos. Chiapas; Honduras; Costa Rica.

390 FIELDIANA: BOTANY, VOLUME 24

A medium-sized or large tree, sometimes 25 meters high, the twigs slender, 1-2
mm. thick, sparsely pilose and brown, becoming glabrate and gray, with a few
scarcely prominent lenticels; buds subglobose, 3 mm. long, glabrate, the scales
ciliate; leaves moderately thick and hard, 6-24 cm. long and 2.5-7 cm. wide,
usually about 15 cm. long and 4 cm. wide, lanceolate or oblanceolate, attenuate-
acute, subcordate to cuneate at the base, coarsely crenate-serrate above the
middle, the teeth directed forward, mucronate, somewhat lustrous on both surfaces,
the costa and nerves somewhat sericeous at first but soon glabrate, the lateral
nerves 12-15 pairs; petioles to rarely 4 mm. long, much thickened at the base,
glabrate.

The acorns of this species are not known with certainty, but it
is believed that a detached acorn from the vicinity of Coban, de-
pressed-globose and 4.5 cm. broad, belongs here.

Quercus pilicaulis Trelease, Mem. Nat. Acad. Sci. 20: 67. pi.
77. 1924 (type from Guatemala, the locality unknown, Warscewicz
43). Q. tomentosa var. bullata A. DC. in DC. Prodr. 16, pt. 2: 33.
1864 (type from Guatemala, Warscewicz 15). Q. tomentosa var.
abbreviata A. DC. loc. cit. (type from Guatemala, Warscewicz 43).
Q. pilicaulis f. macrodonta Trelease, Mem. Nat. Acad. Sci. 20: 68.
pi. 78. 1924 (type from Guatemala, Warscewicz 15). Q. pilicaulis f.
obovalis Trelease, loc. cit. pi. 79 (type from Guatemala, Warscewicz
18). Q. pilicaulis f. Hurteri Trelease, loc. cit. pi. 80 (type from
Quezaltenango, Trelease 32). Q. pilicaulis f. armata Trelease, loc.
cit. pi. 79 (type from Volcan de Santa Maria, Quezaltenango, E. W.
Nelson 3721). Q. pilicaulis f. concava Muller, Amer. Midi. Nat. 18:
851. 1937 (type from Chichavac, Chimaltenango, A.F. Skutch 685).
Q. pilicaulis f. elongata Muller, loc. cit. (type from Chichavac,
Skutch 687). Q. pilicaulis f. exserta Muller, op. cit. 852 (type from
Chichavac, Skutch 661). Q. peduncularis subsp. pilicaulis A. Camus,
Monogr. Quercus 2: 601. pi. 197. 1939. Q. peduncularis subsp.
Hurteri A. Camus, op. cit. 603. pi. 197. Encino; Masket (Volcan
de Santa Maria, Quezaltenango).

Dry or moist plains or hillsides in the mountains, in mixed, oak,
or pine-oak forest, sometimes on white-sand hillsides, 1,100-2,800
meters; Baja Verapaz; Zacapa; Jutiapa; Guatemala; Sacatepe"quez;
Chimaltenango; Solola; Huehuetenango; Quezaltenango; San Marcos.
Chiapas.

A large or medium-sized tree, the twigs stout, 2-4 mm. thick, brown or becom-
ing gray, with or without prominent lenticels, at first densely fulvous-tomentose,
soon glabrate or rather persistently pubescent; buds 2-3 mm. long, broadly ovoid
to rounded, brown, glabrate; leaves very thick and leathery, 5-12 or even 20 cm.
long, 2.5-7 or as much as 12 cm. wide, typically obovate to subrotund or almost
oblanceolate, usually broadly rounded at the apex, cordate or subcordate at the

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 391

base, subentire to undulate, apiculate-undulate, or with coarse, acute or obtuse
teeth, the margins cartilaginous and narrowly revolute, the upper surface at first
puberulent, soon glabrate except along the costa and nerves, somewhat lustrous,
the lower surface persistently tomentose with a dense or sparse, buff tomentum,
the denuded surface rather prominently bullate, the lateral nerves 7-10 pairs;
petioles 3-7 mm. long, dark brown, tomentose or glabrate; staminate catkins
6-10 cm. long, fulvous-pilose, loosely flowered, the anthers glabrous, much exserted;
pistillate catkins 1-10 cm. long or longer, 2-15-flowered distally or along the stel-
late-pubescent peduncle; fruits annual, on long or short peduncles, 2-15 cm. long;
cup usually about 14 mm. broad, sometimes 20 mm. or only 8 mm., usually
hemispheric but sometimes deeply cup-shaped or deeply saucer-shaped, the
margin straight or sometimes flaring, the scales narrowly ovate or cuneate, some-
times thickened basally, the apices narrowly rounded, thin, often laxly appressed,
densely puberulent or tomentose, buff or the glabrate tips deep red; acorn 15-23
mm. long, 8-15 mm. broad, ovoid to oblong or long-conic, obtuse or acute, light
brown, glabrate, one-third to one-fifth included in the cup.

This is a common tree of the mountains, especially in the Occi-
dente. It is particularly abundant on Volcan de Santa Maria and
is apparently the only oak there at middle and high elevations. It
ascends there to about 3,450 meters, a higher elevation than is
reached commonly in Guatemala by oak trees.

Quercus polymorpha Schlecht. & Cham. Linnaea 5: 78. 1830
(type from Veracruz). Q. guatimalensis A. DC. in DC. Prodr. 16,
pt. 2: 78. 1864 (type from Guatemala, Warscewicz 37, the locality
unknown). Q. turbinata Liebm. Dansk. Vid. Selsk. Forh. 186.
1854, not Q. turbinata Blume, 1825 (type, Warscewicz 37). Roble.

Moist or dry slopes, usually in oak forest, 1,000-2,000 meters;
Zacapa; Chiquimula; Jalapa; Huehuetenango. Mexico; Honduras.

A small or medium-sized tree, the twigs 2-3 mm. thick, reddish brown, at
first laxly buff -tomentose, soon glabrate, with numerous pale lenticels; buds 3-5
mm. long, ovoid or lance-ovoid, acute, reddish brown, glabrate or rather persist-
ently pubescent; leaves rather thick and stiff, 5-15 cm. long, 3-8 cm. wide, oblong-
elliptic to ovate, lanceolate, or obovate, broadly or narrowly rounded at the
apex and often emarginate, rarely acuminate but with a rounded tip, cordate or
rounded at the base, entire or crenately coarsely mucronate-dentate above the
middle, the upper surface dull or lustrous, glabrate, the lower surface somewhat
waxy-glaucous, rather persistently buff-floccose or glabrate, the lateral nerves
10-14 pairs, impressed above, very prominent beneath, the veins also prominent
and sharply elevated beneath; petioles 6 to usually 15-25 mm. long, glabrate,
reddish brown; staminate catkins 6-8 cm. long, floccose, rather densely flowered,
the anthers glabrous, much exserted; pistillate catkins 5-10 mm. long, 1-2-flowered;
fruits annual, solitary or geminate on a peduncle 5-30 mm. long; cup about 15 mm.
broad, hemispheric, the scales conspicuously thickened basally, acute and appressed
at the apex, puberulent; acorn 20-25 mm. long, 12 mm. broad, about one-half
included in the cup.

392 FIELDIANA: BOTANY, VOLUME 24

Quercus purulhana Trelease, Mem. Nat. Acad. Sci. 20: 63.
pi 64. 1924.

Moist or wet, mountain forest; Baja Verapaz (type from Cuesta
de Quililha, near Purulha, H. Pittier 163; collected also near Rabinal,
between Sabinal and Rabinal, and between Santa Rosa and Salama).
British Honduras (Great Southern Pine Ridge).

Twigs moderate or rather coarse, 2-4 mm. thick, reddish brown, at first sparsely
villous, becoming glabrous; buds 2 mm. long, ovoid, pilose or glabrate; leaves
moderately thick, 10-20 cm. long, 4-13 cm. wide, obovate to oblong-elliptic, broadly
rounded to acute at the apex, subcordate or deeply cordate at the base, obscurely
to deeply undulate, with rounded or mucronate teeth, the cartilaginous margins
flat or narrowly revolute, somewhat lustrous on the upper surface, at first minutely
puberulent, becoming glabrate, densely fulvous-velutinous beneath with short
hairs, the nerves glabrate, the surface waxy where denuded, the lateral nerves
11-15 pairs, slightly impressed above, rather prominent beneath; petioles 3-5
mm. long, reddish brown, puberulent or glabrate; staminate catkins about 4 cm.
long, laxly flowered, the rachis tomentose; pistillate catkins 2-4 cm. long, 2-4-
flowered; fruits annual, solitary, geminate, or aggregate, the peduncle 1.5-2.5 cm.
long; immature cup 10 mm. broad, apparently becoming deeply cup-shaped, the
scales prominently thickened basally, the apices thin, narrow, laxly appressed,
densely tomentulose.

This species probably is one of the common oaks in the open forest
of pine and oak on the dry mountains of Baja Verapaz. We have
done little collecting there and have not collected this species.

Quercus sapotaefolia Liebm. Dansk. Vid. Selsk. Forh. 185.
1854. Q. microcarpa Leibm. op. cit. 184, not Q. microcarpa Lapeyr.
1813 (type from somewhere in Guatemala, Warscewicz 8). Q.
elliptica var. microcarpa A. DC. in DC. Prodr. 16, pt. 2: 71. 1864.
Q. guatimalensis A. DC. op. cit. 78, in part (Guatemala, Warscewicz
37). Q. parviglans Trelease, Proc. Amer. Phil. Soc. 54: 8. 1915
(new name for Q. microcarpa). Q. parviglans f. polycarpa Trelease,
Mem. Nat. Acad. Sci. 20: 152. pi 299. 1924 (type collected between
Salama and Purulha, Baja Verapaz, 0. F. Cook 291). Q. parviglans
f. Tejadana Trelease, loc. cit. (type from Alotepeque, Chiquimula,
Tejada 209). Q. apanecana Trelease, op. cit. 152. 1924 (type from
El Salvador). Q. correpta Trelease, op. cit. 153. pi 300, in part (Gua-
temala, Warscewicz 25). Q. Donnell-Smithii Trelease, op. cit. 162
(El Zapote, Dept. Guatemala, J. D. Smith 1967). Q. siguatepe-
queana Trelease in Standl. Journ. Arnold Arb. 11: 25. 1930 (type
from Honduras). Q. perseaefolia var. achoteana Trelease in Yuncker,
loc. cit. (type from Honduras). RoUe; Encino; Xaccoy (Baja Vera-
paz).

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 393

Wet to dry, usually mixed oak-pine forest, 800-2,600 meters;
Alta Verapaz; Baja Verapaz; El Progreso; Jalapa; Guatemala;
Quiche* ; Huehuetenango; San Marcos. Chiapas; British Honduras;
El Salvador; Honduras; Costa Rica.

A small to large tree, sometimes 30 meters high, or often only a large shrub,
the twigs 1.5-2.5 mm. thick, laxly tomentose or stellate-pubescent, becoming
glabrate, or persistently floccose, light brown to dark reddish brown, with numerous
pale lenticels, becoming grayish; buds 3 mm. long, acute, glabrous, the scales
sometimes ciliate, reddish brown; leaves often persistent for several years, thick
and coriaceous, 4-8 or often 12 cm. long, 1-4 cm. wide, oblanceolate to oblong or
elliptic-oblong, often broadest above the middle, narrowly rounded and aristate-
tipped to usually broadly or narrowly rounded and not apiculate at the apex,
cuneate to usually narrowly rounded at the base, or sometimes broadly rounded
or even subcordate, entire, the margins coarsely revolute, sometimes crispate,
the upper surface dull or lustrous, glabrous, the lower surface not bullate, some-
what lustrous in age, sometimes waxy-glaucous, usually glabrous, or inconspicu-
ously barbulate in the axils of the nerves, sometimes rather persistently floccose,
the lateral nerves 10-18 pairs, slightly or not at all depressed on the upper surface;
petioles 2-4 or even 7 mm. long, stellate-pubescent to glabrate; staminate catkins
5-8 cm. long, rather laxly flowered, sparsely short-villous, the anthers apiculate,
well exserted; pistillate catkins 1-3-flowered or rarely 6-8-flowered, subsessile or
usually pedunculate, the peduncle 2-20 mm. long; fruits annual, solitary, geminate,
or in 3's or more, subsessile or usually pedunculate, the peduncle sometimes 3 cm.
long, with scattered fruits; cup small or moderate, 8 mm. broad, cup-shaped, the
scales broadly ovate, very thin and tightly appressed, sparsely sericeous or glabrate
and light brown and lustrous; acorn about 15 mm. long and 7 mm. broad, ovoid
or usually narrowly ellipsoid, finely sericeous or glabrate, light brown, about
one-fourth included in the cup.

Sometimes called "encino curtidor" in Honduras; "encino mal-
cote" (El Salvador). This has been reported from Guatemala as
Q. nectandraefolia Liebm. and Q. totutlensis A. DC. It is one of the
abundant oaks in the oak forests that cover much of the Department
of Guatemala, and it is common also in the Coban region, but the
species seems to have a curious, locally restricted distribution in
Guatemala. Since its distribution is so wide in the Department
of Guatemala, it might well be expected in all the areas where oak
forests occur.

Quercus segoviensis Liebm. Dansk. Vid. Selsk. Forh. 186.
1854. Encino.

Moist or dry, mountain forest, with other oaks or in mixed associa-
tions, 1,000-2,400 meters; Zacapa; Jalapa; Guatemala; Chimalte-
nango; Solola; Huehuetenango; San Marcos. Honduras; Nicaragua.

A medium-sized or large tree, the twigs 1.5-5 mm. thick, stellate-tomentose,
becoming glabrate and reddish brown, with numerous prominent pale lenticels;

394 FIELDIANA: BOTANY, VOLUME 24

buds 4-5 mm. long, ovoid or narrowly conic, acute, reddish brown, glabrate or
persistently pubescent about the apex; leaves often drying yellowish brown, 8-15
cm. long, 4-9 cm. wide, broadly obovate to oblanceolate-obovate, broadly rounded
at the apex, usually narrowed to the cordate or rarely cuneate base, erose-crenate
with mucronate crenations, or entire toward the base, the upper surface dull,
glabrate, the lower surface stellate-velutinous, becoming glabrate or somewhat
persistently pubescent, the surface low bullate-granular, the lateral nerves 8-11
pairs, prominent on both surfaces; petioles 3-7 mm. long, reddish, glabrate; cup
20 mm. broad, hemispheric, the scales broadly ovate with narrowed obtuse apices,
closely appressed, tomentulose; acorn broadly ellipsoid, about 20 mm. long and
17 mm. broad, pale grayish brown, glabrate, scarcely one-third included in the cup.

The pink galls found commonly on the leaves are called "chin-
chines de encino."

Quercus Skinneri Benth. Gard. Chron. 1841: 16. 1841; PI.
Hartweg. 90. 1842. Q. grandis Liebm. Dansk. Vid. Selsk. Forh.
183. 1854 (type collected in Guatemala by Warscewicz). Q. chi-
apasensis Trelease, Proc. Amer. Phil. Soc. 54: 9. pi. 2. 1915. Q.
salvadorensis Trelease in Standl. Field Mus. Bot. 8: 5. 1930. Q.
hemipteroides Muller, Amer. Midi. Nat. 18: 853. 1937 (type from
Finca San Francisco, Quiche", A.F. Skutch 1875). Chicharro; Encino.

Usually in moist or wet, mountain forest, or often left in planta-
tions when forest is cleared, frequently seen in cafetales of the
Pacific slope, sometimes planted in regions where not native, as
about Coban, 900-2,100 meters; based on Hartweg 615, the localities
reported by the collector as Pacific mountain slopes of Acatenango,
Medio-monte, and Quezaltenango; Alta Verapaz (probably only in
cultivation); Baja Verapaz; Escuintla; Guatemala (perhaps only
in cultivation); Sacatepe"quez; Chimaltenango; Solola; Quiche";
Quezaltenango; San Marcos. Chiapas; El Salvador; Honduras.

A medium-sized or often very large tree, the twigs 2-4 mm. thick, glabrous or
at first fulvous-tomentose and becoming glabrate, dark reddish brown; buds
4-7 mm. long, ovoid, obtuse, sparsely pubescent or glabrate, light brown; leaves
thin and membranaceous, 8-12 or even 30 cm. long, 3-6 or even 12 cm. wide,
broadly lanceolate or obovate to lanceolate or oblanceolate, attenuate or acuminate,
cuneate to subcordate at the base, coarsely or finely dentate with low or attenuate
teeth, the teeth long-aristate, or the leaves sometimes entire and the teeth repre-
sented by aristae, glabrous on both surfaces or with inconspicuous tufts of hairs
beneath in the axils of the nerves, the lateral nerves 10-15 pairs, prominent on both
surfaces; petioles 2-5 cm. long, or rarely very short; pistillate catkins 5 mm. long,
1-2-flowered at the apex; fruit biennial, large, solitary on a peduncle about 5 mm.
long, very polymorphic at different stages of growth; cup 22-45 mm. broad and
8-20 mm. high at maturity, saucer-shaped to hemispheric, the scales ovate, or
narrowed at the apex, corky-thickened basally, the thin apex closely appressed,
fulvous-tomentulose or in age glabrate; acorn 18-40 mm. long and broad, sub-

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 395

globose to depressed, short-cylindric, or globose-ovoid, the ends usually truncate
or rounded, at first loosely tomentose but soon glabrate, about one-fourth included
in the cup, the shell very thick and hard.

This is one of the most easily recognizable of the local species,
noteworthy for its very large acorns, much larger than those of any
northern oaks. The tree is most abundant in the Pacific bocacosta,
where it grows in dense mixed forest, and is the only species of oak
found in most parts of that area. It often occurs in great abundance,
but seldom is the dominant tree. Many of the trees must reach a
height of much more than 35 meters, with a proportionately thick,
often very high and clean trunk. The name "chicharro," rather
curiously, is given commonly to this species and to no other oak,
at least on the Pacific slope, although the common term "encino"
sometimes is applied to it. The trees seem to grow more rapidly
than other native oaks, and they often are planted about fincas, not
only in the coffee plantations of the Pacific bocacosta but about
Coban, where handsome rows of large trees may be seen in some
localities. The very large acorns are much used in games by children
of the Pacific slope, and the boys often fashion ingenious spinning
tops from them.

Quercus tristis Liebm. Dansk. Vid. Selsk. Forh. 174. 1854
(type from Guatemala, Warscewicz 12, the locality unknown). Q.
castanea A. DC. in DC. Prodr. 16, pt. 2: 72. 1864, in part, not Ne'e,
1801. Q. castanea var. sublobata A. DC. loc. cit. (type from Guate-
mala, Warscewicz 10). Q. tristis f. sublobata Trelease, Mem. Amer.
Acad. Sci. 20: 171. pi. 343. 1924. Q. tristis f. vukani Trelease, op.
cit. 172. pi. 342 (type from Volcan de Pacaya, Guatemala, Scherzer).
Q. tristis f . mixcoensis Trelease, loc. cit. (type collected above Mixco,
Guatemala, Trelease 31). Q. Scherzeri Trelease, loc. cit. pi. 343 (type
from Honduras). Q. consociata Trelease, loc. cit. pi. 345 (type from
Guatemala, Warscewicz 9). Roble; Encino; Encino de la herradura;
Machichi (Huehuetenango).

Moist or dry, often rocky plains and hillsides of the mountains,
generally in oak or pine-oak forest, 1,000-2,600 meters; Chiquimula;
Jalapa; Jutiapa; Guatemala; Sacatep^quez; Chimaltenango; Solola;
Huehuetenango; Quezaltenango; San Marcos. Chiapas; El Salvador.

A large or medium-sized tree, the twigs 1-2.5 mm. in diameter, brown or
reddish brown with scarcely evident or rarely prominulous lenticels, sparsely stel-
late-tomentose at first, soon glabrate or rarely persistently pubescent; buds 4 mm.
long, ovoid to conic, acute, pubescent or glabrate, light or dark brown; leaves
thin but very hard and coriaceous, 3-6 or often 12 cm. long, mostly 1-2.5 but

396 FIELDIANA: BOTANY, VOLUME 24

sometimes 5 cm. wide, oblong or very narrowly obovate-elliptic, almost always
broadest above the middle, acute to broadly rounded at the apex and usually
apiculate or aristate-tipped, subcuneate to rounded or usually cordate at the base,
subentire to characteristically low-aristate-dentate or coarsely dentate with
aristate-tipped teeth, especially toward the apex, the upper surface glabrate or
somewhat pubescent about the base of the costa, somewhat lustrous, impressed-
veined, the lower surface stellate-tomentose becoming glabrate, or somewhat
floccose along the costa and nerves, with prominent reticulate venation, the surface
prominently bullate-granular, the lateral nerves 10-12 pairs; petioles 2-5 or even
15 mm. long, glabrate; staminate catkins 5-6 cm. long, laxly flowered, the rachis
shortly stellate-pubescent, the anthers well exserted; pistillate catkins on very
short peduncles, 1-2-flowered; fruits annual, subsessile or on a peduncle 5 mm. long,
solitary or geminate; cup 10-15 mm. broad, shallowly goblet-shaped or cup-shaped,
the scales ovate to triangular, with rounded apices, very closely appressed, puberu-
lent or becoming glabrate and lustrous, light brown; acorn 12-15 mm. long, 10-12
mm. broad, ovoid-rounded, very obtuse, minutely sericeous becoming glabrate,
about one-third included in the cup or covered at the base only.

This is one of the most abundant of all the Guatemalan oaks.
It forms a high percentage of the oak forests of the central and
western highlands.

URTICACEAE. Nettle Family

Annual or perennial herbs, shrubs, or small trees, often with stinging hairs;
leaves simple, alternate, opposite, or verticillate, entire, dentate, or lobate, usually
bearing punctiform or linear cystoliths on one or both surfaces; stipules usually
present; flowers small, commonly green or whitish, variously arranged, in uni-
sexual or androgynous glomerules, dioecious or monoecious; perianth with 2-5
segments or lobes, sometimes none; stamens as many as the perianth segments and
opposite them, the filaments reflexed in bud, with reversed anthers, the filaments
porrect in anthesis, the anthers erect; ovary superior or slightly inferior (Pouzolzia),
1-celled; style simple, the stigma filiform or penicillate-capitate; ovule 1, erect
or ascending; fruit an achene, sometimes enclosed in the accrescent and fleshy
perianth; embryo straight; endosperm none or scant.

About forty genera, in temperate and tropical regions of both
hemispheres. One other genus is known from Central America,
Gyrotaenia, in Panama.

Leaves opposite or verticillate, those of a pair sometimes very unlike (rarely

appearing alternate in species of Pilea). Plants mostly herbaceous.

Plants armed with stinging hairs Urtica.

Plants without stinging hairs.

Pistillate perianth parted, not enclosing the achene Pilea.

Pistillate perianth shallowly lobate, enclosing the achene Boehmeria.

Leaves alternate.

Pistillate flowers subtended by conspicuous green bracts. Leaves mostly entire.

Plants shrubs; pistillate flowers paniculate Hemistylis.

Plants small herbs; pistillate flowers glomerate in the leaf axils.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 397

Stigma filiform, papillose-plumose on one side; staminate flowers short-
racemose Rousselia.

Stigma penicillate, short; staminate flowers in small cymules. . .Parietaria.
Pistillate flowers not subtended by bracts.
Perianth none in the pistillate flowers.

Flowers glomerate in the leaf axils Phenax.

Flowers in lax panicles or in long thread-like spikes or racemes.

Myriocarpa.
Perianth present in the pistillate flowers.

Stigma penicillate or very short and in fruit uncinate.
Plants without stinging hairs; leaves white-tomentose beneath.

Debregeasia.
Plants with stinging hairs; leaves not white-tomentose beneath.

Plants herbaceous; stigma uncinate in fruit Fleurya.

Plants shrubs or trees; stigma penicillate-capitate Urera.

Stigma linear or filiform, persistent or deciduous.

Plants with stinging hairs; stems succulent, thick Laportea.

Plants without stinging hairs; stems not succulent.

Pistillate perianth strongly nerved; achene crustaceous, lustrous;

stigma deciduous; leaves often entire Pouzolzia.

Pistillate perianth nerveless; achene not crustaceous, dull; stigma
persistent; leaves dentate Boehmeria.

BOEHMERIA Jacquin

Small trees, shrubs, or perennial herbs, without stinging hairs; leaves alternate
or opposite, petiolate, dentate, 3-nerved, often with punctiform cystoliths; flowers
monoecious or dioecious, in small unisexual glomerules, these axillary and sessile
or spicate, the spikes often leafy above, or the inflorescence cymose-paniculate;
staminate perianth 3-5-parted or 4-lobate, a rudimentary ovary present; pistillate
flowers tubular or urceolate, 2-4-dentate; stigma filiform, usually pubescent on
one side; achene enclosed in the persistent withering perianth.

About 45 species, widely distributed in tropical and temperate
regions. One other species is known from southern Central America.

Flower clusters cymose-paniculate; leaves white-tomentose beneath. . . .B. nivea.
Flower clusters sessile in the leaf axils or spicate; leaves not white-tomentose

beneath.
Flower clusters sessile in the leaf axils; leaves all alternate.

Leaves coarsely crenate B. ulmifolia.

Leaves appressed-serrate or subentire B. Pavonii.

Flower clusters spicate, the spike sometimes leafy at the summit.

Spikes spreading or pendent, leafless; leaves all opposite; a shrub or small

tree B. caudata.

Spikes erect or suberect, leafy at the summit; stem leaves opposite, those of
the branches alternate; herb B. cylindrica.

Boehmeria caudata Swartz, Prodr. Veg. Ind. Occ. 34. 1788.
B. guatemalensis Gandoger, Bull. Bot. Soc. France IV. 19: 287. 1919
(type collected in Alta Verapaz, Turckheim).

FIG. 53. Boehmeria caudata. A. Habit of upper portion of pistillate plant;
X M- B. Habit of upper portion of staminate plant; X M- C. Staminate petal
from within; X 13. D. Staminate petal from side; X 13. E. Staminate flower
viewed from above; X 7. F. Stamen; X 7. G. Group of staminate flowers in
position; XI. H. Group of pistillate flowers in position; X 3. I. Seed; X 21.
J. Pistil from side; X 7. K. Pistil from the front; X 7. L. Pistillate flower; X 7.

398

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 399

Alta Verapaz (Senahu, 1,050 meters); San Marcos. Southern
Mexico; Honduras; Costa Rica; West Indies; South America.

A shrub or small tree, sometimes 9 meters high, the young branches densely
white-hirsute; leaves long-petiolate, broadly ovate to elliptic, 5-20 cm. long, 3-10
cm. wide, acuminate or long-acuminate, acute to rounded at the base, closely and
evenly crenate, hispidulous above and rough to the touch, densely velutinous-
pilose beneath; spikes very slender, interrupted, leafless, as much as 25 cm. long;
fruiting perianth ovate or obovate, its margins thin, membranous. (Fig. 53.)

Boehmeria cylindrica (L.) Swartz, Prodr. Veg. Ind. Occ. 34.
1788. Urtica cylindrica L. Sp. PI. 984. 1753.

Damp or wet thickets, often in shallow water, 350-1,400 meters;
Alta Verapaz. United States and Mexico; Honduras; Costa Rica;
Panama; West Indies; South America.

A simple or branched perennial, commonly 50-75 cm. tall, the stems puberulent
or almost glabrous, often densely leafy; leaves on long or short petioles, narrowly
lanceolate to ovate-lanceolate, mostly 5-11 cm. long, long-acuminate or attenuate-
acuminate, obtuse or rounded at the base, coarsely dentate, deep green and
punctate above, somewhat paler beneath, almost glabrous or sometimes copiously
rough-pubescent with short hairs; flower spikes short and about equaling the
petioles, or often greatly elongate, naked below but usually leafy toward the apex.

Like most members of the genus, this plant contains a tough
fiber.

Boehmeria nivea (L.) Gaud, in Freyc. Voy. Bot. 499. 1826.
Urtica nivea L. Sp. PI. 985. 1753. Ramie; Ramio. Ramie.

Native probably of China, but grown in many remote regions;
sometimes planted in central Guatemala, experimentally or as a
curiosity.

A coarse herb commonly 1-1.5 meters tall, the young branches hispid; leaves
large, thin, long-petiolate, broadly ovate or rounded-ovate, often 15 cm. long
and 12 cm. wide, acuminate or caudate-acuminate, broadly cuneate to cordate
at the base, very coarsely dentate, thinly hirsute above, green, very rough to the
touch, whitish beneath and densely tomentose; inflorescences small or large,
often much branched, the flower heads small, globose.

Ramie has been planted in Costa Rica to prevent erosion in
gullies. In some parts of the world it is grown extensively for its
fiber, and it has been planted in large amounts in California and
Louisiana. The fiber is notable for its fineness and strength as well
as its endurance. The greatest obstacle to its development in the
United States has been the difficulty of devising means of removing
the too plentiful resin. If this could be overcome, the fiber would
be found highly useful for the manufacture of long-wearing textiles.

400 FIELDIANA: BOTANY, VOLUME 24

Boehmeria Pavonii Wedd. Ann. Sci. Nat. IV. 1: 202. 1854.

Moist or wet thickets along rocky stream banks; El Progreso
(Sierra de las Minas, near Finca Piamonte, Steyermark 43749).
Peru; Bolivia.

A shrub or small tree, commonly about 2.5 meters high, the branches slender,
pilose with short, white, appressed or ascending hairs; leaves alternate, those of
adjacent nodes similar but usually very unequal, oblong-lanceolate or elliptic-
lanceolate, appressed-serrate or subentire, conspicuously 3-nerved, on rather short
petioles, attenuate-acuminate, acute or obtuse at the base, flat or slightly rugose,
appressed-hispidulous or glabrate above and often lustrous, pale beneath and short-
sericeous, the hairs pointing toward the middle of the areolae; larger leaves 4-16
cm. long, the smaller ones 1-3 cm. long and sessile or short-petiolate; flowers dioe-
cious, rarely monoecious, the flowers densely clustered in the leaf axils, the clusters
6-8 mm. in diameter, unisexual, rarely androgynous.

Boehmeria ulmifolia Wedd. Arch. Mus. Paris 9: 347. 1856. B.
fallax var. ulmifolia Wedd. in DC. Prodr. 16, pt. 1: 198. 1869.

Chichicastillo ; Chichicaste.

Moist or wet forest or thickets, sometimes in dry thickets, often
in second growth, 350-2,800 meters; Pet6n; Alta Verapaz; El
Progreso; Izabal; Chiquimula; Jutiapa; Santa Rosa; Escuintla;
Guatemala; Sacatepe"quez; Suchitepequez; Retalhuleu; Quezalte-
nango; San Marcos. Southern Mexico; British Honduras; Honduras;
Panama.

Usually a shrub of 1-3 meters, sometimes a tree of 6 meters, the branches very
slender, often much elongate, pilose or puberulent to almost glabrous; leaves on
short or long petioles, broadly ovate to narrowly oblong-lanceolate, thin, very
variable in size, attenuate-acuminate, rounded to subacute at the base and often
oblique, coarsely or rather finely crenate or serrate, hispidulous and appressed-
pilose above, densely short-pilose beneath or often almost glabrous, the alternate
leaves unequal, the larger ones 7-15 cm. long and 3-9 cm. wide, the smaller ones
much shorter and sometimes very small and stipule-like; flower clusters small,
dense, brownish, sessile in the leaf axils or at defoliate nodes; fruiting perianth
oblong, narrowed to each end, curved, 1 mm. long, thin-margined, densely pubes-
cent.

Here may belong a Guatemalan record of B. ramiflora Jacq. var.
cuspidata Wedd., although it is possible that this species does occur
in Guatemala. B. ulmifolia is highly variable in respect to leaf form
and pubescence.

DEBREGEASIA Gaudichaud

Shrubs; leaves alternate, petiolate, serrate-crenate, 3-nerved, often gray or
whitish beneath; stipules connate; flowers monoecious or dioecious, glomerate-
capitate, the pistillate receptacle somewhat fleshy-thickened, the glomerules arising

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 401

in the leaf axils or at defoliate nodes, sessile or in cymes, the bracts scarious;
staminate perianth 4-parted, depressed-globose in bud; stamens 4, rarely 3 or 5;
pistillate perianth ovoid or obovoid, contracted at the mouth and minutely dentate,
in fruit succulent and juicy.

About five species, in tropical Asia and eastern Africa.

Debregeasia longifolia (Burm. f.) Wedd. in DC. Prodr. 16,
pt. 1: 235-24. 1869. Urtica longifolia Burm. f. Fl. Ind. 297. 1768.

Abundantly naturalized in some places about Coban, Alta
Verapaz, 1,300 meters, in hedges or wet thickets. Southeastern
Asia and Malaysia.

A shrub 1.5-2.5 meters tall, the stout branches hirsute or hispid; leaves long-
petiolate, thin, lanceolate or oblong-lanceolate, 10-17 cm. long, acuminate or
long-acuminate, rounded or obtuse at the base, rather finely and closely crenate-
serrate, green above, hispidulous, rough to the touch, closely white-tomentose
beneath; flower heads very small, in small lax cymes; fruit heads orange-yellow
or red, about 8 mm. in diameter.

The fruits are said to be edible, and it may be on this account
that the plant was introduced into the Coban region. We have no
knowledge of its having become naturalized elsewhere in America,
and it seems to be scarce in cultivation.

FLEURYA Gaudichaud

Annuals, somewhat succulent, usually provided with stinging hairs; leaves
alternate, petiolate, dentate, 3-nerved, with linear cystoliths; stipules connate;
flowers monoecious or dioecious, the glomerules unisexual or androgynous, spicate
or paniculate in the leaf axils; staminate perianth 4-5-parted, globose or depressed
in bud; stamens 4-5; pistillate perianth of 4 segments, these imbricate, subequal
or very unequal; ovary at first straight, soon oblique, the stigma oblique-ovate or
linear, finally uncinate-inflexed; achene oblique, compressed, exserted from the
perianth.

About eight species in the tropics of both hemispheres. Only
one species is found in North America.

Fleurya aestuans (L.) Gaud, in Freyc. Bot. Voy, 497. 1826.
Urtica aestuans L. Sp. PI. ed. 2. 1397. 1763.

Wet thickets or wet or boggy, open ground, at sea level; Izabal.
British Honduras; Nicaragua; Costa Rica; Panama. West Indies;
tropical South America; Africa.

Plants erect or nearly so, succulent, usually less than a meter tall, the stems
densely hirsute or almost glabrous, often very glandular and usually armed with
stinging hairs; leaves long-petiolate, broadly ovate, 15 cm. long and 12 cm. wide
or usually smaller, acute or acuminate, rounded or subcordate at the base, coarsely

FIG. 54. Fleurya aestuans. A. Habit of upper part of plant; X M- B. Fruit
with bract; X 11. C. Staminate flower; X 6. D. Staminate flower in bud; X 6.

402

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 403

dentate or crenate, sparsely or rather densely hispid on both surfaces, slightly paler
beneath; flowers paniculate, the panicles slender-pedunculate, mostly shorter than
the leaves, green, flower clusters mostly androgynous; achene ovate. (Fig. 54.)

The plant stings. It is of only occasional occurrence along the
Atlantic coast of Central America, and in many places is absent.
Called "nettle," "cow-itch," and "pica" in British Honduras.

HEMISTYLIS Bentham

Shrubs or small trees; leaves alternate, long-petiolate, 3-nerved, entire or
dentate; stipules free; flowers monoecious, the staminate glomerate and spicate,
the pistillate geminate and subtended by two foliaceous bracts, the spikes solitary
or geminate in the leaf axils; pistillate involucres usually inserted near the base of
the spike; staminate perianth 4-parted, subglobose in bud, the segments acute,
valvate; stamens 4; pistillate perianth ovoid, contracted and 2-4-dentate at the
mouth, compressed in fruit and 2-carinate or 2- winged; stigma filiform, curved,
deciduous; achene inclosed in the accrescent perianth, ovoid-conic.

A single species is known in Central America.

Hemistylis odontophylla Wedd. in DC. Prodr. 16, pt. 1:
235-53. 1869.

Damp or wet thickets or forest, 600-900 meters; Santa Rosa.
Venezuela and Colombia.

A slender shrub 2-3 meters tall with few long weak branches, these densely
or sparsely hispidulous; leaves on long or rather short petioles, apparently decid-
uous during the dry season, broadly ovate, 20 cm. long and 15 cm. wide or smaller,
the upper ones reduced, acuminate or long-acuminate, rounded at the base, rather
coarsely dentate, above green, densely puncticulate and hispidulous, slightly rough
to the touch, beneath somewhat paler, soft-pilose; inflorescences forming a long
narrow dense leafy-bracted panicle 25 cm. long; pistillate flowers subtended by
rather large, green, lanceolate to broadly ovate bracts.

This plant presents a rather unusual case of discontinuous dis-
tribution. The large panicles, in which the numerous crowded bracts
are conspicuous, give it an appearance distinct from any other
urticaceous plant of Central America.

LAPORTEA Gaudichaud

Herbs, shrubs, or small trees, often with stinging hairs; leaves alternate,
usually dentate, penninerved or 3-nerved; stipules distinct or connate; flowers
monoecious or dioecious, glomerate or cymose and paniculate, the panicles axillary,
the flowers and fruits often reflexed; staminate perianth 4-5-parted, depressed
in bud, the segments valvate or subimbricate; stamens 4-5; pistillate perianth
of four lobes or segments, these equal or very unequal; ovary at first straight,
soon oblique, the stigma linear; achene oblique, compressed or slightly ventricose,

404 FIELDIANA: BOTANY, VOLUME 24

the perianth membranaceous, scarcely accrescent; pericarp membranaceous or
fleshy.

About 25 species, in tropical and subtropical regions of both
hemispheres, mostly in the Old World. One other Central American
species grows in Nicaragua.

Laportea mexicana (Liebm.) Wedd. in DC. Prodr. 16, pt. 1:
84. 1869. Discocarpus mexicanus Liebm. Dansk. Vid. Selsk. Skrivt.
V. 2: 309. 1851. Laal (Pet&i, Maya); Ortiga (Pet&i).

Wet or moist thickets, 1,200 meters or less; Pete*n; reported from
Guatemala (Rio Amatitlan). Southern Mexico; Salvador.

A shrub or small tree 2-5 meters tall, the branches thick, soft, pale, glabrous;
leaves clustered at the ends of the branches, long-petiolate, broadly ovate, 5-10
cm. long, 3-6 cm. wide, acute, obtuse or rounded at the base, repand-crenate,
green, pilose, abundantly furnished with stinging hairs; flowers dioecious; cymes
laxly much branched, the branches retrorse-setulose, longer than the leaves,
many-flowered; achenes suborbicular, strongly compressed, green, lustrous,
glabrous, the margins narrowly thickened.

In general appearance this is much like the genus Urera, but the
fruits are quite different. To judge from the specimens, the shrub
must bloom when leafless or when the new leaves are beginning to
unfold. Called "chichicaste" and "pan caliente" in Salvador.

MYRIOCARPA Bentham

Shrubs or small trees, without stinging hairs; leaves alternate, usually large,
petiolate, dentate, penninerved and subtrinerved, with cystoliths, these often
radiating from the bases or rudiments of the hairs; stipules connate; flowers
dioecious or sometimes monoecious, in very long and slender, almost thread-like
spikes or racemes, these solitary or fasciculate in the leaf axils or at defoliate
nodes, simple or branched, the minute flowers crowded or usually remote along
the rachis, the staminate mostly sessile, the pistillate sessile or pedicellate; stami-
nate perianth 4-5-parted, the segments obtuse, imbricate; stamens 4-5; pistillate
perianth none; ovary sessile or short-stipitate, compressed; stigma obliquely
oblong, villous-papillose, a conspicuous style present; achene ovoid, compressed,
the margins often thickened, the pericarp thin-crustaceous.

About seven species, ranging from Mexico to Brazil. One other
Central American species is known in Nicaragua.

Pistillate inflorescence paniculate, with short branches; achene long-stipitate,

ciliate with very long hairs. Leaves obovate, glabrous M. obovata.

Pistillate inflorescence simple or of branched but very long and slender, pendulous

racemes; achene sessile or short-stipitate, eciliate or short-ciliate.
Leaves mostly small and less than 5 cm. wide, the cystoliths irregularly dis-
tributed over the upper surface M. heterostachya.

Leaves large, mostly 7-15 cm. wide or larger, the cystoliths of the upper surface
radiating from the center of the areole.

STANDLEY AND STEYERMARK: FLORA OF GUATEMALA 405

Achenes long-ciliate, narrowed to an evident stipe; bractlets divaricate.

M. longipes.
Achenes not ciliate, scarcely narrowed at the base; bractlets appressed.

M. yzabalemis.

Myriocarpa heterostachya Bonn. Smith, Bot. Gaz. 13: 29.
1888. M. heterospicata Bonn. Smith, Bot. Gaz. 12: 133. 1887.

Moist or wet forest, sometimes in thickets on limestone, also
in second growth thickets, 1,600 meters or less; Pete"n; Alta Verapaz
(type from Pansamala, Turckheim 892); Escuintla; Huehuetenango.
British Honduras; Nicaragua.

A shrub or a small tree, sometimes 9 meters high but usually much lower;
leaves on rather short or sometimes long and slender petioles, lanceolate or lance-
ovate, sometimes elliptic-lanceolate, mostly 5-15 cm. long and 3-6 cm. wide,
acuminate or caudate-acuminate, obtuse at the base, 3-nerved, entire or undulate-
dentate, strigose beneath on the veins but elsewhere glabrous or nearly so; flowers
monoecious, the spikes unisexual, very long and slender, pendent, white, pinkish
or dull reddish; staminate spikes few, 7 cm. long or less, once furcate; pistillate
spikes up to 20 cm. long, once furcate near the base; achenes elliptic, conspicuously
stipitate, sparsely strigillose.

Myriocarpa longipes Liebm. Dansk. Vid. Selsk. Skrivt. V. 2:
306. 1851. M. malacophylla Rob. & Bartlett, Proc. Amer. Acad.
43: 50. 1907 (type from Gualan, Zacapa, C. C. Deam 361). Chichi-
caste; Chichicaste manso.

Moist thickets or forest, sometimes in second growth, 200-2,600
meters or less; Alta Verapaz; Pete"n; Izabal; Zacapa; Solola; Suchi-
tepequez; Jalapa; Santa Rosa; Chimaltenango; El Quiche"; Retal-
huleu; Quezaltenango; San Marcos. Southern Mexico; El Salvador;
Nicaragua; Costa Rica.

A large shrub or small tree, usually 3-6 meters tall; leaves thin, on long or
short petioles, broadly ovate or elliptic to obovate-elliptic, mostly 10-30 cm. long
and 8-15 cm. wide, acute or short-acuminate, rounded to broadly cuneate at the
base, closely crenate-dentate, often rugulose and rough above, rather densely
and usually softly pilose or hispidulous beneath; flowers dioecious or rarely
monoecious, the spikes long and slender, pendent, once or twice furcate near the
base, whitish or pinkish, sometimes pale purplish; achenes elliptic, 1-1.5 mm.
long, long-ciliate, narrowed to a short but distinct stipe.

The material is variable in shape and pubescence of the leaves,
and it may be that ultimately it can be divided into two or more
species. The shrub is easily recognized, with other members of the
genus, by its very long and thread-like, whitish spik