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Full text of "Flora of Guatemala"

B R.AR X Y 

OF THE 

UN IVER.SITY 
OF ILLINOIS 



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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^ 

9 



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 



18 



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. 



B 



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. 




D 

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. 



20 



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. 




B 



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 



24 



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. 



29 



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. 



35 



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 



43 



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 



46 



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. 



49 



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.) 




51 



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