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"L I B RAR.Y 

OF THE 

UNIVERSITY 
OF ILLINOIS 

FSo.S 



v. 24- 
pt.? 



BIOLOGY 



10. 7 

FLORA OF GUATEMALA 

PAUL C. STANDLEY 

AND 

LOUIS 0. WILLIAMS 



FIELDIANA: BOTANY 
VOLUME 24, PART VII, NUMBER 1 

Published by 

CHICAGO NATURAL HISTORY MUSEUM 
NOVEMBER 13, 1961 



FLORA OF GUATEMALA 

PART VII 



FLORA OF GUATEMALA 



PAUL C. STANDLEY 

Curator Emeritus of the Herbarium 



AND 



LOUIS O. WILLIAMS 

Curator, Central American Botany 



FIELDIANA: BOTANY 

VOLUME 24, PART VII, NUMBER 1 

Published by 

CHICAGO NATURAL HISTORY MUSEUM 
NOVEMBER 13, 1961 



Library of Congress Catalog Card Number: ^8-3076 



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



6 



7 



CONTENTS 



Families Included in Part VII, Number 1 



Dilleniaceae 2 

Actinidiaceae 10 

Ochnaceae 10 

Marcgraviaceae 16 

Quiinaceae 23 

Theaceae 24 

Guttiferae 36 

Cistaceae. . 61 



Cochlospermaceae 67 

Violaceae 70 

Flacourtiaceae 82 

Turneraceae 109 

Passifloraceae 115 

Caricaceae 146 

Loasaceae. . .152 



Bixaceae 65 Begoniaceae 157 



LIST OF ILLUSTRATIONS 

TEXT FIGURES 

PAGE 

1. Davilla aspera var. Matudae 5 

2. Doliocarpus dentatus 8 

3. Tetracera volubilis 11 

4. Ouratea lucens 14 

5. Sauvagesia erecta 17 

6. Quiina Schippii 25 

7. Cleyera theaeoides 29 

8. Ternstroemia Tepezapote 35 

9. Mammea americana 53 

10. Rheedia edulis 56 

11. Bartholomaea mollis 85 

12. Pleuranthodendron mexicana 101 

13. Erblichia odorata var. mollis Ill 

14. Passiflora capsularis 123 

15. Passiflora edulis 127 

16. Passiflora quadrangularis 142 

17. Carica Papaya 149 

18. Begonia Calderonii 162 

19. Begonia convallariodora 165 

20. Begonia crassicaulis 166 

21. Begonia Franconis 167 

22. Begonia gracilis 169 

23. Begonia heracleifolia 170 

24. Begonia Heydei 172 

25. Begonia peltata 177 

26. Begonia Popenoei 179 

27. Begonia pustulata 180 

28. Begonia Sartorii 182 

29. Begonia trichosepala 184 



Flora of Guatemala 

PARIETALES 

The group of families in part VII, number 1 of the Flora of 
Guatemala is a large order divided by Engler and Diels into 31 
families. More than half (17) of these families are represented in 
Guatemala. Several are common in our flora, and many are not 
well understood. The Guttiferae and the Flacourtiaceae are per- 
haps the most difficult, and the generic entities in both families 
need careful study. Passiflora is the largest genus of this order 
in our flora. 

Plants of the Parietales are most variable and perhaps the group- 
ing is unnatural, but usually they may be characterized by a biseriate 
perianth in which the sepals and the petals may be distinguished 
one from the other, or in some genera the petals may be lacking 
(especially in Flacourtiaceae) or the perianth of tepals (Begon- 
iaceae); the perianth is often pentamerous, but in many genera 
of the order it is not (2-10-merous) ; the petals and the calyx lobes 
may be imbricate (usually) or valvate; the stamens may be as many 
as the petals (or sepals) or often much more numerous; the ovary 
is most often of 3 carpels and unilocular, with parietal placentation 
(axile sometimes in Ochnaceae) and many or numerous ovules; it 
is most often superior but in the Loasaceae and Begoniaceae it 
is inferior, and it is partly inferior in some Flacourtiaceae and 
Theaceae. Unisexual flowers are not uncommon in the order. 

The Parietales are a grouping for convenience rather than a 
phylogenetic one. It may be expected that some of the families 
will be removed to other orders and perhaps new orders established 
for some of these families. 

The genus Saurauia, in this flora, has been separated as the 
Saurauiaceae and will be found in the last pages of the preceding 
part of the flora. The genus has been most often included in the 
Dilleniaceae; but it has also been placed in the Actinidiaceae, which 
would seem to be a reasonably satisfactory disposition of it. 

l 



2 FIELDIANA: BOTANY, VOLUME 24 

DILLENIACEAE 

Trees or shrubs, often scandent, the pubescence usually harsh; leaves alternate, 
entire or dentate, usually with numerous parallel lateral nerves; stipules none, or 
winglike and adnate to the petiole, commonly deciduous; flowers small or medium- 
sized, perfect or polygamous; sepals 4-6, strongly imbricate, persistent; petals 
5-6 or fewer, imbricate, often crumpled in bud, generally deciduous or caducous, 
usually thin and delicate; stamens numerous, rarely definite, hypogynous, free 
or variously united into bundles at the base, commonly persistent; anther cells 
opening by lateral slits or apical pores; carpels free, rarely only 1; ovules 1 or 
more in each carpel, erect from the base or from the inner angle of the carpel; 
styles free; fruiting carpels dehiscent or baccate, the seeds usually with a cristate 
or laciniate aril; endosperm copious, carnose, the embryo minute. 

Eleven genera, in the tropics of both hemispheres. Only the 
following are known in North America. 

Sepals dissimilar, the 2 inner ones erect, valvate and enclosing the fruit; petals 

bright yellow; flowers mostly in terminal panicles; plants scandent. .Davilla 

Sepals all alike, spreading or ascending, not enclosing the fruit; petals usually 

white, not bright yellow. 
Flowers chiefly in terminal panicles; carpels 1-5, distinct, glabrous; woody 

vines Tetracera. 

Flowers axillary or lateral, in panicles or umbels; carpels 1, or 2 and united. 

Carpel of the fruit 1, or didymous; plants scandent Doliocarpus. 

Carpels of the fruit 2; plants erect Curatella. 



CURATELLA L. Sandpaper tree 

Small trees with very harsh, rough pubescence; leaves penninerved; flowers 
white, small, crowded in small, lateral, sessile panicles; sepals 4-5, spreading; 
petals 4-5; anthers oblong, the cells almost parallel; carpels 2, globose at ma- 
turity, dorsally dehiscent or fleshy and indehiscent, coherent at the base; ovules 
geminate, erect; seeds almost completely surrounded by a membranaceous aril. 

Species 2, one West Indian. 

Curatella americana L. Syst. Nat. ed. 10. 1079. 1759. C. ameri- 
cana var. pentagyna Donn.-Sm. Bot. Gaz. 46: 109. 1908 (type from 
Salama, Baja Verapaz, W. A. Kellerman 6499). Chaparro; saha 
(Pete"n, fide Lundell); lengua de vaca. 

Chiefly on dry open or brushy hillsides, often on pine-clad slopes, 
1,200 meters or less; Pete"n; Alta Verapaz; Baja Verapaz; Izabal; 
Chiquimula. Western Mexico; British Honduras to El Salvador and 
Panama. Cuba; South America. 

Mostly 6 meters high or less, the trunk short and thick, the crown spreading, 
the bark with large thin grayish scales that fall easily, the inner bark reddish or 
light brown; leaves short-petiolate or sessile, oval to elliptic-ovate, mostly 12-30 
cm. long, rounded or very obtuse at the apex, often emarginate, abruptly cuneate, 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 3 

decurrent at the base, thick, sinuate or irregularly shallow-denticulate, very sca- 
brous on both surfaces; ill-scented; flowers white or pinkish, long-pedicellate, in 
short sessile lateral panicles, mostly on old wood; petals 5-6 mm. long, glabrous; 
mature carpels 6-7 mm. long, densely hirsute; seeds black. 

Called "yaha" in British Honduras; "caticon" (Tabasco); "ho- 
jaman" (Oaxaca); "malcajaco," "lengua de vaca" (El Salvador). 
Var. pentagyna is a presumably abnormal form with 5 rather than 
the usual 2 carpels of the ovary and fruit. According to Seler, the 
seeds are or were used in Oaxaca for flavoring chocolate. The rough 
leaves contain much silica and are used commonly in tropical 
America as a substitute for sandpaper, to polish articles of wood 
or metal. The bark is utilized in some regions for tanning skins. 
The wood is brown or reddish brown, variegated by the prominent 
rays, rather hard and heavy, with a specific gravity of about 0.77; 
the grain is interwoven, and the texture is coarse; the wood is 
hard to cut and to plane and it is durable. It is employed in various 
regions for fuel, charcoal, fence posts, or even cabinet work, and in 
Central America particularly for making saddle frames. In Guate- 
mala the tree seems to be most plentiful on the pine-clad hillsides 
of Izabal and Baja Verapaz, where often it is associated with Byr- 
sonima crassifolia. This association, without the pines, is character- 
istic of many subarid regions of the whole Pacific coast from El 
Salvador to Panama, and in South America. 

DAVILLA Vandelli 

Scandent shrubs; leaves coriaceous, usually very rough, without stipules; 
flowers paniculate, bright yellow, the panicles mostly small, terminal or arising 
in the upper leaf axils; sepals 5, very unequal, the 2 interior ones larger, very 
concave, accrescent, becoming coriaceous, enclosing the fruit and simulating a 
globose capsule; petals 1-6; filaments thickened at the apex, the anthers short, 
their cells divergent; carpels of the ovary 1-3, the ovules geminate, erect; mature 
carpels indehiscent or rupturing irregularly; seeds almost completely enclosed 
in the aril. 

Species about 30, mostly in Brazil. Only the following are found 
in Central America. 

Larger sepals hirtellous; leaves densely soft-pilose beneath D. Kunthii. 

Larger sepals glabrous; leaves glabrate beneath, usually somewhat pilose along the 
nerves D. aspera var. Matudae. 

Davilla aspera (Aubl.) Ben. var. Matudae (Lundell) L. Wms. 
Fieldiana, Bot. 29: 350. 1961. Davilla Matudae Lundell, Phytolo- 
gia 1: 371. 1940 (type from near Palenque, Chis., Matuda 3647a). 
Chaparro. 



4 FIELDIANA: BOTANY, VOLUME 24 

Moist or wet thickets or forest at or little above sea level, some- 
times in mangrove swamps; Izabal. Mexico through Central Amer- 
ica, except El Salvador. Figure 1. 

A small to large woody vine or liana, the bark exfoliating, ferruginous, the 
branches glabrous or sparsely appressed-hirsute; leaves petiolate, coriaceous, 
mostly oval to rounded-obovate, sometimes oblong-elliptic, 5-16 cm. long, 2-6.5 
cm. wide, rounded to acute at the apex, rounded to acute at the base and often 
abruptly decurrent, entire or undulate, rough or smooth above and glabrous or 
nearly so, rough or smooth beneath, usually with at least a few stiff hairs along the 
costa, often papillose; panicles small or large, 12 cm. long or less, usually many- 
flowered; outer sepals sericeous, the inner ones larger, glabrous or nearly so, in 
fruit 5-8 mm. long; petals bright yellow, fugacious; carpel 1-seeded, the seed 
2.5 mm. long. 

This variety has been confused commonly with Davilla Sagraeana 
A. Rich, and Davilla rugosa Poir., both of which may be synonyms 
of D. aspera var. aspera. It has also been confused with species of 
Tetracera. There is considerable variation in the Central American 
specimens referred to this variety. Called bejuco quemador and bejuco 
corralero in adjacent Mexico. 

Davilla Kunthii St. Hil. PI. Usuel. Bras. 6. 1824-28. Bejuco 
chaparrdn; chaparro; sajab (Pete"n, Maya, fide Lundell). 

Moist or wet forest, often in hilly pine forest, 350-450 meters 
or less; Pete'n; Alta Verapaz; Izabal; Huehuetenango. Southern 
Mexico; British Honduras to Panama. South America. 

A small or large, woody vine, the branches densely hirsute with spreading 
hairs, the bark exfoliating; leaves petiolate, coriaceous, oblong-elliptic to sub- 
orbicular, 6-18 cm. long, broadly rounded to subacute at the apex, rounded to 
acute at the base, scabrous above and very rough to the touch, often lustrous, 
the nerves impressed, the surface rugose, densely soft-pilose beneath, the lateral 
nerves very conspicuous and elevated; panicles small or large, many- flowered; 
sepals densely pilose or hirsute, the inner ones becoming 7-10 mm. long; carpel 1. 

Called "chaparro" in British Honduras and "bejuco de tachicon" 
in Tabasco; "lija" (Oaxaca). The heavy, very rough leaves contain 
much silica and often are used like sandpaper and for cleaning pots 
and dishes. 



DOLIOGARPUS Roland 

Woody vines, glabrous or pubescent, usually not or but slightly scabrous; 
leaves dentate to subentire; panicles few- to many-flowered, or the flowers solitary 
or fasciculate in the leaf axils or at defoliated nodes or lateral, white or yellowish; 
sepals 3 or 5, equal to subequal, spreading; petals usually 3-4, sometimes 2; fila- 
ments thickened at the apex, the anthers short, the cells almost parallel or slightly 




FIG 1 Davilla aspera var. Matudae. A, Branch; X H- B, Flower; X 5. 
C, Flower in fruiting condition with two large sepals enclosing fruit; X 5. 



6 FIELDIANA: BOTANY, VOLUME 24 

divergent; carpels of the globose ovary 1 or 2, the ovules geminate, erect; fruit 
at maturity baccate and indehiscent or coriaceous and 2-valvate or didymous; 
seeds surrounded by a membranaceous or fleshy aril. 

About 25 species in tropical America, chiefly in Brazil. Four 
additional species are known from Central America. Possibly one 
additional species in Guatemala, as yet undescribed. 

Leaves acute to acuminate at apex, elliptic-oblong to oblanceolate-oblong, entire 
to coarsely dentate; flowers fasciculate; pedicels 10-15 mm. long; style 1. 

D. dentatus. 

Leaves rounded at apex, suborbicular-obovate to obovate-oblong, entire or subun- 
dulate; flowers paniculate; pedicels 2-6 mm. long; styles 2 D. coriaceus. 

Doliocarpus coriaceus (Mart. & Zucc.) Gilg, in Engl. & Prantl, 
Nat. Pflanzenfam. 3, Abt. 6: 114. 1893. Pinzona coriaceus Mart. & 
Zucc. Abh. Akad. Muench. 1: 371. 1837. D. nicaraguensis Standl. 
Field Mus. Bot. 4: 233. 1929 (type from region of Braggman's Bluff, 
Nicaragua, Englesing 277). D. belizensis Lundell, Field & Lab. 13: 
6. 1945 (type from British Honduras, Gentle 4389). Water tietie 
(British Honduras). 

Wet lowland forest, British Honduras; Nicaragua; Colombia; 
British Guiana; Brazil. 

A small or large woody vine, the branches glabrous or strigillose, the older 
ones with exfoliating bark; leaves petiolate, the petiole thickened below, narrowly 
margined, 1.3-2 cm. long, 2-2.5 mm. wide, the blade coriaceous, obovate-elliptic 
to broadly or suborbicular-obovate, 7-14 cm. long, 4-10 cm. wide, rounded at 
apex and short-apiculate, obtuse, rounded, or cuneate at the decurrent base, 
entire or subundulate, mostly glabrous on both sides, midrib above minutely 
scabridulous to glabrate, below glabrous to sparsely puberulous, lateral nerves 
slender, prominent, 6-10 pairs, elevated beneath, glabrous or minutely hispidulous 
on the upper surface, glabrous to minutely puberulous on the lower surface, veins 
prominently finely reticulate, especially below; flowers paniculate, the panicles 
axillary, sessile, contracted, closely and many-flowered in early anthesis, becoming 
more lax and more elongated in fruiting stage, the panicles 1-3.5 cm. long; pe- 
duncles 6-10 mm. long, puberulous; bracts ovate to lance-oblong, strigillose 
without; pedicels 2-6 mm. long, slender, puberulous; flowers whitish or greenish, 
fragrant; sepals usually 3, suborbicular to broadly obovate-oblong, 3-3.8 mm. 
long, 2-3 mm. wide, rounded at the apex, persistent, reflexed, ciliolate; petals 
usually 2, obovate-oblong, 4-4.5 mm. long, 2.5-3 mm. wide, rounded at apex; 
stamens numerous, shorter than the sepals; styles 2, 1.25-2 mm. long; fruit didy- 
mous, 4-5 mm. high, 4-7 mm. wide, the locules subglobose, connate above, smooth, 
glabrous. 

This species belongs to the section Pinzona of the genus, char- 
acterized by having two styles instead of one. Study of type material 
of D. belizensis and D. nicaraguensis leaves no doubt that they are 
conspecific with the South American D. coriaceus. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 7 

Doliocarpus dentatus (Aubl.) Standl. Journ. Wash. Acad. Sci. 
15: 286. 1925. Tigarea dentata Aubl. PI. Guian. 920, pi. 351. 1775. 
D. oaxacanus Szysz. Diss. Math. -Nat. Acad. Litt. Cracov. 27: 139. 
1894 (type from Oaxaca). 

Chiefly in wet lowland forest; 200 meters or less; Pete"n; Alta 
Verapaz; Izabal; Huehuetenango. Mexico; British Honduras, along 
the Atlantic coast to Panama; northern South America. Figure 2. 

A small or large, woody vine, the branches glabrous or nearly so, the older 
ones with dark ferruginous, exfoliating bark; leaves petiolate, coriaceous, elliptic- 
oblong to oblong or oblanceolate-oblong, mostly 9-21 cm. long, acute or acumi- 
nate or abruptly acute, narrowly or broadly cuneate at the base, coarsely dentate 
to almost entire, glabrous above and smooth to the touch, dull, paler beneath and 
brownish, often lustrous, at first short-pilose along the costa and veins but in age 
glabrate, usually barbate in the nerve axils, the lateral nerves slender, prominent, 
about 10 pairs; flowers fasciculate on the older branches, slender-pedicellate, the 
pedicels mostly 1-1.5 cm. long, puberulent; flowers 3-4 mm. long, whitish, the 
sepals minutely appressed-pilose; fruit globose, 8-12 mm. in diameter, red. 

This has been reported from Guatemala under the name D. Ro- 
landri Gmel. 

TETRACERA L. 

Scandent shrubs, the pubescence usually rough; leaves coriaceous, conspicu- 
ously nerved; flowers paniculate, the panicles terminal or from the upper leaf axils, 
many-flowered, lax or dense; sepals 4-6, spreading; petals 4-6 or sometimes fewer; 
filaments dilated at the apex, the anthers small, their cells somewhat divergent; 
carpels of the ovary 1-5, acuminate, the ovules numerous, biseriate; mature car- 
pels coriaceous, lustrous, 2-valvate or dehiscent along one suture; seeds 1-5, sur- 
rounded by a lacerate aril. 

About thirty species in the tropics of both hemispheres. No 
other species are known from Central America. 

Sepals sericeous within; carpels of the fruit 2-5. 

Leaves scabrous beneath, very rough to the touch T. volubilis. 

Leaves pilose beneath, not rough to the touch T. mollis. 

Sepals glabrous within; carpels of the fruit 1, or 2-4. 

Carpels of the fruit 1, dark brown; leaves oblong to obovate, mostly closely den- 
tate or serrate T. sessiliflora. 

Carpels of the fruit 2-4, tawny; leaves broadly ovate-elliptic to broadly ovate- 
oblong, entire to sparsely denticulate T. jamaicensis. 

Tetracera jamaicensis DC. Syst. Nat. 1: 399. 1818. T. beli- 
zensis Lundell, Contr. Univ. Mich. Herb. 6: 44. 1941 (type from 
British Honduras, Gentle 2794). Chaparo. 

Thickets or forests. Izabal. British Honduras; Honduras; Nica- 
ragua; Panama; Jamaica. 




FIG. 2. Doliocarpus dentatus. A, Fruiting branch; X ^. B, Fruit; X 
r lower; X 5. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 9 

Woody vine, the branches at first densely and minutely stellate-scabrid inter- 
mixed with scattered, straight, appressed hairs, finally becoming scabrous and 
brownish; leaves long-petiolate, the petioles 1.2-2.5 cm. long, sparingly strigose 
and minutely stellate-scabrid, slightly winged above; leaf blade broadly ovate- 
elliptic to broadly ovate, 7-17 cm. long, 5.5-10 cm. wide, rounded and short 
apiculate at apex, rounded at base, mostly entire or subentire or with a few short 
teeth near apex, both surfaces finely scabrous, the costa bearing a few straight 
hairs, these appressed on lower surface, spreading above, costa and veins slightly 
impressed above, prominent below, the main lateral nerves 11-13, prominently 
reticulate beneath; panicles dense, dioecious; staminate flowers with slender stel- 
late-scabrid pedicels to 8 mm. long; sepals subequal, suborbicular, 5-7 mm. long, 
ciliolate, glabrous within, minutely scabrous without with the outer bearing a few 
simple appressed hairs; petals obovate, to 10 mm. long, glabrous; stamens about 
5 mm. long; carpels of the fruit 2-4, tawny, short-rostrate, 10-15 mm. long, 
glabrous. 

Tetracera mollis Standl. Field Mus. Bot. 8: 25. 1930. Saha 
(Pete'n, Maya, fide Lundell). 

Wet thickets, 300 meters or less, Pete'n; Izabal. British Hon- 
duras, the type from Honey Camp, Orange Walk, Lundell 47. 

A large or small vine, the bark dark ferruginous, exfoliating, the branches 
densely pilose with intermixed, simple and stellate hairs; leaves petiolate, broadly 
oblong or elliptic-oblong, 8-14 cm. long, 4-7 cm. wide, rounded, subacute, or 
abruptly acuminate at the apex, obtuse or rounded at the base and often abruptly 
decurrent, serrulate or repand-dentate, at least toward the apex, very scabrous 
above, densely velutinous-pilose beneath with simple and stellate hairs, the lat- 
eral nerves about 13 pairs, prominent; panicles small or rather large, shorter or 
longer than the leaves; sepals obovate, 7-9 mm. long, obtuse, sericeous on both 
surfaces; carpels of the ovary 3, acuminate, lustrous, 1 cm. long, sparsely ap- 
pressed-pilose; seeds solitary; flowers yellowish white. 

Tetracera sessiliflora Triana & Planch. Ann. Sci. Nat. IV. 17: 
21. 1862. 

Reported by Hemsley from Guatemala; western and southern 
Mexico; British Honduras; Costa Rica; Panama; Colombia. 

A large vine, the young branches hirsute and stellate-pubescent; leaves short- 
petiolate, oblong to obovate, 6-20 cm. long, rounded or obtuse at the apex, often 
abruptly short-pointed, obtuse to rounded at the base and abruptly decurrent, 
remotely dentate to subentire, usually very scabrous on both surfaces, sometimes 
sparsely pilose beneath; panicles mostly large and many-flowered, the flowers 
white; sepals orbicular, densely pubescent; carpel of the fruit only 1, very lustrous, 
6-8 mm. long. 

The names "sumac" (Maya) and "berisa" are reported from 
British Honduras. The flowers are sweet-scented. The stems, as 
in other species, are employed as a substitute for cordage, especially 



10 FIELDIANA: BOTANY, VOLUME 24 

for binding the framework of huts. The rough leaves are sometimes 
used like sandpaper. 

Tetracera volubilis L. Sp. PI. 533. 1753. Chaparro tietie. Be- 
juco de agua; sahaac (Petn, Maya, fide Lundell); cachicdn (Pete"n, 
fide Lundell). 

Moist or dry thickets, sometimes in open forest, 360 meters or 
less; Pete*n; Alta Verapaz; Santa Rosa; Escuintla; Suchitepe"quez; 
Retalhuleu; San Marcos. Southern Mexico; British Honduras to 
El Salvador and Panama; West Indies; South America. Figure 3. 

A small or large vine, the branches brownish, with exfoliating bark, the young 
branches more or less hirsute, glabrate in age; leaves short-petiolate, obovate- 
oblong to obovate, 7-18 cm. long, rounded or abruptly short-pointed at the apex, 
cuneate at the base and often decurrent, serrulate or subentire or often coarsely 
dentate, very scabrous on both surfaces, usually appressed-hirsute beneath on 
the nerves; panicles often very large and dense; sepals orbicular, unequal, 3-5 mm. 
long, sericeous within, scabrous outside, ciliate; carpels of the fruit 3-5, short- 
rostrate, 7-10 mm. long, sparsely pilose at the apex. 

Called "lengua de vaca" in El Salvador. The vine is very com- 
mon in thickets and dry forest of the Pacific lowlands. It is one of 
the more familiar water vines of Central America, the larger stems, 
when cut, yielding a large amount of clear sap that may be drunk in 
place of water when the sources of this fluid are not available, as so 
often happens during the dry months in the lowlands of Central 
America. 

ACTINIDIACEAE 

The genus Saurauia would have been found in this place had the 
system proposed by Engler and Gilg been strictly adhered to. The 
account of the genus will be found under Saurauiaceae, vol. 24, pt. 6: 
428-438. 1949, of this Flora. 

OCHNACEAE 

Trees, shrubs, or herbs, mostly glabrous; leaves simple, alternate, coriaceous 
or membranaceous, usually dentate, often with fine close venation; stipules en- 
tire or pectinate; flowers small or large, yellow, pink, or white, solitary in the leaf 
axils or in terminal or axillary racemes or panicles; sepals 5; petals 5; petaloid 
staminodia often present, 5, distinct, opposite the petals and separated from them 
by a whorl of sterile filaments; fertile stamens 5 or 10; ovary 3-celled at the base, or 
1-celled and 5-6-lobate with each of the lobes 1-celled; fruit capsular or drupaceous. 

About 17 genera, widely distributed in tropical regions. One 
other genus, Cespedesia, a large tree with showy yellow flowers, is 
represented in southern Central America. 




FIG. 3. Tetracera volubilis. A, Fruiting branch; X 
mature carpels; X 4. C, Flower; X 4. 



* B, Flower, showing 



11 



12 FIELDIANA: BOTANY, VOLUME 24 

Plants shrubs or trees; petals yellow; stamens 10; fruit drupaceous Ouratea. 

Plants low herbs; petals pink or white; fertile stamens 5; fruit capsular. 

Sauvagesia. 

OURATEA Aublet 

References: L. A. M. Riley, The Mexican and Central American 
species of Ouratea, Kew Bull. 101-114. 1924. John D. Dwyer, The 
taxonomy of the Mexican, Central American, and West Indian spe- 
cies of Ouratea, Lloydia 7: 121-145. 1944. 

Glabrous shrubs or small trees; leaves alternate, persistent, coriaceous, lus- 
trous, serrate, finely and closely veined; stipules free in American species; flowers 
showy, yellow, in terminal or terminal and axillary panicles or racemes, bracteate, 
the pedicels articulate at the base; sepals 5, usually colored, persistent or decid- 
uous, imbricate; petals 5, somewhat longer than the sepals, obovate, unguiculate, 
imbricate; torus thick, elevated to form a gynophore, lobate; stamens 10, inserted 
on the base of the torus, erect, connivent, the filaments very short; anthers rugose, 
dehiscent by pores; ovary 5-6-parted, the lobes obliquely inserted on the torus; 
styles connate, the stigma simple; ovules solitary in each cell, ascending from the 
base; fruits drupaceous, 5 or by abortion fewer, sessile on the enlarged torus; seed 
erect, the testa membranaceous; cotyledons carnose, plano-convex; radicle very 
short, descending to the hilum. 

Species described more than 100, mostly from the American 
tropics with a few from Africa and Asia. 

The Central American species of Ouratea have "suffered" two 
revisions in relatively recent years but are still in need of a revision 
that takes into account the South American as well as the North 
American species. Some of the names in this treatment will possibly 
be changed when such a revision is made. 

Leaves mostly 30-45 cm. long, usually clustered at the ends of the branches. 
Leaves oblong, the veins not evident beneath; panicles half as long as the leaves. 

O. lucens var. podogyna. 
Leaves narrowly oblanceolate-oblong, the veins conspicuous beneath; panicles 

equaling the leaves 0. Tuerckheimii. 

Leaves smaller, mostly 6-18 cm. long, scattered along the branches. 

Nerves of the leaves impressed on both surfaces 0. nitida. 

Nerves of the leaves not impressed O. lucens. 

Ouratea lucens (HBK.) Engler in Mart. Fl. Bras. 12, pt. 2: 
350. 1876; L. Wms. Fieldiana, Bot. 29: 352. 1961. Xcanlol, xcolol 
(British Honduras). Gomphia lucens HBK. Nov. Gen. & Sp. 7: 219. 
1825 (type from Colombia, Humboldt 1445). Ouratea guatemalensis 
Engler, I.e. 345 (type from Guatemala, Friedrichsthal 780). 0. Peckii 
Riley, Kew Bull. 1924: 109. 1924 (type from British Honduras, Peck 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 13 

617). 0. stenobotrys Riley, I.e. (type from British Honduras, Hooper). 
0. Wrightii (Van Tiegh.) Riley, I.e. 110 (type from Nicaragua). 

Moist and wet thickets, often in second growth, at 700 meters 
or less; Pete"n; Alta Verapaz; Izabal; Esquintla; Retalhuleu. Mexico 
through Central America to Colombia. Figure 4. 

A glabrous shrub or small tree of 1-6 meters, the branches slender, often flex- 
uous; leaves coriaceous, on petioles 3-4 mm. long, lance-oblong or narrowly oblan- 
ceolate, mostly 10-18 cm. long and 3-5.5 cm. wide, acuminate, rounded to acute 
at the base, serrate toward the apex or throughout, the veins conspicuous, not 
impressed; stipules triangular-acuminate, 8-9 mm. long, denticulate, striate; pan- 
icles usually racemiform, few-many-flowered, dense or lax; pedicels 5-10 mm. long 
or in age longer; sepals ovate-lanceolate, the outer ones about 7 mm. long, obtuse; 
petals obovate, 6-9 mm. long; anthers obclavate, about 5 mm. long; drupes 1-5, 
ellipsoid, black, juicy; torus 5 mm. in diameter, bright red. 

Called "coyolillo" and "naranjitto" in El Salvador. Because of 
its bright yellow flowers, this is a rather showy plant, like other 
members of the genus. The young leaves are tinged with bronze. 
The base of the trunk is often enlarged with thick knobby buttresses 
of distinctive form and appearance. The material referred here was 
distributed by Dwyer among five species based upon characters that 
vary too much to serve for specific distinction in the case of these 
Central American plants. 

Ouratea lucens var. podogyna (Donn.-Sm.) L. Wms. Fieldiana, 
Bot. 29: 352. 1961. 0. podogyna Donn.-Sm. Bot. Gaz. 18: 2. 1893 
(type from Guatemala, von Turckheim 1034). 0. insulae Riley, Kew 
Bull. 1924: 106. 1924 (type from Honduras, Gaumer 89). 0. pyra- 
midalis Riley, I.e. 107 (type from Mexico, Rovirosa 495). 

Alta Verapaz; Izabal; Pete*n. Mexico, British Honduras and 
Honduras, possibly Panama. 

Similar to the species but with a more floriferous, larger and more spreading 
panicle; the leaves larger and more prominently serrate. 

Ouratea nitida (Swartz) Engler in Mart. Fl. Bras. 12, pt. 2: 310. 
1876. Ochna nitida Swartz, Prodr. Veg. Ind. Occ. 67. 1788. 

Wet forest, at little above sea level; Pete"n; Izabal. Yucatan 
peninsula of Mexico; British Honduras; Honduras; Nicaragua; Pan- 
ama; Cuba and Jamaica; northern South America. 

A glabrous shrub or a small tree, 6 meters high or less, the trunk as much as 
10 cm. in diameter; leaves on very short petioles, coriaceous, elliptic to elliptic- 
oval, 6-13 cm. long, acute or rather abruptly acuminate, obtuse or rounded at 
the base, serrulate, the nerves and veins impressed on both surfaces; flowers bright 




FIG. 4. Ouratea lucens. A, Flowering branch; X J^. B, Torus, showing two 
persistent drupes; X 2^. C, Sepal; X 2%. D, Portion of inflorescence, showing 
partially dissected flower (3 stamens removed) and buds; X 2H- E, Stamen, 
showing apical pores; X 5. 



14 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 15 

yellow, in pyramidal panicles, these often much exceeding the leaves; pedicels 
5-7 mm. long, or in fruit longer; sepals about 7 mm. long; petals slightly longer 
than the sepals; drupes 1-5, dark blue at maturity, globose, 7 mm. long; seed 
large, globose. 

Called "bastard blossom berries" in British Honduras. 

Ouratea Tuerckheimii Donn.-Sm. Bot. Gaz. 33: 249, t. 10. 
1902. Yepu (Alta Verapaz). 

Wet mixed forest, 350 meters or less; Pete"n; Alta Verapaz (type 
from Cubilqiiitz, Tuerckheim 7829); Panama. 

A glabrous shrub or small tree 3-6 meters high; leaves mostly in clusters of 
3-4 at the ends of the branches, on petioles 6-12 mm. long, oblanceolate-oblong, 
coriaceous, 35-45 cm. long, 8-11 cm. wide, acuminate, obtuse or rounded at the 
base, deep green above, pale green beneath, acutely serrate, the nerves and veins 
prominent beneath; panicles much branched, almost as long as the leaves, the ped- 
icels 10-13 mm. long; sepals linear-oblong, 1 cm. long; petals spatulate-obovate, 
11-12 mm. long; anthers 9 mm. long; drupes globose, 9 mm. in diameter. 

The flowers are said to have the odor of lily of the valley (Conval- 
laria). This species is closely allied to 0. crassinervia Engler. 

SAUVAGESIA L. 

Glabrous, usually small and wiry, erect, annual or perennial herbs; leaves 
small, membranaceous, alternate, entire or serrulate; stipules pectinate-ciliate; 
flowers very small, axillary or in terminal racemes, white or pink; sepals subequal; 
petals equal, convolute; outer staminodia numerous or 5 and alternating with the 
petals, the 5 interior staminodia petaloid, opposite the petals and connivent about 
the stamens and ovary; filaments of the fertile stamens very short, alternate with 
the inner staminodia; anthers linear, the cells laterally dehiscent; ovary with 3 
placentae, the style simple, the stigma obtuse; fruit a small capsule, septicidally 
3-valvate; seeds numerous, small, the testa crustaceous, usually favose-scrobicu- 
late; endosperm carnose, the radicle longer than the cotyledons. 

About 18 species, mostly American, a few in the Old World 
tropics. One additional species is known from Panama. 

Lobes of the stipules filiform, bearing small glands at about the middle; plants 

mostly 10 cm. high or less S. pulchella. 

Lobes of the stipules without glands. 

Sepals 2-2.5 mm. long; plants mostly 10 cm. high or less S. tenella. 

Sepals 5 mm. long; plants mostly 30 cm. high or taller S. erecta. 

Sauvagesia erecta L. Sp. PI. 203. 1753. Yerba deljudio (Izabal, 
fide Blake). 

Usually in moist or wet soil of savannas, open grassy banks, 
moist thickets, or pine forest, 1,450 meters or less; Alta Verapaz; 



16 FIELDIANA: BOTANY, VOLUME 24 

Izabal; Huehuetenango. Southern Mexico; British Honduras to 
Panama; West Indies; South America. Figure 5. 

Plants erect, annual or probably at times perennial, the stems wiry, often red 
or brown, simple or sparsely branched, usually 20-40 cm. high; leaves almost ses- 
sile, elliptic-lanceolate to oblong-lanceolate, usually 1-2 cm. long, acute at each 
end, serrulate; stipules conspicuous, 3-6 mm. long, pectinate-ciliate with very long, 
slender divisions; flowers axillary, the pedicels filiform, mostly shorter than the 
leaves; sepals lanceolate or ovate-lanceolate, 5 mm. long, acuminate, mucronate; 
petals obovate, pink or almost white, 5-6 mm. long; capsule slightly longer than 
the persistent sepals; seeds minute, ellipsoid, deeply pitted. 

This and other species of Sauvagesia are characteristic plants of 
wet savannas and banks of creeks. 

Sauvagesia pulchella Planch, in Seem. Bot. Voy. Herald 80. 
1852. 

Grassy open pine or oak forest, 1,000-1,500 meters (to be ex- 
pected also at much lower elevations); Chiquimula; Jalapa. El Sal- 
vador; Honduras; Costa Rica; Panama. South America. 

Plants annual, erect, mostly 10 cm. high or less, sometimes taller, simple or 
sparsely branched; leaves almost sessile, ovate-lanceolate or oblong-lanceolate, 
10-12 mm. long; stipules 5 mm. long, pectinate-ciliate, the divisions glandular 
and thickened at about the middle; flowers solitary or binate in the upper leaf 
axils, the pedicels about 5 mm. long; sepals 2 mm. long, the petals longer, pink; 
capsule 4 mm. long, acute; seeds scrobiculate. 

Sauvagesia tenella Lam. 111. 2: 119. 1793. 

Open, wet, usually grassy places at low elevations; reported by 
Hemsley from Barbasco, Bernoulli 947, probably in Zacapa. 

Southern Mexico(?); British Honduras; Honduras; Costa Rica; 
Panama. South America. 

Plants annual, usually simple and 10 cm. high or less; leaves almost sessile, 
obovate-lanceolate or oblong-lanceolate, 5-10 mm. long, acute, attenuate to the 
base, remotely glandular-denticulate; stipules 2-4 mm. long, pectinate-ciliate, the 
divisions eglandular; flowers 1-3 in the upper leaf axils, the pedicels 5-10 mm. 
long; open flowers 7-9 mm. broad; sepals lanceolate, aristate-attenuate, subserru- 
late at the apex, white-marginate; petals white or pink, slightly longer than the 
sepals, obovate, obtuse; capsule ovoid, not exceeding the calyx; seeds minute, 
scrobiculate. 

MARCGRAVIACEAE 

Reference: E. Gilg & E. Werdermann, Nat. Pflanzenfam. ed. 2, 
21:94-106. 1925. 




FIG. 5. Sauvagesia erecta. A, Habit; X Y 2 . B, Flower; X 4. 



17 



18 FIELDIANA: BOTANY, VOLUME 24 

Usually woody vines, often epiphytic; leaves alternate, coriaceous, entire, all 
alike, but the leaves of fertile and sterile branches often different; flowers perfect, 
terminal, racemose, umbellate, or paniculate, the inflorescences often pendent; 
bracts often brightly colored, highly variable in form, often modified into nectar- 
secreting organs; bractlets 2, similar to the sepals and usually appressed to them; 
sepals 4-5, imbricate; petals 4-5, united at the base or higher and deciduous as a 
cap; Stamens 3-many, free or united with one another and with the bases of the 
petals; anthers globose or elongate; ovary superior, at first 1-celled, by intrusion 
of the placentae becoming in age 2-many-celled; ovules numerous; style simple, 
short, the stigma 5-radiate; fruit capsular, coriaceous-carnose, irregularly ruptur- 
ing or indehiscent; seeds numerous, small; endosperm scant or none. 

Five genera, confined to tropical America. One other genus, 
Norantea, is represented in southern Central America. 

Flowers umbellate, the central ones sterile; petals united to form a cap. 

Marcgravia. 
Flowers racemose, all fertile; petals free, or united only at the base. 

Bracts spur-like or hat-shaped, hollow; ovary 5-celled Souroubea. 

Bracts globose to semiglobose or carnose-spatulate, solid, not hollow . .Ruyschia. 



MARCGRAVIA L. 

Plants usually scandent and epiphytic, glabrous or nearly so, sometimes ter- 
restrial vines, often very large; sterile plants very different in appearance from the 
adult ones, usually repent on tree trunks; flowers in terminal umbelliform racemes, 
the bracts sac-like, stipitate in the center of the umbel, free from the pedicels; 
sepals 5, imbricate; petals united to form a cap-like structure that falls from the 
flower as a cap; stamens 12-40, free from the corolla, generally somewhat connate 
at the base, the anthers linear, basifixed; ovary many-celled, the stigma sessile, 
obscurely radiate; ovules numerous, horizontal or ascending, anatropous, the pla- 
centae carnose; fruit globose, somewhat car nose or coriaceous, indehiscent or 
irregularly ruptured. 

Species 40 or more, in tropical America, extending northward into 
Mexico. Several others are known in southern Central America. 

Flowers not oblique on the pedicel, the flower in the same axis as the pedicel; 
leaves sessile or practically so. 

Pedicels densely lenticellate, the lenticels elevated M. rectiflora. 

Pedicels not lenticellate M. Schippii. 

Flowers inserted obliquely on the pedicel, the axis of the flower oblique to that of 

the pedicel; leaves short-petiolate. 
Bracts or nectaries with their pedicels about 6 cm. long; leaves of fertile branches 

long-acuminate M. nepenlhoides. 

Bracts with their pedicels 2.5-3 cm. long. 

Leaves narrowly long-acuminate M. guatemalensis. 

Leaves acute or subobtuse M. Gentler. 

Marcgravia Gentle! Lundell, Contr. Univ. Mich. Herb. 6: 46. 
1941. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 19 

Known only from the type, Stann Creek District, Stann Creek 
Valley, Antelope Ridge, British Honduras, P. H. Gentle 3176. 

A woody vine; leaves on stout petioles 3-5 mm. long or less, rigid-coriaceous, 
lance-oblong to obovate-oblong, 4.5-10 cm. long, 2.5-4 cm. wide, acute or sub- 
obtuse, subtruncate or rounded at the base and shallowly emarginate, the lateral 
nerves almost horizontal, inconspicuous; umbels about 15-flowered, the pedicels 
2.5 cm. long or less, puberulent, the flowers inserted obliquely; bracts clavate- 
tubular, about 16 mm. long, the pedicels 8 mm. long; stamens 11-12, the filaments 
broad, compressed, about as wide as the anthers, abruptly contracted at the apex. 

We have seen no material of this species. 

Marcgravia guatemalensis Standl. Contr. U. S. Nat. Herb. 20: 
220. 1919. 

On trees in wet forest, 800-950 meters; Alta Verapaz (type from 
Finca Sepacuite", 0. F. Cook & R. F. Griggs 230). Costa Rica. 

A slender, woody vine, glabrous almost throughout; leaves on petioles 2-3 mm. 
long, narrowly lance-oblong, 9-15 cm. long, 2-4 cm. wide, narrowly long-acuminate, 
rounded or very obtuse at the base, thin when dried, the lateral nerves obscure, 
about 13 pairs, divergent at a rather wide angle; umbels about 15-flowered, pe- 
dunculate, the pedicels 2.5 cm. long, puberulent, the flowers inserted obliquely; 
bracts about 4, tubular-clavate, 20-23 mm. long and 3.5 mm. thick, the pedicels 
10-12 mm. long; sepals 1-1.5 mm. long, much broader than long, very broadly 
rounded at the apex; corolla ovoid, 8 mm. long, obtuse; stamens about 12; fruit 
depressed-globose, 1 cm. in diameter. 

Sterile plants, especially juvenile creeping ones, probably of this 
species, were observed in the wet forest above Tactic at a consider- 
ably higher elevation. So far as known at present, sterile or juvenile 
plants can not be referred to definite species and are not worth pre- 
serving in the herbarium. There are at hand several such collections 
from Zacapa and San Marcos, but it is impossible to guess at their 
identity. 

Marcgravia nepenthoides Seem. Journ. Bot. 8: 245. 1870; 
Hemsl. Biol. Centr. Am. Bot. 1: 90, t. 6. 1879. 

Wet mixed forest, 2,500 meters or less; reported from Alta Vera- 
paz; Izabal; Zacapa. British Honduras to Panama. 

A large, woody vine, sometimes 15 meters long or more, epiphytic or terres- 
trial; leaves on stout petioles 7 mm. long or shorter, chartaceous, oblong-lanceolate, 
10-18 cm. long, 2.5-4.5 cm. wide, long-acuminate, obtuse at the base, the nerves 
usually conspicuous beneath and reticulate; umbels 20-25-flowered, usually pend- 
ent, the stout pedicels 3-3.5 cm. long, verrucose, the flowers inserted obliquely; 
bracts several, almost 4 cm. long, galeiform, rounded at the apex, 1.5 cm. broad 
at the base, on long stout pedicels; corolla about 12 mm. long, somewhat conic. 



20 FIELDIANA: BOTANY, VOLUME 24 

Called cachimba in Honduras, in reference to the inverted (when 
growing) bracts, which suggest tobacco pipes. This plant doubtless 
is common along most of the Atlantic lowlands of Central America, 
but the vines usually are so high in the trees that it is impractical to 
obtain specimens, and herbarium material is unfortunately meager. 
One often finds fallen bracts of flowers on the ground, but not asso- 
ciated with leaves. The plants of this genus have attracted much 
attention from naturalists, particularly Thomas Belt, because of their 
curious inflorescences. The bracts or nectaries excrete a kind of 
nectar, which is much sought by birds. 

Marcgravia rectiflora Triana & Planch. Ann. Sci. Nat. IV. 17: 
364. 1862. 

On trees in wet forest, 2,000 meters or lower, usually most plenti- 
ful little above sea level; Izabal (Cerro San Gil); Huehuetenango 
(Sierra de los Cuchumatanes) . Honduras to Panama; West Indies; 
northwestern South America. 

A large, woody vine, glabrous throughout; leaves sessile, thick-coriaceous, 
oblong or lance-oblong, 6-14 cm. long, short-acuminate, obtuse and somewhat 
oblique at the base, the nerves obsolete, the costa prominent beneath; leaves of 
the sterile plants much smaller, oval or oblong, obtuse or rounded at the apex; 
umbels rather few-flowered, the stout pedicels 4-6 cm. long, bearing numerous 
large elevated lenticels; bracts cylindric-clavate, stipitate, about 2 cm. long; sepals 
rounded; flowers continuous with the axis of the pedicels; corolla whitish, about 
12 mm. long; fruit globose, deep red, 1-1.5 cm. in diameter. 

Marcgravia Schippii Standl. Carnegie Inst. Wash. Publ. 461: 
71. 1935. 

Type from Pete"n, in forest, Camp 32, British Honduras boundary, 
840 meters, W. A. Schipp 1273; also in Huehuetenango (Cerro Chi- 
blac, 1,200-2,000 meters). 

A glabrous, woody vine 25 meters long, the stems as much as 7.5 cm. in diam- 
eter; leaves sessile, rigid-coriaceous, oblong or lance-oblong, 7-8 cm. long, 2-3 cm. 
wide, narrowly obtuse, very oblique at the base, rounded on one side, acute on the 
other, the costa stout and prominent beneath, the lateral nerves obsolete on both 
surfaces; umbels about 10-flowered, the pedicels 5 cm. long, the axis of the flower 
continuous with that of the pedicel; bracts not seen; sepals broader than long, 
4-6 mm. long, truncate or broadly rounded at the apex; corolla oblong-ovoid, 
almost 1 cm. long, broadly rounded at the apex; ovary globose, abruptly con- 
tracted into the short style. The collector states that the flowers are cream- 
colored, the fruit red. 

RUYSGHIA Jacquin 

Epiphytic shrubs, usually more or less scandent, sometimes much elongate; 
leaves on very short, thick petioles, spirally inserted, coriaceous; inflorescences 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 21 

terminal, racemose, many-flowered; bracts inserted at the middle or above the 
middle of the pedicel, sessile, globose, semiglobose, or spoon-shaped, not hollow; 
sepals 5; petals 5, united at the base; stamens 5, alternate with the petals and 
united with them at the base; anthers globose; ovary 2-celled, the style evident or 
obsolete; radicle thicker than the cotyledons and twice as long. 

Species 7-8 or perhaps more numerous, in tropical America. 
Two or three others are found in southern Central America. 

Ruyschia enervia Lundell, Phytologia 1 : 245. 1937 (type from 
British Honduras, Lundell 6308). R. longistylis Standl. & Steyerm. 
Field Mus. Bot. 23: 175. 1944 (type from Guatemala, Steyermark 
49545). 

In wet forest not far above sea level; Huehuetenango; Alta Vera- 
paz. British Honduras. 

A glabrous epiphytic woody vine, the branches stout, subterete, ochraceous; 
leaves short-petiolate, thick-coriaceous, the thick petiole 5 mm. long; leaf blades 
oblanceolate-oblong, 10-12 cm. long, 3-3.5 cm. wide, broadest above the middle, 
obtuse and mucronate-apiculate, gradually narrowed to the obtuse or subacute 
base, yellowish-green above when dry, slightly paler beneath, the lateral nerves 
obsolete; inflorescence in fruit 11 cm. long, rather remotely many-flowered, the 
rachis almost 3 mm. thick, the pedicels thick, 2.5-4 mm. long, spreading or slightly 
ascending; bracts inserted at the base of the calyx, semiglobose or obovate, 4 mm. 
long, obtuse, narrowed to the base and substipitate, very thick; sepals strongly 
unequal, suborbicular, the inner ones 3 mm. long, thick, rounded at the apex, 
appressed; immature fruit subglobose, 4 mm. long, the thick columnar style 
2.5 mm. long. 

SOUROUBEA Aublet 

Shrubs or trees, terrestrial or epiphytic, sometimes scandent, often with numer- 
ous long pendent aerial roots; leaves short-petiolate, coriaceous, lustrous, with few 
or numerous glands on the lower surface; inflorescence racemose, usually elongate; 
bracts deciduous, the limb modified into a cylindric-clavate spur, this often deeply 
2-parted at the base, or the limb sometimes sessile and spur-like or hat-like, in- 
serted at or below the apex of the pedicel, the limb usually hollow; sepals 5, strongly 
imbricate; petals 5 or 3, connate below, reflexed in anthesis, circumscissile at the 
base and deciduous; stamens 5 or 3, the filaments complanate, adnate to the bases 
of the petals; anthers ovate, dorsifixed above the base; ovary ovoid, usually 5- 
celled, the stigma sessile, radiate; ovules numerous, ascending or subhorizontal; 
fruit fleshy-coriaceous, depressed-globose, indehiscent or irregularly ruptured; 
seeds 3-6 or fewer in each cell, the testa areolate. 

About 15 species in tropical America, from Mexico southward. 
One additional species occurs in southern Central America. The 
genus has been monographed by A. C. de Roon but the monograph 
has not yet been published. The annotations by the monographer 
are not necessarily followed here. 



22 FIELDIANA: BOTANY, VOLUME 24 

Lobes of the bracts about as long as the saccate portion; bracts about 25 mm. long. 

S. guianensis. 

Lobes of the bracts inconspicuous or none; bracts to about 10 mm. long. 

Inflorescence minutely puberulent; leaves not vernicose S. exauriculata. 

Inflorescence glabrous; leaves vernicose or not. 

Leaves vernicose, shining, mostly less than 10 cm. long S. triandra. 

Leaves not vernicose, dull, mostly more than 10 cm. long S. Gilgii. 

Souroubea exauriculata Delpino, Att. Soc. Ital. Sci. Nat. Mi- 
lan 12: 180. 1869; S. belizensis Lundell, Field & Lab. 13: 8. 1945 (type 
from British Honduras, Gentle 4648); S. puberula Standl. & Steyerm. 
Field Mus. Bot. 23: 175. 1944 (type from Guatemala, Steyermark 
46862). 

In wet forest or wet open slopes in the mountains. Suchitepe"- 
quez; Huehuetenango. Mexico; British Honduras; Honduras and 
Nicaragua. 

A scandent epiphytic shrub or occasionally a small tree, glabrous except the 
puberulent inflorescence, the branches stout, brownish or ochraceous, obtusely 
subangulate; leaves short-petiolate, subcoriaceous, oblanceolate-oblong, 5-12 cm. 
long and 2-4.5 cm. broad, obtuse to very obtuse at the apex, mucronate-apiculate, 
broadest above the middle, gradually narrowed to acute or attenuate at the base, 
lateral nerves slender and inconspicuous; petioles short, 5-9 mm. long; inflores- 
cence terminal, sessile or short-pedunculate, dense, many-flowered, 7-20 cm. long, 
minutely puberulent in all parts, pedicels 6-10 mm. long, stout; bracts inserted 
at the base of the calyx, about 10 mm. long; sepals closely imbricated, orbicular, 
2.5-5 mm. long, rounded at the apex. 

Perhaps the commonest species of the mountains of northern 
Central America. Souroubea mexicana Baill. is another synonym. 

Souroubea Gilgii Al. Richter ex Gilg & Werderm. in Engler & 
Prantl, Nat. Pflanzenfam. ed. 2, 21: 102. 1925, nomen subnudum. 

Wet forest area, mostly at low elevations. Izabal. Honduras; 
Nicaragua; Costa Rica; Panama. 

Weak vines or shrubs; leaves oblanceolate to oblong-obovate, obtuse to some- 
what acute, apiculate, cuneate or attenuate to the base, 10-18 cm. long and 
4-7 cm. broad, subcoriaceous; petioles thick and short, 1-3 mm. long; inflores- 
cence to 25 cm. long, spicate, the rachis thickened and fleshy; sepals suborbicular, 
2-3 mm. long; corolla sympetalous, 4-5-lobed, 5-8 mm. long; bract 10-14 mm. 
long, biauriculate at the apex, the auricles 2-4 mm. long, the sac subcylindric, 
scrotiform at the base, 8-10 mm. long. 

The species seems not to have been properly published but it 
appears to be distinct. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 23 

Souroubea guianensis Aubl. PL Guian. 244, t. 97. 1775. Ruys- 
chia guianensis Sw. Fl. Ind. Occ. 1: 504. 1797. Souroubea sympetala 
Gilg in Engler, Bot. Jahrb. 25, Beibl. 60: 32. 1898. 

Dense, wet, mixed forests at sea level or but little above. Izabal. 
British Honduras to Panama; northern South America. 

A small to large epiphytic vine with thick branches and usually numerous long 
flexible aerial roots; leaves obovate to oblong-obovate, 5-15 cm. long, rounded to 
subacute at the apex, obtuse or acute at the base, more or less coriaceous, fleshy 
when fresh, the lateral nerves inconspicuous, bearing few to numerous large glands 
on the lower surface, petiole short; racemes usually elongate and many-flowered, 
the pedicels remote, stout, 1.5-3.5 cm. long; bracts red, the spur hollow, slender, 
extending below the point of attachment into two slender auricles of equal length, 
the whole bract to 2.5 cm. long; sepals ovate-oblong, yellow, reflexed in anthesis; 
petals yellow; stamens 5, half as long as the petals; fruit subglobose; seeds usually 
few, 4-5 mm. long, black. 

The Central American material is apparently considered by the 
monographer, de Roon, to be a distinct species and has been anno- 
tated under the name of S. sympetala Gilg. I am unable to distin- 
guish the species from the older S. guianensis. 

Souroubea triandra Lundell, Phytologia 1 : 244. 1937 (type from 
British Honduras, Lundell 6492); S. micrantha Standl. & Steyerm. 
Field Mus. Bot. 23: 62. 1944 (type from Guatemala, Standley 72793). 

Wet forests at relatively low elevations. Izabal; Alta Verapaz. 
British Honduras; Costa Rica. 

A glabrous epiphytic shrub; leaves oblanceolate to obovate-elliptic, 5-12 cm. 
long and 2-6 cm. broad, rounded or very obtuse at the apex, attenuate or acute at 
the base, subcoriaceous, vernicose or lustrous, petioles 3-8 mm. long; racemes 5- 
16 cm. long, 1.5-2 cm. broad, many-flowered, the pedicels stout, divaricate or 
somewhat reflexed, 5-9 mm. long; bracts tubular-cylindric, 8-10 mm. long, 
inserted at the base of the calyx, short stipitate; sepals orbicular, 1.8 mm. long or 
less; corolla about 4.5 mm. long, 3-4-lobate to the middle; irregularly cleft in 
anthesis, the lobes reflexed, obtuse; stamens 3-4; ovary 3-4-celled. 

The monographer, de Roon, has applied an unpublished specific 
combination to this species which, when published, may replace the 
name S. triandra. 

QUIINACEAE 

Trees or shrubs, sometimes scandent; leaves opposite or verticillate, simple 
and entire or dentate or sometimes pinnate, penninerved, finely transverse-lineo- 
late between the nerves; stipules usually geminate, narrow, stiff, persistent; flowers 
small, polygamous, in terminal or axillary panicles; sepals 4-5, imbricate; petals 
usually 4-5, sometimes 6-8, alternate with the sepals, imbricate or contorted in 
bud; stamens 15-30, free or connate at the base, the filaments filiform; anthers 



24 FIELDIANA: BOTANY, VOLUME 24 

small, globose-didymous, the cells dehiscent by slits; ovary 2-3-celled or 7-11- 
celled; styles distinct, linear, stigmatose at the apex; ovules 2 in each cell, ascend- 
ing; fruit baccate, finally dehiscent, often by abortion 1-celled, 1-3-seeded, the 
endocarp fibrous; seeds ovoid or globose, tomentose. 

Genera about 3, all tropical American. One other, Lacunaria, is 
represented in Panama. 

QUIINA Aublet 

Trees or shrubs, sometimes scandent; leaves opposite, entire or dentate or 
sometimes pinnate, short-petiolate, coriaceous; stipules foliaceous, erect; flowers 
very small, short-pedicellate, in clusters of 2-3 in the axils of bracts, arranged 
in very small, few-flowered panicles; sepals 4-5; petals 4-5, rarely 6-8; stamens 
15-30; ovary 2-3-celled; styles 2-3, linear; fruit an almost dry berry, usually 
1-celled and 1-3-seeded. 

About 15 species, chiefly in the Guianas and Brazil, with only the 
following species in North America. 

Quiina Schippii Standl. Field Mus. Bot. 8: 26. 1930. 

British Honduras, in wet forest little above sea level, the type 
from Middlesex, Schipp 238; to be expected in Pete*n and Izabal; 
Honduras (Lake Yojoa, 600 meters). Figure 6. 

An almost glabrous tree 6-18 meters high, the young branchlets appressed- 
pilose with fulvous hairs; stipules linear or linear-subulate, 7-15 mm. long; leaves 
opposite, on stout petioles 3-5 mm. long, oblong-lanceolate, 8-20 cm. long, 2-7 cm. 
wide, narrowly long-acuminate, gradually attenuate to the base, green and lus- 
trous above, paler beneath, the lateral nerves about 9-13 pairs, ascending, curved; 
racemes solitary or fasciculate, 1-4.5 cm. long, simple, with few-many flowers, the 
bracts ovate, acute, ciliate, the rachis densely hirtellous, the pedicels 1.5-3 mm. 
long, glabrous; sepals 4, oval or rounded, 1.5-1.8 mm. long, rounded at the apex, 
ciliate; petals 4, rounded, 2.5 mm. long, glabrous, white, ciliolate; stamens shorter 
than the petals; fruit oblong, 8 mm. long, bright red, obtuse or subtruncate at 
the apex. 

The name "pigeon plum" is reported from British Honduras. 
THEACEAE. Tea Family 

Trees or shrubs with deciduous or usually persistent leaves; leaves alternate, 
simple, entire or serrate, without stipules; flowers usually perfect, regular, small or 
large, axillary or subterminal, generally solitary; sepals 5-7, rarely more numerous, 
imbricate, usually persistent; petals 5, rarely 4 or more, distinct or connate at the 
base; stamens very numerous, rarely only 5, 10, or 15, free or united at the base, 
often adnate to the petals; ovary superior or rarely inferior, 2-10-celled, the cells 
1-many-ovulate; styles as many as the carpels or connate into one; fruit a loculici- 
dal capsule, or indehiscent and dry or drupaceous; seeds 1-many; endosperm none 
or scant; embryo usually curved. 




FIG. 6. Quiina Schippii. A, Branch showing leaves and inflorescence. B, Flower; 
X 10. C, Portion of branch, showing fruits. 



25 



26 FIELDIANA: BOTANY, VOLUME 24 

About 18 genera, with most of their species in tropical or sub- 
tropical regions of both hemispheres. One other genus, Pelliciera, 
is represented in southern Central America. 

Anthers versatile; fruit capsular. 

Seeds winged; native trees Laplacea. 

Seeds not winged; cultivated shrubs Camellia. 

Anthers basifixed; fruit indehiscent. 

Leaves, at least usually, densely tomentose or sericeous beneath; anthers gla- 
brous Freziera. 

Leaves glabrous or glabrate beneath, at least in age. 

Ovary inferior; leaves crenate-serrate Symplococarpon. 

Ovary superior. 

Anthers pilose; leaves (in Guatemalan species) serrulate; bractlets minute. 

Cleyera. 

Anthers glabrous; leaves entire or obscurely serrulate; bractlets large and 
conspicuous Ternstroemia. 

CAMELLIA L. 

Trees or shrubs; leaves persistent, coriaceous or membranaceous, serrate; 
flowers often large and showy, axillary, solitary or clustered, sessile or short- 
pedicellate; sepals 5-6, unequal; petals short-coherent at the base, strongly im- 
bricate; stamens numerous, many-seriate, united at the base or almost to the 
apex, the inner ones free, the anthers versatile; ovary 3-5-celled, the styles free 
almost to the base, or more or less united; ovules 4-5 in each cell, pendulous; 
capsule ligneous, loculicidally dehiscent; seeds usually solitary in each cell; endo- 
sperm none. 

About 45 species, in tropical and subtropical Asia. 

Camellia japonica L. Sp. PI. 698. 1753. Camelia. 

Native of China and Japan; planted widely for ornament in 
warmer regions of the earth; grown commonly for ornament in 
Guatemala, chiefly in the mountains, at lower elevations planted in 
the ground, in the higher mountains often grown in pots. 

A shrub or small tree, glabrous throughout or nearly so, with leathery, deep 
green, ovate to elliptic leaves 5-10 cm. long. The large flowers are primarily red, 
but they vary to white, and there are many varieties. 

The camellia seems to thrive in Guatemala at middle elevations, 
but at the highest elevations, as in San Marcos, it is sometimes in- 
jured by the cold, and it does not grow well out of doors at Quezalte- 
nango, where the flowers bring a price that is very high for Guatemala. 

Camellia sinensis (L.) Kuntze, Act. Hort. Petrop. 10: 195. 1887. 
Thea sinensis L. Sp. PI. 515. 1753. Camellia Thea Link, PI. Hort. 
Berol. 2: 73. 1822. Te. Tea. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 27 

Native of China and India; planted occasionally in Guatemalan 
gardens for ornament or as a curiosity; formerly planted on a com- 
mercial scale at Finca Chirripec near Coban, Alta Verapaz. 

The tea of commerce is obtained from the dried, young and tender 
leaves of this shrub. Tea from the Orient is imported normally into 
Guatemala, principally for use by foreign residents of the country. 
It is not a common beverage among native Central American people. 
The tea plantation at Finca Chirripec was a fairly large one, and is 
said to have furnished the greater part of the tea used in Guatemala. 
The plant for drying and aging the leaves was a small building, and 
it is evident that the amount produced was not very great. It was 
marketed in two distinct grades, in cartons weighing from one ounce 
upward. The Guatemalan tea gardens were very neat and pretty in 
appearance and looked just like the pictures of the tea gardens of 
India and Ceylon, the terrain and the climate being much the same. 
For the successful production of tea, a mild cool climate with per- 
manently moist air is needed, and these conditions seem to have 
been found at Chirripec. So far as we know, this was the only large 
commercial tea plantation in Central America. The labor was per- 
formed by Quecchi Indians of the vicinity, principally by women. 
Little or no tea is harvested in Guatemala now because of the high 
cost of harvesting. 

CLEYERA Thunberg 

Reference: Clarence E. Kobuski, Studies in the Theaceae, VII. 
The American species of the genus Cleyera, Journ. Arn. Arb. 22: 
395-416. 1941. 

Trees or shrubs; leaves persistent or deciduous, serrulate (in Guatemalan spe- 
cies) or entire; flowers perfect, axillary or borne at leafless nodes, solitary or fascic- 
ulate, the pedicels usually about 1 cm. long, thickened slightly toward the apex, 
2-bracteolate, the bracteoles small to minute, subopposite or alternate near the 
apex of the pedicel; sepals 5, imbricated, unequal, the outer ones smallest, ciliolate 
usually; petals 5, imbricate, connate at the base; stamens about 25, the anther 
setose, 2-celled, opening by longitudinal slits; ovary usually glabrous, 2-3-celled, 
many-ovulate; styles elongate, 3-4-fid; fruit indehiscent, baccate, globose or ovoid, 
many-seeded; endosperm scant, the embryo curved. 

About ten species are known from mountain regions of tropical 
America, and a few others occur in Asia. No other species is known 
in Central America. 

Cleyera theaeoides (Sw.) Choisy, Mem. Soc. Phys. Hist. Nat. 
Geneve 14, 1: 110. 1855; L. Wms. Fieldiana, Bot. 29: 353. 1961. 



28 FIELDIANA: BOTANY, VOLUME 24 

Eurya theaeoides (Sw.) Blume, Mus. Bot. Ludg.-Bat. 2: 105. 1856. 
Cleyera Matudai Kobuski, Journ. Arn. Arb. 22: 403. 1941 (type from 
Mexico, Matuda 2560). C. revoluta Kobuski, I.e. 405 (type from Gua- 
temala, Steyermark 36208). C. tacanensis Kobuski, I.e. 406 (type 
from Guatemala, Steyermark 36052). C. Skutchii Kobuski, I.e. 405 
(type from Guatemala, Skutch 545). Tabojilla; cormen; baratillo; 
fruta de cabro. 

Often in open cloud forests or in drier pine-oak forests, 1300 to 
3000 meters. El Progreso; Zacapa; Alta Verapaz; Baja Verapaz; 
San Marcos; Quiche"; Chiquimula; Jalapa; Guatemala; Suchitepe"- 
quez; Chimaltenango; Quezaltenango. Mexico (Chiapas); El Sal- 
vador; Honduras; Nicaragua; Costa Rica; Panama. Figure 7. 

A rather large shrub or, when well grown, an attractive tree to 20 m. tall; 
branchlets pubescent but usually becoming glabrous; leaves short-petiolate, ob- 
long-elliptic to obovate or broadly oval, coriaceous, glabrous or slightly pubescent 
on the petiole and on the back of the leaf, crenulate-dentate, occasionally obscurely 
so, cuneate to the base, apex acuminate, (usually) obtuse, blade 2-15 cm. long 
and 1-6 cm. broad, lateral nerves 10-12 pairs; flowers solitary or fascicled from 
the twigs or the leaf axils; pedicels 5-20 mm. long, glabrous or puberulent, with 
two subopposite deciduous bracts near the flower; sepals 5, unequal, rounded, the 
outer ones pubescent, puberulent, or glabrous, sometimes verriculate, usually cilio- 
late, 2-4 mm. long; petals usually 5, white or greenish-yellow, sometimes emargi- 
nate, 5-8 mm. long; stamens about 25, the anther sparsely pubescent or glabrous; 
ovary glabrous or puberulent, 3-celled; styles 3, free from about the middle to 
almost the base. 

The Indian name "Yxinche" is reported by Garcia Salas for Chi- 
maltenango. The tree is called coyolillo or cacao in Honduras, both 
names better known for other plants. Most Guatemalan specimens 
of this species have been reported as Eurya theoides or as Cleyera 
Skutchii. The leaves on flowering growths are usually smaller than 
those on sterile growths or those of a previous season. Kobuski has 
divided this species into several which it seems impossible to sepa- 
rate. The characters relied upon for separation are too inconsistent 
for use. 

FREZIERA Swartz 

Reference: Clarence E. Kobuski, Studies in the Theaceae, VIII. 
A synopsis of the genus Freziera, Journ. Arnold Arb. 22: 457^496. 
1941. 

Trees or large shrubs; leaves mostly coriaceous, lightly serrulate or subentire, 
petiolate; flowers small, dioecious, fasciculate in the leaf axils or on short flowering 
branchlets; bractlets 2, opposite, at the apex of the pedicel; sepals 5, imbricate, 




B 



FIG. 7. Cleyera theaeoides. A, Portion of flowering branch. B, Flower; X 4. 
C, Fruit; X 4. 



29 



30 FIELDIANA: BOTANY, VOLUME 24 

persistent; petals 5, imbricate, unequal, usually free to the base; stamens 15-30 
in the staminate flowers, 1-seriate, the filaments adnate to the bases of the petals; 
anthers linear or ovate-elliptic, glabrous, usually dehiscent to the base; ovary 
glabrous, 3-5-celled, attenuate into an entire style, the ovules numerous; staminodia 
of the pistillate flower fewer than the stamens; fruit indehiscent, conic-globose; 
seeds 10-50 in each cell, small. 

Species about 34, in tropical America, mostly in mountain re- 
gions. Two or three other species are known from southern Central 
America. 

Leaves covered beneath with a dense ferruginous tomentum of matted and en- 
tangled hairs F. guatemalensis. 

Leaves pilose beneath with spreading or appressed, not matted, mostly straight 

hairs. 
Leaves densely and conspicuously papillose along and near the costa on the 

upper surface F. macrophylla. 

Leaves smooth on the upper surface, not at all papillose F. Grisebachii. 



Freziera Friedrichsthaliana (Szysz.) Kobuski (Eurya Friedrichs- 
thaliana Szysz.) was reported originally as a native of Guatemala, 
and based upon Friedrichsthal 996. The plants of this collection, 
while all labeled as coming from Guatemala, were collected in large 
part in other Central American countries. The species has been 
collected twice in Nicaragua, and not recently in Guatemala. Since 
the collector is known to have collected plants in Nicaragua, it is 
best to delete this species from the list of Guatemalan plants, at 
least for the present. 

Freziera Grisebachii Krug & Urban, Bot. Jahrb. 21: 542. 1896. 
Eurya lancifolia Standl. Field Mus. Bot. 8: 317. 1931 (type from 
British Honduras, Schipp 455). F. lancifolia Kobuski, Ann. Mo. 
Bot. Gard. 25: 354. 1938. 

Moist or wet, mixed forest, 120-825 meters; Pete'n (Camp 36, 
British Honduras boundary, W. A. Schipp S710). British Hon- 
duras. 

A tree of 15-18 meters, the trunk 30 cm. in diameter, the branches thinly 
appressed-pilose with fulvous hairs, in age glabrate; leaves on petioles 1-1.5 cm. 
long, thin, lance-oblong, 10-14 cm. long, 3-4.5 cm. wide, acuminate or long- 
acuminate, rounded at the base, serrulate, glabrous on the upper surface except 
along the costa, slightly paler beneath, pilose with usually closely appressed hairs, 
the lateral nerves arcuate; flowers axillary or at defoliated nodes, fasciculate or 
on very short peduncles, the pedicels in fruit rather slender and 3-5 mm. long; 
sepals suborbicular, 3-3.5 mm. long, densely sericeous; fruit subglobose, 7 mm. 
long, 3-celled, glabrous. 



STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 31 

Freziera guatemalensis (Donn.-Sm.) Kobuski, Ann. Mo. Bot. 
Gard. 25: 354. 1938. Eurya guatemalensis Donn.-Sm. Bot. Gaz. 46: 
109. 1908. Huel Colorado (Huehuetenango) ; alajix (Coban, Quecchi). 

Mostly in moist or wet pine forest, 900-2000 meters; Alta Verapaz 
(type from Coban, Tuerckheim II. 1824); Guatemala; Huehuete- 
nango. Mountains of Honduras. 

A small or large tree sometimes 12 meters high, with long, rather slender, 
rufous-torn entose branches; leaves on short stout petioles, lance-oblong, 10-20 cm. 
long, 3-6 cm. wide, acuminate or long-acuminate, rounded or obtuse at the base, 
entire or serrulate, green above, glabrous or practically so, covered beneath with 
a dense persistent tomentum of ferruginous matted hairs, coriaceous or chartaceous, 
the lateral nerves in numerous pairs, almost straight; flowers axillary or at de- 
foliated nodes, on very short, few-flowered, stout peduncles, the pedicels 5-7 
and 4 mm. long or less, thick; flowers 8 mm. long; sepals suborbicular, ferruginous- 
tomentose; petals 7 mm. long; stamens glabrous. 

The wood is used commonly about Coban for firewood. The 
wood in this genus is dull brownish or reddish brown, hard and 
heavy or moderately so. Kobuski refers to F. guatemalensis collec- 
tions which we find better referable to F . macrophylla. 



Freziera Karsteniana (Szysz.) Kobuski, Ann. Mo. Bot. Gard. 
25: 354. 1938. Eurya Karsteniana Szysz. in Engl. & Prantl, Nat. 
Pflanzenfam. II. 6: 190. 1893. This was described originally as 
from Guatemala, but it has been found that the species is actually 
native in Colombia. 

Freziera macrophylla Tulasne, Ann. Sci. Nat. Bot. III. 8: 330. 
1842. Durazno de monte. 

Moist or wet, mixed, mountain forest, 1,400-2,500 meters; Que- 
zaltenango; San Marcos. Mexico (Oaxaca). 

A small tree, sometimes 9 meters high, the branches densely fulvous-pilose 
with spreading or subappressed hairs; leaves on short thick petioles, lance-oblong, 
12-25 cm. long, 4-6.5 cm. wide, acuminate or narrowly long-acuminate, obtuse 
or rounded at the base, serrulate, green above (when dry yellow-green), glabrous 
except along the costa, conspicuously and densely verrucose near the costa, fulvous- 
pilose beneath with soft, spreading or often appressed hairs, the lateral nerves 
20-25 pairs, almost straight or slightly arcuate; flowers borne in the leaf axils 
and at defoliated nodes, subsessile or on pedicels 2-4 mm. long; sepals suborbicular, 
coriaceous, 3-4 mm. long, the outer ones densely pubescent; petals glabrous, creamy 
white, twice as long as the sepals; stamens about 15; ovary glabrous; fruit globose, 
5-celled. 

The specimens from Guatemala, in part referred by Kobuski 
to F. guatemalensis, seem to agree perfectly with authentic material 
of F. macrophylla, and to be certainly different from F. guatemalensis. 



32 FIELDIANA: BOTANY, VOLUME 24 

LAPLACEA HBK. 

Reference: Clarence E. Kobuski, Studies in the Theaceae, XX. 
Notes on the South and Central American Species of Laplacea, 
Journ. Am. Arb. 31: 405^29. 1950. 

Shrubs or large trees, glabrous or pubescent; leaves coriaceous, more or less 
dentate; flowers pedicellate, axillary, sometimes large and showy; sepals 5, un- 
equal, the calyx bracteolate at the base; petals 5, slightly coherent, strongly im- 
bricate; stamens numerous, the outer ones or all of them more or less connate 
or in 5 groups, adherent to the base of the petals, the anthers versatile; ovary 
5-10-celled, the styles short and divergent or the stigmas sessile and distinct; 
ovules 4-8 in each cell, pendulous; fruit capsular, often ligneous, broadly oblong, 
loculicidally dehiscent, the columella persistent; seeds compressed or flat, produced 
at the apex into an oblong wing; endosperm none, the embryo straight, the cotyle- 
dons oblong, flat, the radicle short, superior. 

About 30 species, in tropical America and the East Indies. Two 
other species occur in southern Central America. 

Laplacea coriacea L. Wms. Fieldiana, Bot. 29: 354. 1961 (type 
from Guatemala, Steyermark 42880). 

Probably cold cloud forest. Endemic. Zacapa. 

Shrubs or trees of unknown size; branchlets many, terete, glabrous. Leaves 
toward the apex of the branches and branchlets, stiff-coriaceous, completely gla- 
brous, symmetrical or slightly asymmetrical, broadly oval to broadly obovate, ob- 
tuse, the margins entire or slightly undulate, the blade 4-8 cm. long and 2.5-5 cm. 
broad, often decurrent into the petiole; the nerves 7-12 pairs, nearly plain above, 
slightly canaliculate beneath, the mid-rib plain above, elevated beneath; the petiole 
to 1.5 cm. long, subglandular, sometimes winged; inflorescences axillary, flowers sol- 
itary; peduncle terete, 1-2.5 cm. long; sepals 5, imbricated, about 3 mm. long and 
twice as broad, fleshy and coriaceous, ciliolate, otherwise glabrous; petals unknown; 
stamens unknown; ovary elliptic-ovoid, densely sericeous; stigma stellate, 5-lobed; 
capsule ovoid, about 1.5 cm. long and 0.8 cm. thick, 5-loculate. 

This species has been confused by both Standley and Kobuski 
with Laplacea grandis and into it have been placed specimens of 
that species, the specimen on which this species is based, and several 
specimens belonging in the Sabiaceae and in another family. The 
very broad, thick, obtuse leaves of L. coriacea are characteristic. 

SYMPLOCOCARPON Airy-Shaw 

Reference: Clarence E. Kobuski, Studies in the Theaceae, VI. The 
genus Symplococarpon Airy-Shaw, Journ. Arnold Arb. 22: 188-196. 
1941. 

Trees, glabrous or pubescent; leaves alternate, entire or serrate, more or less 
coriaceous; flowers perfect, axillary, fasciculate, the pedicels minutely bracteate 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 33 

at the base and with 2 persistent bractlets, opposite or nearly so, at the juncture 
of the pedicel and hypanthium; sepals 5, imbricate, persistent; petals 5, alternate 
with the sepals, imbricate, slightly connate at the base; stamens 25-40, uniseriate, 
the filaments connate at the base and adnate to the base of the corolla; anthers 
oblong-elliptic, narrowed at the apex into a subulate mucro; ovary inferior, 2-celled; 
ovules usually solitary in each cell, sometimes more numerous; styles 2, rarely 3, 
free; fruit indehiscent, subglobose, crowned by the persistent calyx and styles, 
2-celled, usually 2-seeded. 

Seven species, ranging from central Mexico to Panama. Three 
are known in Costa Rica and Panama. 

Symplococarpon lucidum Lundell, Bull. Torr. Bot. Club 69: 
394. 1942. Paso, macho. Wild raisin male. 

Known only from British Honduras, type Gentle 3248; Conservator 
of Forests 85. 

A small tree, the bark dark green, glabrous except for the whitish-strigose buds, 
the branchlets slender, reddish; leaves on rather stout petioles 2.5-4.5 mm. long, 
membranaceous or chartaceous, elliptic-oblong to oblanceolate or lanceolate, 4-10 
cm. long, 2-4.5 cm. wide, yellowish green and usually lustrous above, obtuse- 
acuminate, cuneate at the base, crenate-serrate, the lateral nerves 6-9 pairs, the 
veins reticulate; flowers in clusters of 9 or fewer, the pedicels slender, in bud 1 cm. 
long, usually recurved, the bractlets ovate, ciliate, 9.6-0.8 mm. long; hypanthium 
glabrous; calyx lobes depressed-orbicular, ciliate; stamens about 30; free portion 
of the ovary sparsely hirsute; styles 2; cells of the ovary 2-3-0 vulate. 

It is questionable whether or not this species is distinct from 
S. Brenesii Kobuski from southern Central America. The wood is 
reported to be pink. 

TERNSTROEMIA Mutis 

Reference: Clarence E. Kobuski, Studies in the Theaceae, XIII. 
Notes on the Mexican and Central American species of Ternstroemia, 
Journ. Arnold Arb. 23: 464-478. 1942. 

Glabrous trees or shrubs; leaves spirally disposed, often congested or verti ciliate 
at the ends of the branches, usually coriaceous, mostly entire or nearly so; flowers 
axillary, solitary, bracteolate; bractlets generally 2, opposite or subopposite, at 
the base of the calyx or one of them somewhat distant; flowers perfect; sepals 
usually 5, imbricate, persistent, coriaceous; petals mostly 5, free or more or less 
connate; stamens 25-300, usually 2-seriate, the filaments connate, the outer ones 
adnate to the bases of the petals; anthers mostly longer than the filaments, oblong 
or linear, the connective usually produced into an appendage; ovary 2-3-celled, 
rarely 1-celled, or falsely 4-6-celled; ovules 2-20 in each cell, rarely solitary, 
pendulous from the apex of the placenta; style 1, entire or rarely 2-3-parted; fruit 
indehiscent, coriaceous; seeds few, the testa usually smooth. 



34 FIELDIANA: BOTANY, VOLUME 24 

A tropical genus, with 85 species or more, mostly in tropical 
America and Asia. Two or perhaps three other species are known 
from Central America. 

Ternstroemia Tepezapote Schlect. & Cham. Linnaea 6: 420. 
1831. T. Seleriana Loes. Bull. Herb. Boiss. II. 3: 213. 1903 (type 
from Chacula, Huehuetenango, Seler 3099). T. impressa Lundell, 
Bull. Torr. Bot. Club 66: 599. 1939 (type from Chiquihuite, Volcan 
de Tacana, Chiapas, E. Matuda 2814). Taonabo sphaerocarpa Rose, 
Contr. U. S. Nat. Herb. 8: 322. 1905. Ternstroemia sphaerocarpa 
Melchior, Nat. Pflanzenfam. ed. 2. 21 : 142. 1935. Baratillo (Jalapa) ; 
trencillo (Chiquimula) ; chucul (Huehuetenango); hualicuc (Huehue- 
tenango); uixlilil-caax (Pete"n, Maya, fide Lundell); panool, chique, 
roble (Pete"n, fide Lundell). 

Moist or wet thickets or mixed forest or pine forest, sometimes on 
limestone, 3,150 meters or lower; Pete'n; Alta Verapaz; El Progreso; 
Izabal; Zacapa; Chiquimula; Jalapa; Santa Rosa; Guatemala; Suchi- 
tepe"quez; Huehuetenango; Chimaltenango; Quiche 1 ; Solola. Southern 
Mexico; British Honduras; Honduras; El Salvador. Figure 8. 

A shrub or tree, sometimes 15 meters high, with a trunk 15 cm. or more in 
diameter; leaves oblong-obovate or oblong-oblanceolate, 7-13 cm. long, mostly 
3-4 cm. wide, obtuse or rounded at the apex or sometimes subacute, cuneate or 
attenuate at the base, entire or obscurely crenulate, the nerves inconspicuous on 
both surfaces, sometimes impressed above; pedicels 1.5-2.5 cm. long, the bractlets 
2, opposite, unequal, broadly ovate or suborbicular, 2-3 mm. long; sepals unequal, 
the outer ones suborbicular, about 8 mm. long, glandular-denticulate, often con- 
spicuously so, or sometimes entire, petals pinkish white, lanceolate or ovate, 8 mm. 
long, acute, united half their length; stamens about 50 and 2-seriate; ovary 2-celled, 
each cell 4-5-ovulate, the style 6-7 mm. long, the stigma punctiform; fruit conic 
or ovoid-conic, 1-2 cm. long, 1-1.5 cm. broad at the base. 

Called "river craboo" in British Honduras; "trompillo" (El Salva- 
dor). Kobuski recognizes as a distinct species Ternstroemia impressa, 
principally because the nerves are slightly impressed on the upper 
surface. To it he refers a specimen from Soloma, Huehuetenango, 
and another from Jalapa could be placed with it. This character, 
however, appears to be a matter of degree, and the species appears 
to be a quite untenable one, the more so when he reduces to synonymy 
under T. Tepezapote four species described from Mexico that probably 
have quite as good claims to specific status. Several of the Guate- 
malan specimens could be referred to T. megaloptycha Kobuski 
(Journ. Arnold Arb. 23: 474. 1942), which was based upon numerous 
collections from Honduras. It is keyed by Kobuski as having larger 
flowers than T. Tepezapote, but there appears to be every gradation 




FIG. 8. Ternstroemia Tepezapote. A, Flowering branch from tree; X K- 
B, Fruits; X J^. C, Fruit; X 1^. D, Flower, showing bractlets and calyx. 
E, Flower; X 4. 



35 



36 FIELDIANA: BOTANY, VOLUME 24 

in this respect, and it should be referred to synonymy under the 
latter. Apparently there is nothing besides geographic range to 
separate the two. This tree is remarkable for its wide altitudinal 
distribution in Guatemala, although in this respect it is equalled 
by many other trees. In Huehuetenango it is reported that a decoc- 
tion of the bark is employed as one of the numerous "remedies" for 
snake bites. The specific name Tepezapote is one of the Nahuatl 
names given to this tree in Mexico. In this genus the heartwood 
exhibits various shades of brown; luster low; of medium density but 
hard and strong; grain variable; not difficult to work and finishes 
smoothly; durability doubtful. So far as known, no use is made 
of the wood in Guatemala, unless for firewood. 

GUTTIFERAE 

Reference: Juliano Vesque in DC. Monog. Phan. 8: 1-669. 1893. 

Trees, shrubs, or often herbs, terrestrial or epiphytic, dioecious, polygamous, 
or perfect, sometimes with aerial roots, the sap frequently yellow and resinous, 
the leaves and petals sometimes with black or transparent dots or lines; leaves 
mostly opposite and decussate, penninerved, simple, entire, without stipules, 
sometimes with a marginate pit in the axillary part of the petiole; flowers gen- 
erally white or yellow, sometimes pink, in chiefly terminal inflorescences, some- 
times solitary, regular or nearly so, the pedicels often bracteolate, the bractlets 
usually resembling sepals; sepals variable in number, sometimes unequal; petals 
variable in number, free or nearly so; disk sometimes present; stamens numerous 
in the staminate flowers, rarely definite, free or variously connate, all fertile or 
the inner or outer ones reduced to staminodia; anthers various in form; pistil- 
late flowers with staminodia, the ovary superior, 1-many-celled; ovules 1-many 
in each cell, axile or attached to the septa, rarely basal or parietal; stigmas sessile 
or on free or partly connate styles; fruit capsular, opening septicidally or rarely 
loculicidally, sometimes baccate or drupaceous; seeds often enveloped in an aril, 
without endosperm. 

About 45 genera, the species most numerous in tropical regions. 
Another genus represented in southern Central America is Chryso- 
chlamys. 

Leaves abundantly tomentose on the lower surface with branched hairs . . . Vismia. 
Leaves glabrous. 
Fruit dehiscent. 

Cells of the ovary 1-ovulate Tovomiia. 

Cells of the ovary 2-many-ovulate. 

Flowers in elongate racemes Marila. 

Flowers solitary, cymose, or paniculate, never racemose. 

Flowers dioecious or polygamous; large shrubs or trees, with large, mostly 

coriaceous leaves Clusia. 

Flowers perfect; herbs or small shrubs, rarely small trees, the leaves thin, 
usually very small. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 37 

Sepals 4, in unequal pairs; petals 4 Ascyrum. 

Sepals and petals each 5, the petals equal or nearly so Hypericum. 

Fruit indehiscent. 

Leaves with very numerous, fine lateral nerves, the nerves mostly less than 
1 mm. apart, not or obscurely connected by transverse nerves. 

Calophyllum. 

Leaves with few lateral nerves, or the nerves sometimes numerous but con- 
nected by conspicuous transverse nerves. 

Leaves broadly rounded at the apex Mammea. 

Leaves acute or acuminate. 

Stamens connate into a tube, this split above into 5 spreading lobes; 

flowers red or reddish Symphonia. 

Stamens free; flowers white or yellowish Rheedia. 

ASCYRUM L. 

Low perennials, somewhat woody, densely leafy, in appearance like Hyperi- 
cum; leaves small, entire; flowers yellow, small, subternate at the ends of the 
branches; sepals 4, the 2 outer ones large, or all the sepals subequal; petals 4, 
naked within; stamens short-connate at the base or free; hypogynous glands none; 
ovary 1-celled, the 2-3, rarely 4, placentae parietal; styles free or coherent at the 
very base; ovules numerous; capsule dehiscent along the placentae. 

About 5 species, in eastern and southeastern North America, 
Mexico, Central America, and West Indies. Only the following 
reaches Central America. 

Ascyrum hypericoides L. Sp. PI. 788. 1753. Hierba de sabino 
(fide Aguilar) . 

Brushy hillsides in pine forest, 1,000-1,800 meters; Zacapa; Ja- 
lapa; Quiche"; Alta Verapaz; Huehuetenango. Eastern United 
States; Mexico; Honduras; Cuba; Jamaica. 

A low shrub, much branched from the base or throughout, glabrous, diffuse or 
ascending, the stems 30 cm. long or less, the branches somewhat compressed and 
2-edged; leaves sessile, oblong or obovate, 0.5-2 cm. long, 1-4 mm. wide, obtuse, 
narrowed and biglandular at the base, punctate; flowers terminal and sometimes 
also axillary, golden yellow, short-pedicellate, 12-18 mm. broad; outer sepals oval 
or ovate, sometimes cordate at the base, obtuse, the inner ones narrower and gen- 
erally shorter; petals oblong-linear, about equaling the outer sepals; styles 2; cap- 
sule ovoid, 4 mm. long. 

Only the typical variety is known to occur in Central America, 
and it is not common. The leaves sometimes dimorphic. 

CALOPHYLLUM L. 

Trees with colored sap; leaves opposite, with very numerous and close, par- 
allel lateral nerves, coriaceous, petiolate; flowers small, polygamous, the inflores- 



38 FIELDI ANA: BOTANY, VOLUME 24 

cences axillary or terminal, paniculate or racemose, the nodes not articulate; 
sepals 4, decussate; petals indefinite, none, or 1 or more, the sepals and petals often 
much alike; stamens numerous, free or slightly united by their bases, the filaments 
slender; anthers ovate or oblong, opening by longitudinal slits; ovary 1-celled, the 
ovule 1, erect, anatropous; stigma peltate, sometimes lobate; fruit drupaceous. 

About 100 species, chiefly in the Old World tropics, about 7 in 
America. One other Central American species occurs in Panama. 

Calophyllum brasiliense Camb. in St. Hil. Fl. Bras. Merid. 1: 
321, pi 67. 1825. 

Widely distributed in tropical South America. Represented in 
Mexico and Central America by the following geographic variety: 

Calophyllum brasiliense var. Rekoi Standl. Trop. Woods 30: 
7. 1932. C. Rekoi Standl. Contr. U. S. Nat. Herb. 20: 192. 1919. 
C. chiapense Standl. I.e. Mario; Santa Maria; lech; leche; leche 
amarilla (Alta Verapaz). 

Wet to dry, mixed forest, most common on plains or foot-hills, 
1,200 meters or less; Pete"n; Alta Verapaz; Izabal; Chiquimula; Juti- 
apa; Santa Rosa; Escuintla; Guatemala; Chimaltenango; Solola; 
Suchitepquez; Retalhuleu; Huehuetenango; doubtless in all the 
Pacific coast departments. Southern Mexico; British Honduras to 
El Salvador and Panama. 

A tall glabrous tree, frequently 30 meters high or even more, the trunk often 
a meter in diameter, the crown rather small, spreading, small buttresses sometimes 
present; bark smooth or slightly fissured, grayish brown, the inner bark light 
brown, exuding a yellowish latex; branchlets more or less quadrangular or nar- 
rowly winged, green; leaves short-petiolate, varying from lance-oblong to elliptic 
or obovate, mostly 6-10 cm. long on fertile branches but on sterile branches often 
15-20 cm. long, lustrous, usually subacute or short-acuminate, acute at the base, 
the lateral nerves exceedingly numerous and close together, parallel; inflorescences 
racemose, axillary and terminal, mostly much shorter than the leaves, few-several- 
flowered, the pedicels 1 cm. long or shorter; sepals orbicular to oblong-elliptic, 
4-6 mm. long; petals 1-3 or none, white or yellowish, oblong, 4-6 mm. long; sta- 
mens numerous in the staminate flower, fewer in the pistillate flower; style well 
developed in the pistillate, equaling or longer than the ovary, the stigma peltate, 
slightly lobate; fruit globose or oval, 1-2 cm. in diameter, pale green, yellowish 
at maturity. 

In El Salvador sometimes called "barillo" and "varillo;" "leche 
Maria," "bari," "ocu" (Veracruz, Oaxaca); "lechoso" (Oaxaca). 
This is a well-known tree in Guatemala as well as in many other 
parts of Central America, and it is an important timber tree of 
Guatemala, especially in the Pacific foothills. It is known generally 
by the name "Mario," and the wood often is advertised for sale in 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 39 

the newspapers under this name. The term is said to have no rela- 
tion with the Spanish name Maria, although in some parts of its 
range the people have added, more or less as a matter of course, the 
term "Santa" to the feminine form of the name. Mario is believed 
to be a term of Antillean origin. This species makes an excellent 
shade tree, and often is planted for the purpose, its dense crown of 
deep green leaves reminding one of the Mamey. In El Salvador the 
yellow latex issuing from the trunk is applied to heal or hold the 
navel of young babies, when the navel does not heal promptly. 
Heartwood pink to brick-red, sapwood lighter and distinct; luster 
rather low; hard and heavy or moderately so, the specific gravity 
0.55-0.75, the weight 34-47 pounds per cubic foot; texture medium, 
fairly uniform, the grain usually roey. Lumber holds nails and screws 
firmly. The timber has been utilized for general construction, ship 
and boat building, shingles, furniture, and many other purposes in 
which attractive strong durable material is needed. Small quanti- 
ties have been exported from tropical America to the United States 
and Europe. In Central America the wood is valued for fuel, 
because it burns slowly and for a long time. It is used in some areas 
for railroad ties. This species has been reported from Guatemala as 
C. Caldba Jacq. 

GLUSIA L. 

Glabrous trees or shrubs, often epiphytic and sometimes strangling their hosts 
by their epiphytic roots, sometimes woody vines, the sap yellow, viscid, resinous, 
turning black when dried, dioecious or polygamodioecious; leaves opposite, usually 
coriaceous, usually with numerous parallel lateral nerves; inflorescences more or 
less ternately branched, rarely 1-flowered, articulate at the nodes; flowers small 
or large, unisexual or rarely perfect; bractlets often numerous decussate pairs be- 
low the flower and resembling sepals; sepals 4-6, rounded; petals 4-10, free or 
somewhat connate at the base; stamens usually numerous in the staminate flower, 
free or connate, variable in form, number, and dehiscence, often accompanied by 
staminodia, usually covered with a viscid resinous sap; pistillate flower with vari- 
ously formed staminodia; ovary 4-15-celled, with as many sessile or subsessile, 
radiate stigmas; ovules anatropous, numerous, arillate, fruit a thick-walled, leath- 
ery or fleshy capsule, septicidally dehiscent. 

Species about 145, in tropical America, 2 species in New Cale- 
donia and 1 in Madagascar. Several additional ones are found in 
southern Central America. It is difficult to present a satisfactory 
account of the Guatemalan species. Although many specimens are 
available, not all are in proper condition for study, and there is com- 
plete material of few species. To make matters worse, while some 



40 FIELDIANA: BOTANY, VOLUME 24 

of the species seem to be very uniform in all their characters, others 
are highly variable and inconstant. 

Petioles very short, thick, deeply canaliculate, usually marginate for most or all 
their length, mostly 6-10 mm. broad; leaves thick-coriaceous, rounded at 
the apex. 

Leaf blades broadest at or near the middle C. belizensis. 

Leaf blades broadest above the middle. 
Pistillate inflorescence usually several-many-flowered and long-pedunculate. 

C. confer ia. 

Pistillate inflorescence usually 3-9-flowered, short-pedunculate or sessile. 
Leaves mostly less than 1.5 times as long as broad, very thick, the nerves 

scarcely prominent beneath C. rosea. 

Leaves usually twice as long as broad or longer, thinner, the nerves very 

prominent beneath. 
Peduncle bearing at the middle a pair of reduced leaves 3-4 cm. long; 

staminate flowers 9-15 C. lusoria. 

Peduncle bearing 2 greatly reduced bracts, these not leaf-like. 

Pistillate inflorescences about 9-flowered C. conferta. 

Pistillate inflorescences 1-5-flowered. 

Capsules about 3 cm. long; anthers shorter than the filaments. 

C. Massoniana. 
Capsules 2 cm. long or shorter; anthers much longer than the short 

filaments C. Salvinii. 

Petioles usually elongate, rarely short, slender, not conspicuously canaliculate, less 
and often much less than 5 mm. broad; leaves relatively thin, flexible, rounded 
to acute at the apex. 
Leaves acute or subacute. 

Leaves mostly elliptic or oblong-elliptic, sometimes narrower, broadest at or 

near the middle, mostly acute at the base C. quadrangula. 

Leaves mostly oblanceolate or oblong-oblanceolate, sometimes broader, 
broadest well above the middle, long-attenuate to the base. 

C. guatemalensis. 

Leaves very obtuse or rounded at the apex. 
Leaves all or mostly 12-19 cm. long. 

Capsule as much as 12 cm. long; leaves elliptic-oblong to narrowly cuneate- 

obovate C. Chanekiana. 

Capsule much smaller, mostly 4 cm. long or shorter. 
Leaves broadly cuneate-obovate, 5.5-10.5 cm. wide, broadly rounded or 

almost truncate at the apex C. suborbicularis. 

Leaves oblong-oblanceolate, 4.5-7.5 cm. wide, obtuse or narrowly rounded 

at the apex C. Lundellii. 

Leaves mostly 5-8 cm. long (occasionally in C. flava to 14 cm.). 

Inner sepals about 4 mm. long; leaves when dried thin C. Gentlei. 

Inner sepals mostly 6 mm. long or even longer; leaves when dried rather 
thick and stiff C. flava. 

Clusia belizensis Standl. Carnegie Inst. Wash. Publ. 461: 72. 
1935. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 41 

Known only from the type, W. A. Schipp 1242, collected at 
Camp 33, Petn, on the British Honduras boundary, 870 meters. 

A glabrous tree 9 meters high, the trunk 30 cm. in diameter, the internodes 
short; petioles 8-10 mm. long, very broad, deeply canaliculate and marginate; 
leaf blades mostly oblong or elliptic, broadest at or near the middle, 8-14 cm. long, 
4-7 cm. wide, rounded at the apex, acute at the base, rigid-coriaceous, the lateral 
nerves very numerous and close together, ascending at an angle of 45 degrees; 
pistillate flowers solitary, on a very short thick peduncle; bractlets 2, broadly 
rounded, 8 mm. long; sepals 4, ovate-rounded, broadly rounded at the apex, the 
inner ones 15-17 mm. long. 

Known only from incomplete and somewhat imperfect material, 
but apparently a distinct and valid species. 

Clusia Chanekiana Lundell, Lloydia2: 102, /. 2. 1939. Chunup 

(Maya). 

Known only from the type, British Honduras, on tree in advanced 
forest, Valentin, El Cayo District, C. L. Lundell 6366. 

An epiphytic shrub 4 meters high, the branches pendent from the weight of the 
fruits; leaves coriaceous, the stout petioles 1.5-2.5 cm. long, not marginate; leaf 
blades narrowly cuneate-obovate to elliptic-oblong, 13-22 cm. long, 4.5-9 cm. 
wide, narrowly or broadly rounded at the apex, attenuate or cuneate at the base, 
the lateral nerves 20-26 pairs, ascending at a narrow angle, prominent on both 
surfaces; peduncles stout, 2-3 cm. long, bearing 1-3 fruits, the thick pedicels 1.5- 
2.5 cm. long, bearing 4-6 bractlets, 4 of these subtending the calyx, depressed- 
obovate, 3.5-6 mm. long; sepals 4, subequal, broadly ovate, 15 mm. long, rounded 
at the apex; capsule yellowish white, fleshy, often tinged with rose-red on one side, 
ellipsoid-oblong, as much as 12 cm. long and 4.5 cm. in diameter, 6-celled; pulp 
surrounding the seeds orange-red; persistent stigmas at the apex of the fruit 6, 
teat-like. 

The species is remarkable for its large fruits. The fruits in this 
genus are mostly large and pulpy; they lose their characteristic form 
when dried and are so difficult to preserve for the herbarium that 
they are seldom collected. The leaves are very slow in drying, when 
herbarium specimens are made, and linger in the press longer than 
those of almost any other group of dicotyledonous plants. 

Clusia conferta Standl. Field Mus. Bot. 22: 91. 1940. Jubub 
or jubup (Alta Verapaz). 

Wet, mixed forest, 1,500 meters or lower; so far as known, en- 
demic; Alta Verapaz; Huehuetenango; Izabal (type from banks of 
Rio Dulce, C. L. Wilson 401). 

A glabrous tree, sometimes 9 meters high, often epiphytic and sometimes sub- 
scandent, the branches very thick; leaves almost sessile, rigid-coriaceous, the peti- 
ole scarcely 1 cm. long, broadly winged; leaf blades oblong or cuneate-oblong, 



42 FIELDIANA: BOTANY, VOLUME 24 

20-24 cm. long, 9 cm. wide, rounded at the apex, slightly narrowed toward the 
base, the base broadly cuneate, the costa very thick, the lateral nerves very 
numerous and slender, divergent at a rather broad angle; pistillate inflorescence 
terminating a stout peduncle 5 cm. long, this naked, the flowers capitate-congested, 
about 9, sessile; bracts broadly ovate, coriaceous, closely appressed, 8 mm. long, 
obtuse or rounded at the apex; sepals 4, decussate, rounded, thick-coriaceous, 
mostly broader than long, broadly rounded at the apex; ovary large, subglobose, 
the style column thick, almost 4 mm. long; stigmas 6, ovate-oval, 2 mm. long. 

There may belong here a sterile specimen collected above Ta- 
mahu, Alta Verapaz, in wet forest at 900 meters or more. Its leaves, 
very lustrous when dry, are as much as 38 cm. long and 16 cm. wide. 

Glusia flava Jacq. Enum. PL Carib. 34. 1760. C. utilis Blake, 
Contr. U. S. Nat. Herb. 24: 14, t. 4- 1922 (type from Lago de Izabal, 
Izabal, S. F. Blake 7859). Manzanita de ratdn (fide Aguilar); quie- 
bramuelas; hoja de tortilla (Pete"n); hubuche (Pete*n, Maya); memela 
(Huehuetenango) . 

Moist or wet, mixed forest, 1,300 meters or lower, mostly at low 
elevations; Pete"n; Alta Verapaz; Baja Verapaz; Izabal; Solola; Saca- 
tep^quez; Huehuetenango. Southern Mexico; British Honduras to 
Costa Rica and probably farther southward; Jamaica. 

A shrub or small tree, usually 9 meters high or less, terrestrial or epiphytic, 
with rather slender branches; leaves slender-petiolate, mostly cuneate-obovate, 
generally 5-20 cm. long, sometimes slightly larger, rounded or very obtuse at the 
apex, cuneate-attenuate at the base, the lateral nerves numerous, very slender, 
ascending at a very narrow angle, prominent on both surfaces in the dry state; 
flowers in terminal few-flowered cymes, pink, slender-pedicellate, the pistillate in- 
florescences mostly 3-5-flowered, the pedicels usually bearing several pairs of 
bractlets; inner sepals rounded, about 7 mm. long; petals obovate, obtuse, thick 
and fleshy, 1 cm. long or sometimes larger; stamens numerous; filaments much 
shorter than the short anthers; ovary 6-12-celled; capsule usually 12-35 mm. long, 
subtended by the persistent calyx, subglobose, crowned by the conspicuous radiat- 
ing stigmas; seeds about 6 in each cell, 3.5 mm. long, surrounded by an orange aril. 

Called "matapalo" in British Honduras; "chunup," "canchunup" 
(Yucatan, Maya). This has been reported from Guatemala as C. 
minor L. and C. parvicapsula Vesque. The latter is a Mexican spe- 
cies, perhaps not distinct from C. flava. In Izabal the yellow latex 
is placed on cotton which is applied to cavities in the teeth to re- 
lieve toothache. The same vernacular name, "quiebramuelas," and 
the same use for the plant, are reported also from Honduras. A 
chemical examination of the latex might be well worth while. In 
the Yucatan peninsula the latex of this and other species is some- 
times used to adulterate chicle. It is reported that the bark is em- 
ployed for making pails or similar containers for liquids. The wood 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 43 

is not used except possibly for fuel and fence posts. In this genus the 
heartwood is brownish, often with a tinge of pink, of medium luster, 
rather light but firm to moderately heavy and hard; texture medium 
to coarse, straight-grained, easy to work; probably not very resist- 
ant to decay. 

Clusia Gentlei Lundell, Contr. Univ. Mich. Herb. 6: 48. 1941. 
Matapalo. 

Known only from the type, Stann Creek Valley, Stann Creek 
District, British Honduras, P. H. Gentle 3512. 

An epiphytic shrub, the branches slender, ferruginous, the epidermis, as in 
other species, exfoliating in small pieces that often shrivel and resemble coarse 
hairs; petioles 3-6 mm. long; leaf blades thin, almost membranaceous when dried, 
oblanceolate to narrowly cuneate-obovate, 4-10 cm. long, rounded at the apex, 
narrowly attenuate to the base, the lateral nerves about 15 pairs, ascending at a 
very narrow angle, prominulous; pistillate flowers sessile, the cymes with 9 or 
fewer flowers, the bracts 2 mm. long, triangular-ovate; sepals ovate-orbicular, the 
inner ones 4 mm. long; petals ovate-orbicular, 4 mm. long; ovary 4-celled. 

Clusia guatemalensis Hemsl. Diagn. PI. Mex. 1: 2. 1878. 
C. mexicana Vesque, Epharm. 3: 9, it. 24, 25. 1892. Matapalo; man- 
zana rosa de mico (fide Aguilar) ; palma rosa (San Marcos) ; lima real 
(San Marcos). 

Mostly in wet, mixed forest, 300-2,000 meters; Alta Verapaz; 
Zacapa; Chiquimula; Jalapa; Santa Rosa; Escuintla; Suchitep^quez; 
Guatemala; Sacatep^quez (type from Barranco Hondo, Volcan de 
Fuego, Salvin); Chimaltenango; Quezaltenango; San Marcos. South- 
ern Mexico; El Salvador. 

A shrub or a small tree, usually 9 meters high or less, terrestrial or more often 
epiphytic; leaves thin, on rather long, slender petioles, variable in form but mostly 
oblanceolate to oblong-obovate and 8-16 cm. long, acute to obtuse but always 
more or less pointed at the apex, cuneate to cuneately long-attenuate to the base, 
the lateral nerves numerous, prominulous, ascending at an angle of 45 degrees or 
less; staminate flowers creamy or pinkish white, 2.5-3 cm. broad, in 3-12-flowered 
cymes, pedicellate, the bractlets triangular, acute, 2-4 mm. long; sepals suborbic- 
ular, the inner ones 7 mm. long; filaments shorter than the anthers; fruits usually 
solitary and pendent, sometimes 3, subglobose, 2-4.5 cm. long, pale green tinged 
with red, about 8-celled, the persistent sepals as much as 11 mm. long. 

Known in El Salvador by the names "icaco montes" and "man- 
zana montes." 

Clusia Lundellii Standl. Carnegie Inst. Wash. Publ. 461: 72. 
1935. Matapalo. 



44 FIELDIANA: BOTANY, VOLUME 24 

Wet, mixed forest, chiefly at or little above sea level; Pete'n (type 
from Monte Santa Teresa, C. L. Lundell 3072) ; Alta Verapaz (planted 
at Coban); Izabal. Mexico (Tabasco); British Honduras. 

A tree of 10 meters, or a large coarse vine climbing to the tops of tall trees; 
leaves coriaceous, on rather slender petioles 1.5-3 cm. long, narrowly oblong to 
obovate-oblong or rather narrowly cuneate-obovate, 14-17 cm. long, 4.5-8 cm. 
wide, narrowly or broadly rounded at the apex, cuneate-attenuate at the base, 
the lateral nerves numerous, prominent, ascending at a very narrow angle; pistil- 
late flowers terminal, solitary, the peduncles 3-3.5 cm. long; bractlets about 5 pairs, 
4-10 mm. long, rounded-ovate; sepals unequal, ovate-rounded, 1-1.5 cm. long, 
broadly rounded at the apex; capsule oval, 3.5 cm. long, about 5-celled. 

Clusia lusoria Standl. & Steyerm. Field Mus. Bot. 23: 63. 1944. 
Lechemaria (probably a corruption of leche amarilla). 

Moist or wet, mixed forest, 1,700-2,500 meters; endemic; Zacapa; 
Chiquimula (type from middle slopes of Montana Norte to El Jutal, 
Cerro Brujo, southeast of Conception de las Minas, Steyermark 
30987). 

A tree of 15-18 meters; leaves on very thick, winged petioles 1-2 cm. long, 
oblong-obovate to broadly obovate-elliptic, 9-16 cm. long, 4-10 cm. wide, broadly 
rounded at the apex, cuneate at the base, often broadly so, lustrous above, the 
lateral nerves numerous, divergent at a rather wide angle, prominulous on both 
surfaces; staminate inflorescence 9-15-flowered, borne on a thick peduncle 5-11 cm. 
long, this bearing at the middle 2 foliaceous oblong bracts 3-4 cm. long, the flowers 
sessile or on very short and thick pedicels; bractlets 2 at the base of the calyx, large, 
suborbicular; sepals suborbicular, unequal, the inner ones 8 mm. long; petals oval, 
18 mm. long and 8 mm. wide, greenish white or pale yellow; stamens numerous, 
the linear anthers 5-6 mm. long, the filaments somewhat shorter. 

Clusia Massoniana Lundell, Contr. Univ. Mich. Herb. 4: 17. 
1940. 

Moist or wet, mixed or pine forest, 1,700 meters or less; Izabal. 
British Honduras (El Cayo District, C. L. Lundell 6605). 

A glabrous tree of 7-17 meters, the trunk as much as 25 cm. in diameter; 
leaves thick-coriaceous, the petioles 2 cm. long or shorter, very thick, broadly 
winged; leaf blades obovate or oblong-obovate, 10-19 cm. long, 4-10.5 cm. wide, 
broadly rounded at the apex, cuneately narrowed to the base, the lateral nerves 
18-30 pairs, prominulous on both surfaces; staminate flowers numerous or few, 
cymose, the cymes pedunculate, the flowers sessile; sepals suborbicular, 7 mm. 
long; petals greenish white, oblong or obovate-oblong, 1.5 cm. long or less; sta- 
mens numerous, the anthers much shorter than the filaments; capsules 3-5, oval, 
2-3 cm. long, as much as 2 cm. in diameter, 5-celled. 

This has been reported from British Honduras as C. Salvinii 
Donn.-Sm., to which it is closely related. More and better material 
is necessary to determine the validity and characters of the species 



STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 45 

of this group of the genus. It is quite possible that there are more 
good species than are recognized here. 

Clusia quadrangula Bartlett, Proc. Amer. Acad. 43: 55. 1907. 

Moist or wet, mixed forest, 900 meters or less; Pete"n; Alta Vera- 
paz; Izabal (type from Livingston, C. C. Deam 56). British Hon- 
duras; Honduras. 

A large shrub or a tree, commonly 7-9 meters high, most often an epiphytic 
shrub, the branches rather slender; leaves slender-petiolate, thin when dried, 
mostly elliptic or oblong-elliptic, 7-12 cm. long, 3-6 cm. wide, acute at each end, 
the lateral nerves very numerous, prominulous on both surfaces, ascending at an 
angle of about 45 degrees; stamina te cymes few-flowered, shorter than the leaves, 
the bractlets often as many as 5 pairs, coriaceous, resembling the sepals; sepals 
ovate-cordate, the inner ones 4-5 mm. long; petals creamy white, twice as long as 
the sepals; stamens very numerous, the filaments very short or obsolete, much 
shorter than the anthers; fruit subglobose or oval, 2 cm. long, about 9-celled. 

This has been reported from Guatemala and British Honduras as 
C. mexicana Vesque, from which it is quite distinct. The plants are 
sometimes epiphytic. On sterile shoots the leaves may be as much 
as 18 cm. long and 7 cm. wide. 

Clusia rosea Jacq. Enum. PL Carib. 34. 1760. 

Moist forest, 900 meters or less; Pete"n; Alta Verapaz; Suchitepe"- 
quez. Southern Mexico; British Honduras; Costa Rica; Panama. 
West Indies; northern South America. 

A glabrous tree or an epiphytic shrub, commonly 10-18 meters high; leaves 
thick-coriaceous, on very short, broad, winged petioles, rounded-obovate, 7-15 cm. 
long and almost as wide, broadly rounded at the apex, somewhat cuneately nar- 
rowed to the base, the lateral nerves numerous, prominulous, ascending at an angle 
of about 45 degrees; cymes few-flowered, the branches and pedicels very thick; 
bractlets 2 at the base of the calyx; sepals 4, coriaceous, the inner ones as much 
as 1.5 cm. long; petals 6, rosy white and waxy; capsule 8-12-celled, subglobose, 
5-6 cm. in diameter. 

In Costa Rica this and related species are known as "copey," and 
in the West Indies as "cupey." This term is believed to be of An- 
tillean origin. When fresh, the leaves are very thick, heavy, and 
succulent. Oviedo relates that during and after the conquest of the 
West Indies the Spaniards used the leaves in place of scarce paper 
for sending messages. With a pin or other sharp-pointed instru- 
ment, it is possible to write perfectly legible words upon the leaves. 
They also colored or marked the leaves and used them as playing 
cards. A deck of such cards must have been rather formidable to 
shuffle, in spite of the fact that the Spanish deck contains fewer than 
the customary 52 cards of the deck employed in the United States. 



46 FIELDIANA: BOTANY, VOLUME 24 

Clusia Salvinii Donn.-Sm. Bot. Gaz. 35: 1. 1903. C. Schippii 
Lundell, Contr. Univ. Mich. Herb. 4: 18. 1940 (type from Camp 32, 
Pete"n, British Honduras boundary, W. A. Schipp 1275). Matapalo; 
lengua de venado (fide Aguilar) ; cerbatana. 

Usually. in wet, mixed mountain forest, 2,600 meters or lower; 
Pete"n; Alta Verapaz; Zacapa; Chiquimula; Jalapa; Guatemala; Saca- 
tep^quez (type from Volcan de Fuego, Osbert Salviri) ; Chimaltenango; 
Quiche"; Quezaltenango; San Marcos. Southern Mexico; British 
Honduras; Honduras. 

Terrestrial or epiphytic, commonly a tree of 5-9 meters or sometimes larger, 
sometimes a more or less scandent, epiphytic shrub, the branchlets usually stout; 
leaves thinly or thickly coriaceous, the petioles 1-3 cm. long, usually thick and 
winged, rarely slender, oblong-obovate to broadly obovate, 10-18 cm. long, 5-9 cm. 
broad, rounded at the apex, more or less cuneate at the base, the lateral nerves 
numerous, prominulous on both surfaces, ascending at an angle of about 45 de- 
grees; inflorescences mostly 3-5-flowered, the flowers sessile or on short thick ped- 
icels, the bractlets usually 1 pair, similar to the sepals; sepals suborbicular, 7-9 mm. 
long; stamens 45-50, the anthers about 3 times as long as the short filaments; cap- 
sule globose, about 6-celled, 1.5-1.8 cm. in diameter, the seeds numerous, or most 
of them abortive. 

C. Schippii may be a distinct species, but the material of it now 
available offers no characters by which it can be distinguished from 
C. Salvinii. 

Clusia suborbicularis Lundell, Phytologia 1: 218. 1937. Sello. 

Moist or wet forest or thickets, 300 meters or less; Pete"n; Izabal. 
British Honduras. 

A large or small, glabrous tree, sometimes 17 meters high, with a trunk 25 cm. 
in diameter; leaves rather thinly coriaceous, the petioles rather slender, not winged, 
1.5-2 cm. long; leaf blades cuneate-obovate, usually rather broadly so, 12-16 cm. 
long, 9-12 cm. wide, broadly rounded or almost truncate at the apex, cuneately 
narrowed to the base, the lateral nerves 20-25 pairs, prominulous on both surfaces, 
ascending at an angle of 45 degrees or less; inflorescences 3-7-flowered, the pedicels 
short and thick, the bractlets 3-5 pairs, crowded, ovate-orbicular, 3-9 mm. long; 
sepals depressed ovate-orbicular, the inner ones 11 mm. long; stamens exceedingly 
numerous, the filaments very short, almost obsolete; capsule oval-globose, 5-8- 
celled, 2.5-3.5 cm. long. 

The Maya name is reported as "chunup" in British Honduras. 



Garcinia Mangostana L. is the Mangosteen, a tree native in the 
Malay region. Its fruit is considered by some persons to be the most 
delicious of all tropical fruits. Because of difficulties attending its 
propagation, and especially because it seems to demand special soil 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 47 

and climatic conditions, it never has become common in tropical 
America, and the total number of trees in Central America is very 
small, although they have fruited abundantly at various places along 
the Atlantic coast. A famous tree on the shore of Lake Izabal was 
leased for some time by the United Fruit Company, to obtain the 
seeds for planting. Trees have fruited at Quirigua, and they have 
done well at Tela, Honduras. The rather small, somewhat depressed 
fruits are dark purple, about 5-8 cm. in diameter, and contain several 
large seeds. 

HYPERICUM L. St. Johnswort 

Annual or perennial herbs or shrubs, rarely trees, usually glabrous throughout; 
leaves small, opposite, entire, often gland-dotted, sessile or short-petiolate; flowers 
5-4-parted, yellow, often black-punctate, variously arranged; sepals alike or dis- 
similar; petals usually contorted in bud, often persistent in fruit; stamens numer- 
ous, free or united in 3 or 5 phalanges opposite the petals; ovary 3-5-carpellate; 
styles 3-5, free or more or less united, the stigmas subcapitate; ovules generally 
numerous; fruit capsular, membranaceous or coriaceous, 1-celled or more or less 
completely 3-5-celled; endosperm none; embryo straight or curved. 

About 300 species, in tropical and temperate regions of both 
hemispheres, in the tropics found mostly in the mountains. A few 
other species grown in the mountains of Costa Rica and Panama. 

Leaves linear. 
Leaves minute, 2.5 mm. long or less, scale-like, closely appressed to the stem. 

H. aphyllum. 
Leaves much larger, spreading or appressed. 

Styles 4 (3?); sepals linear; petals obliquely apiculate and 6-9 mm. long; cap- 
sule oblong-cylindric H. fasciculatum. 

Styles 3; sepals linear-lanceolate to lanceolate-oblong; petals not obliquely 

apiculate and mostly less than 6 mm. long; capsule ovoid. 
Styles much longer than the ovary, about as long as the mature capsule; 

leaves imbricated, revolute, erect H. arbuscula. 

Styles usually not longer than the ovary and much shorter than the cap- 
sule; leaves mostly flat, or sometimes revolute, often spreading or only 

ascending, not or little imbricated H. uliginosum. 

Leaves lanceolate to oblong or broader. 

Annuals or if more enduring then with long perpendicular roots, never root- 
stocks H. gymnanthum. 

Perennials with rootstocks, or shrubs or small trees. 

Herbs erect or prostrate, herbaceous or stems suffruticose at the bases. 
Flowers solitary at the ends of branches; stamens 16-20; stems slender and 
weak, usually prostrate or procumbent H. epigeium. 

Flowers in corymbs; stamens numerous; stems stout, stiff and erect. 

H, formosum. 
Shrubs or small trees, the stems woody throughout or nearly so. 

Leaves small, all or mostly less than 1 cm. long H. calcicola. 



48 FIELDIANA: BOTANY, VOLUME 24 

Leaves larger, mostly 2-6 cm. long. 

Sepals about 1 cm. long, long-acuminate; leaves mostly about 7 mm. 
broad or narrower, crowded on the branches H. styphelioides. 

Sepals 5 mm. long or less, broadly ovate, scarcely acute; leaves mostly 
10-20 mm. broad, not crowded H. Steyermarkii. 

Hypericum aphyllum Lundell, Am. Midi. Nat. 29: 477. 1943. 

Type from Monkey River, Toledo District, in open pine ridge 
near Jenkins Creek, British Honduras, P. H. Gentle 4175. Reported 
also from Florida. 

A slender stiff erect annual, 65 cm. high or less, the branches few, erect; leaves 
opposite, reduced to linear-subulate appressed scales 2.5 mm. long or less; flowers 
small, scattered along the erect branches, sessile or subsessile; sepals 5, linear- 
subulate, 2 mm. long; petals linear, yellow, 2.5 mm. long; stamens only 5-6, 
shorter than the petals, the filaments free; ovules very numerous; styles 3, free, 
1 mm. long; capsule fusiform, 3-4.5 mm. long; seeds minute, oblong, 0.25 mm. 
long, almost smooth. 

Related to the common H. gentianoides (L.) BSP. of the United 
States, but said to differ in seed characters. 

Hypericum arbuscula Standl. & Steyerm. Field Mus. Bot. 23: 

63. 1944. 

Endemic; Baja Verapaz (type from Fatal, John R. Johnston 1812; 
taken by the same collector "between Fatal and Salama," and the 
type may have been collected farther southward than Fatal, perhaps 
about Santa Rosa) ; Quiche". 

A low, densely branched shrub about 30 cm. high, the branches very numerous, 
rather stout, woody throughout, erect or nearly so, very densely leafy; leaves ses- 
sile, erect or strongly ascending, imbricate, linear, 1-nerved, 6-9 mm. long, strongly 
revolute, acute, coriaceous, somewhat lustrous, not punctate; flowers few at the 
ends of the branches, short-pedicellate, 5-parted; sepals narrowly lance-oblong, 
acute, nerved, 5 mm. long, epunctate; petals yellow, slightly longer than the 
sepals, narrowly spatulate, inflexed and cucullate at the apex, not punctate; sta- 
mens numerous, distinct, equaling the petals; styles 3, very slender, as much as 
4 mm. long or even longer; capsule ovoid-oblong, 3 mm. long, acute, subtrigonous, 
1-celled; seeds few, oblong, terete, brown, almost smooth, 0.8 mm. long. 

Hypericum calcicola Standl. & Steyerm. Field Mus. Bot. 23: 

64. 1944. 

Known only from the type, on limestone bluffs, common, Dept. 
Huehuetenango, between Tojquia and Caxin Bluff, summit of Sierra 
de los Cuchumatanes, 3,700 meters, Steyermark 50160. 

A very densely branched shrub, woody throughout, erect and 30-60 cm. high 
or depressed and spreading, the older branches blackish, the younger ones reddish, 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 49 

densely leafy; leaves small, short-petiolate, opposite, subcoriaceous, elliptic or 
oval, sometimes broadly ovate, 5-10 mm. long, 3-6 mm. wide, obtuse or narrowly 
rounded at the apex, green above, epunctate, the veins evident and reticulate, 
paler beneath, sparsely and coarsely punctate, the veins and nerves laxly and rather 
conspicuously reticulate; flowers solitary at the ends of short branches, on short or 
elongate pedicels; sepals 4, ovate-oblong, 2.5 mm. long, pale rose-colored, acute, 
punctate; petals rose-colored, reflexed, oblong, 7 mm. long, obtuse; stamens about 
10; styles 3, about 2 mm. long, recurved in anthesis; capsule narrowly ovoid, almost 
5 mm. long, pale rose, acute, 3-sulcate, 3-celled; seeds not numerous, narrowly 
oblong or subfusiform, pale, 1 mm. long. 

The species is a very distinct one, and quite unlike any other 
found in Central America. In general appearance it suggests the 
densely branched forms of Symphoricarpos microphyllus. 

Hypericum epigeium Keller, Bull. Herb. Boiss. II. 8: 183. 1908. 

Moist or wet, usually open forest, most frequently in pine forest, 
sometimes on banks or limestone cliffs, 2,100-3,800 meters; Sacate- 
pe"quez (Volcan de Agua, the type J. D. Smith 2147); Chimalte- 
nango; Huehuetenango (Sierra de los Cuchumatanes) ; Totonicapan; 
Quezaltenango. Mexico. 

A slender perennial herb, rarely somewhat suffrutescent, usually much 
branched, the stems erect or usually procumbent or prostrate, often forming 
large colonies, the stems rather densely leafy; leaves usually lax and spreading, 
thin, sessile, punctate, elliptic to lance-oblong, mostly 1-2 cm. long, sometimes 
larger, obtuse or subacute at the base; flowers solitary at the ends of the branches, 
pedicellate; sepals ovate-lanceolate, acute, 3-4.5 mm. long; petals obovate, rounded 
at the apex, 8-9 mm. long; stamens about 20, free or nearly so; styles 3, about 
3 mm. long; capsule ovoid, 3-sulcate. 

This has been reported from Guatemala as H. mexicanum L. 
Here quite possibly belongs H. chamaecaulon Keller (Bull. Herb. 
Boiss. II. 8: 179. 1908). That name has never been properly pub- 
lished, but is said to pertain to a Guatemalan plant. It appeared in 
a key published by Keller in the place cited. 

Hypericum fasciculatum Lam. Encycl. Meth. 4: 160. 1797. 

Along open stream beds or in shallow sandy-bottomed ponds or 
in savannas. British Honduras; southeastern United States; West 
Indies. 

Slender shrubs or small trees occasionally to 4.5 m. but usually about 1 m., 
much branched; leaves opposite or fasciculate, linear, obtuse or mostly glandular- 
apiculate, rounded above, canaliculate below, punctate-glandular, mostly 1-2 cm. 
long and about 0.1 cm. broad; flowers yellow, solitary on the ends of branches 
but generally surpassed by new shoots, sessile or nearly so; sepals linear, apiculate, 
4-6 mm. long and 1 mm. or less broad, punctate, similar to the leaves but smaller; 



50 FIELDIANA: BOTANY, VOLUME 24 

petals cuneate to the base, subtruncate and obliquely apiculate, 6-9 mm. long; 
capsule 6-7 mm. long, oblong-cylindric, quadrisulcate; styles 4 in our material, 
about as long as the capsule. 

The material from British Honduras differs from that of other 
regions in having the leaves less conspicuously fasciculate or not at 
all so, the styles 4, and the capsule longer and oblong instead of 
conic-ovoid. 

Hypericum formosum HBK. Nov. Gen. & Sp. 5: 196. 1821. 

Moist or wet forest or on open banks, 2,400-2,800 meters; Baja 
Verapaz (Fatal); Quiche" (between Chajul and Nebaj); Huehuete- 
nango (Sierra de los Cuchumatanes). Western United States; 
Mexico. 

An erect perennial herb a meter high or less, sometimes suffrutescent below, 
the stems solitary or clustered, usually branched; leaves sessile, oval or elliptic, 
1.5-3 cm. long, rounded or obtuse at the apex, obtuse or subacute at the base, 
coarsely punctate, paler beneath; flowers bright yellow, corymbose, often numer- 
ous, pedicellate; sepals lanceolate, attenuate, 6 mm. long, often punctate; petals 
oval, 10-12 mm. long; stamens very numerous, free or nearly so; capsule oblong- 
ovoid, 8 mm. long, 3-sulcate; styles 3. 

Hypericum gymnanthum Engelm. & Gray, Bost. Journ. Nat. 
Hist. 5: 212. 1847. 

Moist, open places, or marshes, 1,300-1,400 meters; Alta Verapaz 
(region of Coban and Santa Cruz, apparently scarce). Honduras. 
Widely distributed in the United States. 

A slender, erect annual, simple or branched, usually 40 cm. high or less; leaves 
sessile, thin, ovate or the lowest oval, 1-2 cm. long, sessile and often somewhat 
amplexicaul, acute or the lower ones obtuse, 3-7-nerved at the base, finely punc- 
tate; flowers small, in open terminal cymes, the bracts subulate; sepals lanceolate, 
4.5 mm. long, acuminate; petals equaling or slightly longer than the sepals, yellow; 
capsule 1-celled, 4 mm. long; styles 3; stamens 10-12. 

Hypericum Steyermarkii Standl. Field Mus. Bot. 22: 160. 
1940. 

In forest of pine and Abies, 2,500-3,500 meters; Quezaltenango 
(type from Volcan de Santo Tomas, Steyermark 34760) ; San Marcos. 
Mexico (Chiapas). 

A branched glabrous shrub or tree 2-6 meters high, the branches brown or 
brownish; leaves sessile, membranaceous, densely pellucid-punctate, oblong-ovate 
to lance-oblong, 3-6.5 cm. long, 1-2.2 cm. wide, acute and apiculate, obtusely 
cuneate at the base, paler beneath; flowers few, very large, terminal, fasciculate, 
usually 3-4, the pedicels 1-2 cm. long; sepals broadly ovate, 5-6 mm. long, acute, 
green, black-glandular on the margins; petals deep yellow, 2 cm. long, multilineate, 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 51 

obovate, rounded at the apex, sparsely punctate near the apex; stamens numerous, 
short-connate at the base, in 5 phalanges, half as long as the petals; styles 5, dis- 
tinct; capsule ovoid, 14 mm. long, subobtuse, 5-celled; seeds numerous, oblong, 
2-2.5 mm. long, pale brown, minutely reticulate. 

Hypericum styphelioides A. Rich. Ess. Fl. Cub. 237. 1845. 
H. terrae-firmae Sprague & Riley, Kew Bull. 1924: 12. 1924 (type 
from British Honduras, M, E. Peck 321). 

Along stream banks in pine forest, at or little above sea level, 
British Honduras. Cuba. 

A glabrous branched shrub 2 meters high or less, woody throughout, the 
branches densely leafy; leaves sessile, oblong-lanceolate, ascending, 1.5-2.5 cm. 
long, acute, narrowed to the base, 5-nerved below, punctate; flowers sessile at the 
ends of the branches, surrounded by leaves; sepals lanceolate, acuminate, about 
1 cm. long, about 10-nerved, epunctate; petals about 2 cm. long, yellow, 6 mm. 
wide, many-nerved; stamens very numerous, 1 cm. long; ovary 1-celled, the styles 
5 and 5 mm. long; capsule 5 mm. long; seeds cylindric, slightly curved, 1 mm. 
long, punctate-striate. 

Sprague and Riley contrasted H. terrae-firmae with H. stypheli- 
oides in several minor characters. These do not hold for the more 
ample material now at hand, and it is not at all unusual to find a 
Cuban plant such as this extending to the mainland of Central 
America, more especially to the Yucatan peninsula. 

Hypericum uliginosum HBK. Nov. Gen. & Sp. 5: 194. 1821. 
Mil flores (fide Aguilar) ; retij (Coban, Quecchi) ; ruda de monte. 

Open fields and hillsides, pine, oak, Cupressus, or Alnus forest, 
often in wet or marshy places but frequently in dry stony ground, 
3,400 meters or less, most frequent at middle elevations; Alta Vera- 
paz; Baja Verapaz; El Progreso; Chiquimula; Zacapa; Jalapa; Juti- 
apa; Guatemala; Sacatepe"quez; Chimaltenango; Solola; Quiche"; 
Huehuetenango; Totonicapan; Quezaltenango; San Marcos. Mex- 
ico; British Honduras; Honduras; El Salvador; Nicaragua; Costa 
Rica; Panama. 

Plants essentially annual but sometimes persisting for more than a year, aris- 
ing from a perpendicular tap root or from several stout roots, erect, simple or 
usually branched, mostly 50 cm. high or less; leaves linear or nearly so, mostly 
1-2 cm. long, acute or obtuse and apiculate, narrowed to the sessile base, 1-nerved, 
puncticulate, paler beneath, spreading or ascending, usually flat but sometimes 
revolute; flowers usually numerous, subcorymbose along the short or elongate, 
more or less secund branches, sessile or nearly so; sepals linear-lanceolate, long- 
attenuate, striate, epunctate; petals deep yellow, little longer than the sepals; sta- 
mens about 20, free; styles 3; capsule lance-oblong, 5-6 mm. long, acute, 1-celled; 
seeds very numerous, oval, brownish, scarcely 0.5 mm. long, almost smooth. 

UNIVERSITY OF ILLINOIS 
LIBRARY 



52 FIELDIANA: BOTANY, VOLUME 24 

This has been reported from Guatemala as H. pratense Cham. & 
Schlecht. and as H. paucifolium Wats. It is one of the commonest 
and most widely distributed of Guatemalan plants and is found in 
almost every kind of habitat except dense, wet forest. It is some- 
what weedy by nature and frequently invades cultivated or over- 
grazed land. In Guatemala it is said to be used sometimes in 
domestic medicine. 

MAMMEA L. Mamey 

Trees with resinous sap; leaves opposite, thick, penninerved; inflorescences 
axillary, sessile, 1-3-flowered, the flowers polygamous; calyx closed in bud, split- 
ting into 2 valves; petals 4-6; disk none; stamens numerous, the filaments filiform, 
free, the anthers linear, longitudinally dehiscent; ovary 2-celled, the ovules 2 in 
each cell, or 4-celled with 1-ovulate cells, the ovules basal, erect; style thick, the 
stigma large, peltate, 2-4-parted; fruit large, baccate. 

Four species, the following one and three others in tropical Africa. 

Mammea americana L. Sp. PI. 512. 1753. Mamey; muc 
(Quiche*). 

Native of tropical America, possibly of the West Indies; not 
native in Central America but grown commonly for its edible fruit; 
planted rather frequently in Guatemala, chiefly at 1,000 meters or 
less, and especially frequent along the Pacific foothills and plains; 
rarely planted as high as 1,500 meters, but not thriving at high 
elevations. Figure 9. 

A medium-sized tree, sometimes 15 meters high but usually lower, the crown 
rounded, dense; leaves persistent, somewhat coriaceous, dark green, on petioles 
1-1.5 cm. long, broadly elliptic to obovate, usually broadly rounded at the apex, 
acute or obtuse at the base, 8-15 cm. long or even larger, glabrous, the lateral 
nerves very numerous, parallel, prominent on both surfaces in the dry state and 
reticulately connected by elevated veins, the leaves densely pellucid-punctate; 
flowers part pistillate and part perfect, solitary or clustered, the pedicels 1-1.5 cm. 
long, the buds globose; sepals semiglobose, concave, thick, 1-1.5 cm. long; petals 
4-6, orbicular or oblong, white, obtuse, 1.5-2.5 cm. long; fruit subglobose, apicu- 
late, 10-15 cm. long, the exocarp thick, russet-colored, and rough, the mesocarp 
rather firm and juicy, yellow or reddish-yellow, sweet; seeds 4 or fewer, the testa 
rough, fibrous. 

Sometimes called "mamey de Santo Domingo" in Yucatan; the 
Maya name is reported as "chacalhaaz." For some unknown rea- 
son, the mamey does not seem to be popular in Central America, 
and it is rare in many regions. Many northern people, for some ob- 
scure reason, consider it one of the best of tropical fruits. It is 
somewhat like an apricot or a clingstone peach. Some of the fruits 




FIG. 9. Mammea americana. Portion of a branch and fruits; reduced. 



53 



54 FIELDIANA: BOTANY, VOLUME 24 

are very large, and they are said to weigh rarely as much as 4 kilo- 
grams. Some of those in Guatemalan markets are very fine, as large 
as the largest oranges of the United States. The fruits were observed 
as plentiful in the markets of the central cities at the end of January, 
and they are obtainable at other seasons as well. The juice is some- 
times used in Guatemala for flavoring aguas gaseosas or soda water. 
The fruit may be eaten either raw or cooked and sometimes is made 
into preserves. The flowers are fragrant. The name "mamey" is 
believed to be of Antillean origin. A caserio of the Department of 
Retalhuleu is called El Mamey. A tincture made by dissolving the 
gum of the tree in alcohol is employed externally in Guatemala for 
killing niguas, redbugs, and other insects. In some regions the tim- 
ber is employed for fuel, fence posts, and miscellaneous purposes, 
but in Central America the trees are protected for their fruit. The 
heartwood is dull reddish or purplish-brown, merging gradually into 
the lighter sapwood, and it is hard, rather heavy, strong, of medium 
texture and fairly straight grain, and moderately resistant to decay. 

MARILA Swartz 

Trees or shrubs; leaves opposite, petiolate, penninerved, the primary nerves 
connected by numerous parallel secondary ones; inflorescences axillary, racemose, 
the flowers perfect or apparently sometimes unisexual; sepals 4-5; petals 4-5; sta- 
mens numerous, the filaments linear, free or nearly so, the connective ending in a 
glandular process; anthers opening by longitudinal slits; ovary 3-5-celled; ovules 
numerous in each cell, attached to a simple or longitudinally cleft, axile placenta; 
style simple, the stigma subentire; fruit capsular; seeds comose at each end. 

About 20 species, in tropical America. There are two other spe- 
cies in Central America. 

Marila verapazensis Donn.-Sm. Bot. Gaz. 47: 253. 1909. 

Wet, mixed forest, 350 meters; Alta Verapaz (type from Cubil- 
giiitz, Tuerckheim 8660). British Honduras; Nicaragua; Costa Rica; 
Panama (?) ; Colombia. 

An almost glabrous tree, sometimes 12 meters high, branching near the ground, 
the trunk as much as 25 cm. in diameter; leaves short-petiolate, coriaceous, oblong, 
16-24 cm. long, 5-8.5 cm. wide, abruptly acuminate, rounded to subacute at the 
base, punctate beneath and sometimes puberulent on the nerves, the lateral nerves 
13-15 pairs, very prominent, divergent at a wide angle; racemes about as long as 
the leaves, puberulent, many-flowered, the pedicels remote, spreading, the flowers 
5-parted, green, odorless; sepals ovate or oval, puberulent, 7-8 mm. long, reflexed 
in anthesis; petals 9 mm. long, obovate-oblong; anthers very numerous, produced 
at the apex into a linear appendage; ovary glabrous, the style 5 mm. long. 



STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 55 

Cuatrecasas (An. Inst. Biol. M4x. 20: 95. 1950) has suggested 
that M. verapazensis is a synonym of M . laxiflora Rusby, of Bolivia. 
This has not been verified; however, M. laxiflora var. multinervia 
Cuatr. from the Colombian Choco seems to be the same as the Cen- 
tral American plant. 

RHEEDIA L. 

Trees or shrubs, polygamodioecious, the sap yellowish; leaves opposite or 
verticillate, penninerved, coriaceous, with a marginate pit in the axillary part of 
the petiole; flowers small, solitary or fasciculate in the leaf axils or at defoliate 
nodes; sepals 2, more or less united at the base, petals 4, decussate and imbricate; 
stamens numerous in the staminate flower, free, inserted about and sometimes 
upon a thick fleshy disk, the anthers subglobose or curved, dehiscent by longi- 
tudinal slits; perfect flowers with fewer stamens, these inserted about the base of 
the disk; ovary superior or partly surrounded by the disk, 3-4-celled, the cells 
1-ovulate; stigma peltate-discoid to hemispheric, entire or lobate, subsessile or 
upon a short style; fruit baccate, crowned by the persistent stigma, the skin cori- 
aceous, smooth or verrucose, 1-celled, containing 1-5 seeds, these enveloped in a 
pulpy aril. 

About 40 species, four in Madagascar, the others in tropical 
America. Several additional ones are found in southern Central 
America. 

Flower buds just before anthesis 8-12 mm. in diameter; sepals 8-10 mm. long; 

fruit about 6 cm. long R. macranlha. 

Flower buds before anthesis about 4 mm. in diameter; sepals 4 mm. long or less; 
fruit usually about 2 cm. long. 

Leaves of fertile branches mostly 13-22 cm. long R. intermedia. 

Leaves of fertile branches mostly 8-10 cm. long R. edulis. 

Rheedia edulis Triana & Planch. Ann. Sci. Nat. IV. 14: 310. 
1860. Mameyito; Array an. 

Moist or wet forest, 1,200 meters or less, probably planted at 
higher elevations; Santa Rosa; Guatemala; Sacatepquez; Quiche". 
Southern Mexico; British Honduras to El Salvador and Panama. 
Figure 10. 

A small to large, glabrous tree, sometimes 30 meters high, the branchlets green, 
more or less tetragonous; leaves short-petiolate, coriaceous, often deep red when 
young, elliptic-oblong or elliptic-lanceolate, those of adult branches mostly 8- 
10 cm. long, on sterile branches often much larger, obtuse or subacute, acute at 
the base, somewhat paler beneath, the lateral nerves very numerous, prominent 
on both surfaces; pedicels usually in dense fascicles at defoliated nodes, the slender 
pedicels usually 13 mm. long or shorter; sepals oval or rounded, thin, 3-4 mm. long; 
petals 4, white, ovate-orbicular, 5-7 mm. long; stamens 25-30 in the staminate 
flowers, 10-12 in the perfect flowers; fruit oval, yellow at maturity, usually about 
2 cm. long, smooth, rounded at each end. 




FIG. 10. Rheedia edulis. A, Flowering branch- X 
X 2J4 C, Perfect flower; X 2%- 



B, Staminate flower; 



56 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 57 

Called "chaparrdn" in El Salvador. This is a rather common 
tree on the Pacific plains and foothills and probably grows in all the 
Pacific coast departments. In this genus the heartwood is dull gray- 
ish to pinkish brown, hard, moderately heavy, tough and strong, 
coarse-textured, with irregular grain; not difficult to work but does 
not finish very smoothly; of fair durability. The bark is said to be 
rich in tannin. The timber is used locally for tool handles, general 
construction and carpentry, fence posts, and railway crossties. The 
fruit of all the species, probably, is edible. 

Rheedia intermedia Pittier, Contr. U. S. Nat. Herb. 13: 454, 
/. 81. 1912. Leche amarilla; polo bayo (Pete"n, fide Lundell); crauel 
(Alta Verapaz) ; jocomico; jocote de mico (Izabal, fide Blake; probably 
erroneous names). 

Wet, mixed forest, often on limestone, 500 meters or less; Pete"n; 
Alta Verapaz (type from Rio Lanquin, Tuerckheim 8180); Izabal. 
British Honduras; Honduras; Nicaragua. 

Usually a large glabrous tree, sometimes 18 meters high; leaves coriaceous, 
short-petiolate, elliptic-oblong or lance-oblong, those of fertile branches mostly 
13-22 cm. long, those of sterile branches often larger, acuminate or short-acumi- 
nate, often abruptly so, obtuse or cuneate at the base, slightly paler beneath, the 
lateral nerves very numerous, prominent; flowers fasciculate in the leaf axils or at 
defoliate nodes, few or numerous, the slender pedicels often 2 cm. long; sepals 
rounded, 2-3 mm. long; petals white, ovate-orbicular, 3.5-4 mm. long; fruit oval, 
yellow at maturity, 2 cm. long, generally 1-seeded. 

Called "waika plum" in British Honduras; "caimito," "caimito 
de montana" (Honduras); "sakipa" (Honduras; an Indian name). 
This has been reported from Guatemala as R. macrophylla Planch. & 
Triana, var. Benthamiana Vesque, and from British Honduras as 
R. edulis Triana & Planch. The bark is described as pale brownish 
gray with very small, knobby protuberances. 

Rheedia macrantha Standl. & Steyerm. Field Mus. Bot. 23: 
65. 1944. 

Wet, mixed forest, 900-1,400 meters; often on limestone; en- 
demic; Alta Verapaz (type collected along Rio Frio, about 8 km. 
below Tactic, Standley 90524); Huehuetenango(?). 

A medium-sized tree, 9 meters high or more with a trunk 15-20 cm. in diameter, 
glabrous, the branchlets more or less tetragonous; leaves on thick petioles 1-2 cm. 
long, coriaceous, elliptic-oblong or lance-oblong, 14-24 cm. long, 4.5-8 cm. wide, 
acuminate, cuneate or obtuse at the base, lustrous, paler beneath, the lateral 
nerves very numerous, prominent on both surfaces; flowers large, greenish, 
fasciculate in the leaf axils or at defoliate nodes, the slender pedicels 2.5-4.5 cm. 



58 FIELDIANA: BOTANY, VOLUME 24 

long, the buds 8-10 mm. in diameter; sepals suborbicular, 8-10 mm. long; petals 
broadly oval or almost orbicular, 8-10 mm. long; stamens numerous in the stam- 
ina te flower, the filaments stout, dilated toward the base; ovary deeply sulcate 
vertically, the stigma disciform, thick, 3 mm. broad; fruit ellipsoid, 6 cm. long, 
3.5 cm. thick, obtuse at the base, contracted toward the apex. 



SYMPHONIA L. f. 

Trees or shrubs with yellowish sap; leaves opposite, penninerved, the petioles 
without an axillary marginate pit; flowers perfect, in terminal or axillary, mostly 
umbellifonn cymes, rarely solitary; sepals 5, imbricate; petals 5, larger than the 
sepals, contorted, forming a globose or obovoid bud; disk cupular, extrastaminal, 
slightly 5-lobate; androecium columnar, split in the upper half into 5 narrow 
lobes opposite the petals, each lobe bearing 2-4 linear anthers, the anthers dehiscent 
by lateral slits; ovary 5-celled, the cells 2-8-0 vulate; style elongate, cleft into 5 
conic radiating lobes at the apex, the stigmas minute, pore-like; fruit baccate, 
few-seeded. 

About 16 species, one in tropical America and Africa, the others 
in Madagascar. 

Symphonia globulifera L. f. Suppl. PI. 302. 1781. Barillo; 
leche amarilla; pimientillo. 

Wet, often swampy forest, at or little above sea level; Izabal. 
British Honduras, along the Atlantic coast to Panama; West Indies; 
South America; tropical Africa. 

A large, glabrous tree, sometimes 30 meters high, the trunk sometimes more 
than a meter in diameter, the bark rough, brownish or darker, the trunk some- 
times supported on stilt roots, the crown rounded, with horizontal or often re- 
curved and pendent branches; leaves short-petiolate, coriaceous, lanceolate to 
lance-oblong or obovate-lanceolate, 6-12 cm. long, obtusely long-acuminate, 
narrowed to the acute base, the lateral nerves very numerous and slender, prom- 
inulous beneath; flowers on pedicels 4-13 mm. long, or in fruit 1-2.5 cm. long; 
sepals orbicular or broadly ovate, 2-8 mm. long; petals red, orbicular, 11-14 mm. 
long; fruit globose to ovoid, 3-4 cm. long, dark green at first, brownish or yellowish 
in age; seeds 1-3, with a thin testa. 

Called "waika chewstick," "leche amarillo macho," "wycot," and 
"corb&ti" in British Honduras. One of the common large trees of 
the lowland forests on the Atlantic coast of Central America. When 
growing in the open where they can develop perfectly, the trees often 
look from a short distance strangely like the American elm (Ulmus 
americana L.) of the eastern United States. The bark contains a 
yellowish resin that becomes black and pitch-like upon exposure, 
and is used in some regions for calking boats. The timber is em- 
ployed in tropical America for construction, carpentry, crates and 
boxes, railway crossties, and fuel. Small shipments have been made 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 59 

to Europe and North America, mostly for use as veneers for plywood. 
Heartwood yellowish, grayish, or greenish brown, with a somewhat 
mealy appearance; sapwood whitish; hard and moderately heavy, 
the specific gravity 0.65-0.78; texture coarse, the grain straight or 
irregular; durability fair. 

TOVOMITA Aubl. 

Shrubs or small trees, glabrous or nearly so, dioecious or polygamodioecious; 
leaves opposite, petiolate, thin, penninerved; flowers small, in terminal paniculate 
cymes, the pistillate flowers usually fewer and smaller than the staminate ones; 
sepals 2-4, if 3 or 4 then the outer ones smaller; petals usually 4, sometimes 5-8; 
stamens numerous, free or somewhat connate; ovary 4-5-celled, with one erect 
ovule (sometimes 2) in each cell; style very short; fruit a 4-5-valvate capsule, 
usually with fleshy pericarp; seeds solitary (or two) in the cells; trigonous, sur- 
rounded by a fleshy aril. 

Perhaps 25 species in tropical America. The genus as here de- 
scribed includes Tovomitopsis Planch. & Triana. 

Tovomita nicaraguensis (Oerst.) L. Wms. Tropical Woods 111: 
18. 1959. Chrysochlamys guatemaltecana Donn.-Sm. Bot. Gaz. 13: 
26. 1888 (type from Guatemala, Tuerckheim 989). Tovomitopsis gua- 
temaltecana Standl. & Steyerm. Field Mus. Bot. 23: 66. 1944. 

Wet, mixed forests, usually below 1,000 meters; Pete"n; Alta 
Verapaz; Izabal; Huehuetenango. British Honduras to Panama. 

A glabrous tree 7-15 m. tall, the branchlets terete, stout; leaves oblanceolate 
to oblong-obovate, acute or short-acuminate, cuneate-attenuate to the base, 
10-23 cm. long and 4.5-9 cm. broad, with 10-12 pairs of lateral nerves, very slender, 
prominulous beneath, petioles 2-2.5 cm. long; panicles terminal, short-pedunculate, 
about 10 cm. long and often as broad, many-flowered, several times branched, the 
flowers white, fragrant, long-pedicellate, the buds globose; sepals 4, in two pairs, 
the outer pair smaller, 6 mm. or less long; petals 5, broadly obovate, 8 mm. or 
less long, resinous striate or punctate; stamens in the pistillate flowers reduced 
and perhaps some sterile, many, the filaments somewhat dilated toward the 
base; stigmas 5, cuneate-obovate; ovary 5-celled; fruit obovoid, about 2 cm. long, 
splitting open at maturity, the valves widely spreading, thick and fleshy. 

VISMIA Vandelli 

Trees or shrubs with orange-red sap; pubescence often of a stellate tomentum; 
leaves opposite, penninerved, often glandular-punctate; flowers paniculate, the 
branches of the panicle opposite, the pedicels articulate near the base; sepals gen- 
erally scarious-margined; petals mostly lanate inside; both sepals and petals often 
with glandular dots and lines; stamens numerous, connate almost half their length 
into 5 fascicles opposite the petals, alternating with 5 much shorter, obovate or 
oblong lobes; anthers ovoid or globose, longitudinally dehiscent; ovary 5-celled; 



60 FIELDIANA: BOTANY, VOLUME 24 

ovules numerous or rarely few in each cell; styles 5, almost free, the stigmas sub- 
capitate; fruit baccate; seeds cylindric, straight or slightly curved, verrucose. 

The genus Vismia is in need of revision. The Central American 
species have received little attention in recent years and their rela- 
tion to the species of northern South America is not well understood. 
It is believed that the specific names used here apply properly to the 
species concerned, but there may be an older name for each of them. 

Species about 15 in tropical America and about five in tropical 
Africa. 

Leaves with coarse ferruginous tomentum on the lower surface, the tomentum not 
appressed and often of stellate hairs V. camparaguey. 

Leaves with fine, soft, closely appressed tomentum on the lower surface, often 
brownish at first but in age grayish or white V. mexicana. 

Vismia camparaguey Sprague & Riley, Kew Bull. 1924: 13. 
1924. Achotillo; achiotillo; cuxupche (Pete*n, Maya, fide Lundell); 
camparaguey; sangre de perro; lengua de chivo (fide Aguilar); cam- 
paraguay (Coban, Quecchi?). 

Moist or wet, open forest or brushy plains or hillsides, often in 
pine forest, frequently in second growth, 1,200 meters or less; Pete"n; 
Alta Verapaz; Izabal. Mexico (Chiapas) ; British Honduras; extend- 
ing farther southward. 

A large shrub or a tree, sometimes 10 meters high with a trunk 30 cm. in diam- 
eter, usually much smaller, the branches stellate-tomentose with ferruginous hairs; 
leaves short-petiolate, lanceolate to ovate-oblong, mostly 15-25 cm. long and 5- 
9 cm. wide but sometimes much larger on young plants, acuminate or long-acumi- 
nate, rarely subobtuse, rounded to shallowly cordate at the base, subcoriaceous, 
glabrous above, rather laxly ferruginous-tomentulose beneath, the pubescence 
coarse and not at all appressed, of branched hairs, the lateral nerves about 15 
pairs, stout and conspicuous; inflorescences terminal, thyrsiform, many-flowered, 
often 10 cm. long or larger, ferruginous-tomentose throughout, the flowers on ped- 
icels 5-8 mm. long; sepals ovate-lanceolate, obtuse, about 6.5 mm. long, ferrugi- 
nous-tomentose outside, glabrous within; petals white, oblanceolate, obtuse, 7 mm. 
long, many-nerved, glabrous outside, villous within; fascicles of stamens with 
numerous anthers, villous; ovary glabrous; fruit ovoid, 1 cm. long, tipped with 
the persistent styles, covered below by the persistent sepals. 

Known in British Honduras as "Old William," "can't-be-helped," 
and "yellow sangre." The explanation of one of these names is given 
as follows: "The wood is poor for building huts, but if nothing else is 
available this is used, as 'it can't be helped/ ' The wood is pinkish, 
moderately hard, strong, tough, medium-textured, fairly straight- 
grained, not difficult to work, not highly durable. Little use is made 
of it except as fuel. This has been reported from Guatemala as 
V. macrophylla HBK., V. ferruginea HBK., and V. latifolia Choisy. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 61 

Vismia mexicana Schlecht. Linnaea 10: 245. 1836. Campara- 
guey; camparaguay (Quecchi?) ; achiotillo. 

Moist or wet forest, usually in open pine forest, sometimes in 
thickets, pastures, or second growth, 900-1,450 meters; Pete"n; Alta 
Verapaz; Guatemala(?); Huehuetenango. Southern Mexico; British 
Honduras; Honduras, and southward. 

A shrub or small tree, the slender branches crowned with a close ferruginous 
tomentum; leaves slender-petiolate, ovate to lance-oblong or ovate-oblong, mostly 
12-19 cm. long and 5-7 cm. wide, acute or acuminate, rarely obtuse, at first more 
or less tomentulose on the upper surface but in age glabrous except along the costa, 
densely covered beneath with a minute, closely appressed tomentum, this brown 
or ferruginous at first, becoming pale or whitish in age; panicles small or large, 
many-flowered, densely ferruginous-tomentose, the flowers pedicellate; sepals 
about 5 mm. long, densely ferruginous- tomentulose; petals white, 7 mm. long, 
lineolate; phalanges of stamens bearing as many as 40 anthers, villous; fruit 
broadly ovoid, 10-12 mm. long, glabrous. 

Called "achiotillo" and "guayabon de montana" in Honduras; 
"tesuate Colorado" (Oaxaca). In Honduras this is often a tree of 
9-14 meters, but in Alta Verapaz, where it is abundant, it is usually 
much lower. The bark is thin, scaly, medium-brown in color, and 
exudes a small quantity of yellow-brown resin. Around Coban a 
decoction of the plant (perhaps a tincture of the resin) is employed 
as a mouth wash, to relieve toothache, and the leaves are bound on 
the temples to relieve headache. In Alta Verapaz this species is 
found at the higher levels, being replaced at lower elevations by 
V. camparaguey. This species has been confused with V. guianensis 
(Aubl.) Pers., from which it seems to be distinct, but it may be 
synonymous with some of the other South American species. 

GISTACEAE 

References: W. Grosser, Cistaceae, Pflanzenreich IV. 193. 1903. 
E. Janchen, Cistaceae, Nat. Pflanzenfam. ed. 2. 21: 289-313, ff. 1925. 

Herbs or shrubs; leaves alternate (in Guatemalan species) or opposite, with 
or without stipules; flowers regular, perfect, solitary, cymose-paniculate or cymose- 
racemose; sepals 5 or 3, the outer ones mostly shorter or narrower than the inner; 
petals 5, contorted, or 3 and imbricate, in cleistogamous flowers none; stamens 
numerous, unequal, rarely few, inserted on a hypogynous, usually disk-like recep- 
tacle; anthers laterally or subintrorsely dehiscent; ovary 1-celled and with parietal 
placentae, or incompletely 10-3-celled; ovules numerous or few on each placenta, 
commonly orthotropous, ascending or recurved; capsule loculicidal, rarely dehis- 
cent only at the apex; seeds with endosperm, the embryo curved; cotyledons folia- 
ceous, linear, subterete, or oblong. 

Five genera, of wide distribution, only two of them represented in 
Central America. 



62 FIELDIANA: BOTANY, VOLUME 24 

Petals 3, imbricate in bud; leaves in Guatemalan species linear, strigose. .Lechea. 

Petals 5, contorted in bud; leaves in Guatemalan species stellate-puberulent or 

pilose with spreading hairs Helianthemum. 

HELIANTHEMUM Adanson 

Plants usually perennial herbs, sometimes suffrutescent at the base, branched, 
erect or decumbent; flowers small or rather large and showy, yellow, mostly in 
terminal, simple or bifid racemes or cymes, the lowest often axillary; petals 5, 
broad; stamens numerous or rarely few; ovary with 3 placentae or false septa, the 
ovules numerous; style articulate with the ovary, filiform or subclavate, the stigma 
capitate or 3-lobate; embryo uncinate. 

About 100 species, widely distributed. Only the following are 
known in Central America. 

Stems and leaves pilose with spreading simple hairs H. chihuahuense, 

Stems and leaves minutely stellate-tomentulose. 

Flowers unequally pedicellate, the pedicels of the petaliferous flowers much 
longer than those of the cleistogamous flowers, the latter sessile or nearly so. 

H. glomeratum. 

Flowers all on pedicels of about the same length, the flowers often all petaliferous. 

H. Pringlei. 

Helianthemum chihuahuense Watson, Proc. Am. Acad. 23: 
268. 1888. 

Dry, open hillsides in the mountains, mostly in pine-oak forest, 
1,800-2,500 meters; Jalapa; Chimaltenango; El Quiche"; Huehuete- 
nango. Mexico; Honduras. 

Perennial from a hard woody root, erect or ascending, 30 cm. high or less, the 
stems few or numerous, simple or branched, hirsute; cauline leaves oblong-lanceo- 
late or spatulate-lanceolate, mostly 1-2 cm. long and 2-4 mm. wide, acute, sessile, 
green, pilose with white spreading hairs; leaves of the axillary branches smaller, 
often fasciculate; inflorescences with few-many flowers, the flowers partly peta- 
liferous, and partly cleistogamous, the two kinds on pedicels of equal length; outer 
sepals linear, the interior ones shorter, ovate, acuminate or mucronate, stellate- 
puberulent and pilose, in fruit 6 mm. long; petals yellow, twice as long as the sepals; 
capsule glabrous, ovoid, in the petaliferous flowers 5 mm. long, in the cleistogamous 
flowers 2.5-3 mm. long; seeds dark ferruginous, subpuberulent. 

The plant is not at all common in Guatemala, and usually of only 
sporadic occurrence. 

Helianthemum glomeratum Lagasca ex DC. Prodr. 1: 269. 
1824. Cistus glomeratus Lagasca, Gen. & Sp. Nov. 16. 1816. Halim- 
ium glomeratum Grosser, Pflanzenreich IV. 193: 47, /. 11. 1903. 

Mostly on open grassy or rocky hillsides, generally in oak or pine 
forest, sometimes growing on serpentine, 1,500-2,800 meters; Zacapa; 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 63 

Jalapa; Chimaltenango; Solola; Quiche^ Huehuetenango; Totonica- 
pan; Quezaltenango; San Marcos. Mexico; Costa Rica. 

A slender perennial from a woody, often thick root, the stems rather wiry, 
often numerous, branched or almost simple, usually erect and 40 cm. high or less, 
covered with a whitish or grayish tomentum of minute stellate hairs; leaves short- 
petiolate, lance-oblong or spatulate-lanceolate, 1-2.5 cm. long, 3-9 mm. wide, 
grayish-tomentulose on both surfaces; petaliferous inflorescences subumbellate and 
3-8-flowered, the pedicels 2.5 cm. long or less, the cleistogamous flowers sometimes 
mixed with the petaliferous ones, or often in small dense clusters; outer sepals of 
the petaliferous flowers linear, half as long as the inner, the latter oval, acuminate, 
densely stellate-tomentulose, in fruit 7-8 mm. long; inner sepals of the cleistoga- 
mous flowers 1-1.5 mm. long; capsule of the petaliferous flowers 6 mm. long, that 
of the cleistogamous flowers only 2 mm. long, glabrous; seeds brown, lustrous. 

All through the pine and oak forests of Chimaltenango, Quiche", 
and Huehuetenango, especially those regions closely cropped by 
sheep and cattle, this is one of the most abundant herbaceous plants, 
and during the dry winter months one may travel many miles with- 
out finding any other small plant in flower. Stock evidently does not 
eat it. The flowers are small for the genus, and not showy. 

Helianthemum Pringlei Watson, Proc. Am. Acad. 23: 268. 
1888. Halimium Pringlei Grosser, Pflanzenreich IV. 193: 46. 1903. 

Reported by Grosser as collected in the mountains of both Hue- 
huetenango and Quezaltenango, Seler 2905, 3286. Mexico. 

Plants slender, perennial, erect, 30-50 cm. tall, the stems few or several, 
mostly simple below, racemosely branched above, minutely stellate-tomentulose; 
leaves lanceolate or narrowly spatulate, 1.5-2.5 cm. long, 3-5 mm. wide, densely 
whitish-tomentulose, acute or subobtuse, attenuate to the base, ascending, short- 
petiolate or sessile; inflorescence lax, the flowers mostly solitary, the pedicels all 
long and slender; outer sepals of the petaliferous flower filiform, the inner ones 
twice as long, oval, in flower 6 mm. long, in fruit 7-8 mm.; petals yellow, twice 
as long as the sepals; cleistogamous flowers half as large as the petaliferous ones; 
capsule ovoid, acute, lustrous, in the petaliferous flowers 7 mm. long, in the cleis- 
togamous flowers 3-5 mm. long; seeds ferruginous, smooth. 

The species must be rare in Guatemala, for we have collected 
extensively in the two departments from which it is reported but we 
have not found it. In general appearance it is substantially differ- 
ent from H. glomeratum, especially in its more open inflorescence and 
taller, more strict stems. 

LECHEA L. 

Reference: Albion R. Hodgdon, Taxonomic study of Lechea, Rho- 
dora 40: 29-69, 87-131. 1938. 



64 FIELDIANA: BOTANY, VOLUME 24 

Perennial herbs, rarely suffrutescent at the base, erect or ascending; leaves 
small, alternate or falsely verticillate, ovate to subulate, without stipules; flowers 
very small, numerous; sepals 5, biseriate, the 2 outer ones narrow, the 3 inner 
broad, imbricate in bud; petals 3, dark red, usually shorter than the sepals, mar- 
cescent; stamens 3-25; ovary short-stipitate, the style very short or none, the 3 
stigmas dark red, fimbriate-plumose; placentae of the ovary 3, broad, each bearing 
2 erect subsessile ovules, one on each side of the posterior face; capsule 3-valvate; 
embryo almost straight or curved in the hard endosperm. 

Seventeen species are recognized in the latest monograph, all 
American, chiefly in eastern United States but ranging southward 
to West Indies and Honduras. 

Exterior sepals equaling or exceeding the interior ones L. tripetala. 

Exterior sepals shorter than the interior ones L. Torreyi var. congesta. 

Lechea Torreyi Leggett var. congesta Hodgdon, Rhodora 40: 
105. 1938. 

In savannas and pine lands. British Honduras (Peck 143). 
North Carolina to Mississippi and Florida. 

Stems simple or branching, 20-40 cm. tall; leaves linear to elliptic, acute, 
pilose on midrib and margin, sparingly so elsewhere, glabrous above, 5-20 mm. 
long and to 15 mm. broad; inflorescence a compact panicle, the crowded branches 
ascending or suberect; inner sepals obovate-elliptic, obtuse, about 2 mm. long, 
strongly ferruginous-pubescent, outer sepals shorter; seeds 3. 

The species is known in Central America from a single collection. 

Lechea tripetala (Mocino & Sesse") Britton, Bull. Torr. Bot. 
Club 21: 252. 1894. Helianthemum tripetalum Mocino & Sesse" ex 
Dunal in DC. Prodr. 1: 284. 1824. L. Skinneri Benth. Bot. Voy. 
Sulph. 66. 1844 (type collected in Guatemala by Skinner, the exact 
locality unknown). 

Brushy or grassy slopes, in oak and pine forest in the mountains, 
1,200-2,100 meters; Zacapa; Jalapa; Guatemala; Sacatape"quez; Chi- 
maltenango; Quiche"; Huehuetenango. Mexico; Honduras; Okla- 
homa, U.S.A. 

Plants decumbent or suberect, usually forming dense clumps of many wiry 
stems, 20 cm. tall or less, sparsely or densely strigose on the leaves and stems; 
leaves linear, mostly 6-12 mm. long and 1 mm. wide; flowers very small, usually 
numerous, on pedicels 2-3 mm. long; sepals 2 mm. long, the inner ones ovate, 
strongly carinate; capsule depressed-globose, 2-2.3 mm. broad; seeds usually 6, 
brown, smooth. 

The plant is an inconspicuous one of most ordinary appearance, 
abundant at many places in the pine-oak forests. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 65 

BIXACEAE. Arnotto Family 

Shrubs or trees with yellow or red sap; leaves alternate, simple, long-petiolate, 
membranaceous, entire, palmate-nerved; stipules in pairs, deciduous; flowers per- 
fect, large, in terminal panicles, the pedicels bearing 5 glands below the calyx; 
sepals 5, imbricate, caducous; petals 5, imbricate, fugacious; stamens indefinite, 
inserted on a thick receptacle, the anthers oblong, hippocrepiform with the 
branches united at the back, apically dehiscent by 2 short slits which finally be- 
come a single pore; ovary free, 1-celled, with 2 placentae, the ovules numerous; 
style simple, terminal, the stigma shortly bilobate; capsule bivalvate, densely cov- 
ered with long and rather soft spines, or often unarmed; seeds numerous, obovoid, 
the coat fleshy, containing a red or yellow sap; embryo axial, straight, the cotyle- 
dons broad, the radicle conic-cylindric. 

A single genus, with characters of the family. 



BIXA L. Arnotto 

An American group, consisting of at least two and perhaps three 
or more species, only one of which is found in North America. The 
numerous forms of this genus, at least those referred to B. Orellana, 
are highly variable in the form of the fruit, as well as in the size of 
the flowers. A critical monograph of the variations, based upon 
adequate material (which apparently has never been available in any 
single herbarium), is greatly needed. The generic name is said to be 
a modification of "bija," the Antillean name of the plant. 

Bixa Orellana L. Sp. PI. 512. 1753. Achiote; achiotillo (form 
with very small leaves and fruit); chaya; xayau (Quecchi); oox 
(Chuje); ox ( Jacaltenango) . 

Planted commonly in fincas, chiefly at 1,000 meters or less, occa- 
sionally grown at higher elevations; common in wet or dry thickets 
of the lowlands, and forming extensive, almost pure stands in many 
places, especially in the Pacific plains; almost or quite absent in some 
extensive regions; Pete"n; Alta Verapaz; Baja Verapaz; Izabal (mostly 
in cultivations); El Progreso; Zacapa (said to be rare); Chiquimula; 
Jutiapa; Santa Rosa; Escuintla; Sacatepe"quez; Quezaltenango; Chi- 
maltenango; Suchitepe"quez; Retalhuleu; San Marcos; doubtless found 
in cultivation in all or most of the departments. Mexico to British 
Honduras and Panama; West Indies; South America. 

A shrub or tree, commonly 2-8 meters high, in Guatemala sometimes 12 meters 
tall, usually with a dense rounded crown and a short trunk, the bark dark brown, 
smooth, the inner bark yellow; leaves thin, mostly persistent, on long slender 
petioles, ovate to broadly ovate or deltoid-ovate, mostly 8-20 cm. long, acumi- 
nate or long-acuminate, truncate or rounded at the base and 5-nerved, green and 



66 FIELDIANA: BOTANY, VOLUME 24 

glabrous above, paler beneath, usually rather densely but minutely lepidote; pani- 
cles rather small, few-many-flowered; sepals 12-14 mm. long, brown-lepidote, 
obtuse; petals pink or white, about 2.5 cm. long but variable in size, soon falling; 
capsule ovoid, acute or acutish, commonly 2.5-4.5 cm. long, densely covered with 
long or short, flexible spines, reddish-brown; seeds numerous, covered with abun- 
dant red-orange pulp. 

Numerous forms are found in Guatemala, differing in the shape 
and indument of the fruit and in the size of the petals. The one 
described above is the most widely dispersed in tropical America, 
at least in cultivation, and is the one said to yield the largest amount 
of dye material, although this is by no means certain. The most 
conspicuous variants of Guatemala and other parts of Central Amer- 
ica are the following: 

Bixa Orellana var. urucurana (Willd.) Kuntze ex Pilger, Pflan- 
zenreich ed. 2. 21: 315. 1925. B. Urucurana Willd. Enum. Hort. 
Berol. 565. 1809. Achiote. 

The usual wild form in Guatemala, although often seen in culti- 
vation; specimens seen from Pete'n, Retalhuleu, Jutiapa, Santa Rosa 
and Escuintla, and probably almost as widely distributed in Guate- 
mala as the typical form. Generally distributed through the range 
of the species. 

Characterized by the form of the fruit, which is globose or depressed-globose, 
variable in size, and densely covered with long flexible spines. 

Bixa Orellana var. leiocarpa (Kuntze) Standl. & L. Wms. 
Fieldiana, Bot. 29: 358. 1961. Orellana americana var. leiocarpa 
Kuntze, Rev. Gen. 45. 1891. Achiote. 

Frequent in cultivation in Guatemala; specimens seen from Petn, 
Santa Rosa, Alta Verapaz, Solold, and Sacatepquez, but probably 
found in all the warmer departments. Widely distributed with the 
species. British Honduras. 

Easily distinguished by the fruit, which is smooth and without bristles. 

Bixa is known in British Honduras by the name "atta." The 
Maya name of Yucatan is listed as "cuxul" and "cuxub" ("ciui" is 
the dye in small cakes as prepared for use). The name "achiote" (in 
some regions "achote") is derived from the Nahuatl name, "achiotl." 
In Guatemala this appears in such place names as Los Achiotes and 
El Achiotal (an achiote grove or thicket) on the Pacific coast. This 
tree is one of the best known and most useful ones of all tropical 
America because of the orange-red dye obtained from the dried seeds, 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 67 

or rather from their pulpy covering. Large amounts of it are used in 
Guatemala, and it is sold in all the markets, either the dried pods 
with the seeds or more usually the dried and powdered pulp, in the 
form of small cakes. Achiote is much used through Central America 
for coloring food, especially rice, also tortillas and other articles. In 
some regions apparently rice is not considered fit to eat unless it is 
so colored, but colored rice is not so common in Guatemala. Large 
amounts of arnotto dye are used in the United States and Europe 
for coloring butter and cheese, oils, and varnish, the supply coming 
principally from the West Indies and South America. It gives an 
orange-yellow color to silk and cotton, but the color is said to be 
fugitive. 

By the Indians of some regions arnotto was formerly much used 
and still is, in certain areas for painting the face and body, partly 
for ornament and partly to lessen attacks of mosquitoes and other 
biting insects. At an early date the plant was introduced into the 
Pacific islands, where the people soon learned to use it for the same 
purposes. The plant has been carried to most tropical regions of the 
Old World and has become naturalized in many of them. 

The wood is pinkish yellow or whitish, of about the consistency 
of basswood (Tilia). Some of the Indians used it to obtain fire by 
friction. A gum obtained from the branches is somewhat similar to 
gum arabic. The bark contains a strong fiber suitable for cordage. 

Being always at hand around the country houses, this plant is 
much used in domestic medicine. In Guatemala it is esteemed for 
treating sores and burns, to prevent the formation of scars. It is 
also a favorite remedy for dysentery, and in Yucatan is used as an 
antidote for poisoning by the seeds of Jatropha Curcas or Manihot. 
The specific name, Orellana, was given in honor of Francisco Orellana, 
associate of Pizarro in the conquest of Peru, and discoverer of the 
Amazon River. 

COGHLOSPERMACEAE 

Trees, shrubs, or herbs with colored sap; leaves alternate, usually long-petio- 
late, stipulate, palmately lobate or palmately compound; flowers perfect, large 
and showy, paniculate or racemose, regular; sepals 5, imbricate in bud, deciduous; 
petals 5, yellow, imbricate or subcontorted; stamens numerous, the filaments free, 
equal or unequal, the anthers linear, 2-celled, opening by terminal, short, often 
confluent slits; ovary superior, 1-celled and with 3 parietal placentae projecting 
into the cell, or perfectly 3-celled; ovules numerous; style simple, with a minutely 
denticulate stigma; fruit a 3-5-valvate capsule; seeds glabrous or covered with 
long hairs, straight or cochleate-reniform, the endosperm copious; embryo large, 
the cotyledons broad. 



68 FIELDIANA: BOTANY, VOLUME 24 

Three genera, in the tropics of both hemispheres. Two genera 
are known in America, both represented in Central America. 

Trees or shrubs; ovary 1-celled; filaments equal in length Cochlospermum. 

Herbs; ovary perfectly 3-celled; filaments unequal Amoreuxia. 

AMOREUXIA Mocifio & Sesse" 

Herbs with large tuberous roots; leaves membranaceous, palmately lobate, on 
very long petioles; flowers large and showy, orange, in few-flowered terminal pan- 
icles; sepals 5, imbricate, deciduous; petals 5, large, contorted-imbricate; stamens 
numerous, inserted on an eglandular torus, the filaments very unequal; anthers 
linear; ovary completely 3-celled, the placentae axial, many-ovulate; capsule locu- 
licidally 3-valvate, somewhat inflated, the endocarp membranaceous, separating 
into valves with the epicarp; seeds obovoid, straight or curved, glabrous or mi- 
nutely pubescent, the testa osseous. 

About six species, ranging from Arizona to Bolivia. Only the 
following is known from Central America. 

Amoreuxia palmatifida Mocifio & Sesse" ex DC. Prodr. 2: 638. 
1825. 

Damp thickets, sometimes in open grassy places, 800 meters or 
less; Zacapa; Huehuetenango. Mexico; Nicaragua; Arizona, U.S.A. 

An erect herb 40-60 cm. tall, the stems pale green, simple or branched, abun- 
dantly leafy, puberulent; stipules linear or filiform, deciduous; leaves on slender 
petioles 6-10 cm. long, the blades 4-8 cm. wide, rounded in outline, cordate at 
the base, deeply 5-7-lobate, the lobes thin, paler beneath, glabrous or nearly so, 
obovate, usually rounded at the apex, constricted below, coarsely crenate except 
near the base; sepals green, 1.5 cm. long, puberulent, ovate-lanceolate to linear- 
lanceolate, acuminate; petals bright orange, about 3 cm. long; capsule ovoid, 
3-4 cm. long; seeds turgid, black or dark brown, slightly curved, pubescent. 

A showy and handsome plant, well worthy of cultivation. It 
grows sparingly on brushy plains of the Zacapa Valley, but with 
foliage only during the wetter months, persisting by its large tuber- 
ous roots. In October, 1940, it was growing abundantly in wet places 
near the oil storage tank in the railroad yards at Zacapa. The Maya 
name is reported from Yucatan as "zacyab." 

COGHLOSPERMUM Kunth 

Reference: S. F. Blake, The American species of Maximilianea 
(Cochlospermum), Journ. Wash. Acad. Sci. 11: 125-132, /. 1-2. 1921. 

Trees or shrubs; leaves palmately lobate or digitately compound; flowers 
large, yellow, in axillary racemes or terminal panicles; sepals imbricate, decidu- 
ous; petals contorted-imbricate; stamens numerous, inserted on an eglandular 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 69 

torus, the anthers oblong or linear, sometimes apiculate; placentae of the ovary 
3-5, many-ovulate; capsule large, 3-5-valvate, the endocarp membranaceous; 
seeds cochleate-reniform or spirally twisted, lanate or hirsute, the testa corneous; 
embryo incurved, the cotyledons ovate. 

About 15 species, in tropical Asia, Africa, Australia, and America. 
Only one species occurs in North America. 

Cochlospermum vitifolium Willd. ex Spreng. Syst. 2: 596. 
1825. Bombax vitifolium Willd. Enum. Hort. Berol. 2: 720. 1809. 
C. hisbiscoides Kunth, Syn. PI. Aequin. 3: 214. 1824. Maximilianea 
vitifolia Krug & Urban, Bot. Jahrb. 15: 293. 1892. Tecomasuchil; 
tecomasuche; pumpunjuche; pumpumjuche; tecomajuche; comasuche; 
pochote (Pete*n) ; tsuyuy (Quecchi) ; cho (Pete"n, Maya) ; pomp, pumpo 
(Huehuetenango) ; tecomatillo (Zacapa) . 

Chiefly on dry brushy hillsides or plains, often in thin coastal 
forest, frequently in second growth, mostly at 1,000 meters or lower; 
Pete*n; Alta Verapaz; Baja Verapaz; Izabal; Zacapa; El Progreso; 
Jutiapa; Santa Rosa; Escuintla; Suchitepe"quez; Retalhuleu; San 
Marcos; Huehuetenango. Mexico; British Honduras to El Salva- 
dor and Panama; northern South America. 

A deciduous shrub or tree, sometimes 25 meters high but usually lower, often 
flowering when only 2 meters tall, the crown spreading, not very dense, the bark 
pinkish-brown, the inner bark yellow, the branchlets thick, dark reddish-brown, 
at first sparsely pilose; leaves on petioles 10-25 cm. long, usually 5-lobate for one- 
half to three-fourths their lengths, 10-30 cm. wide, green and glabrate above, 
somewhat paler beneath, pilose along the veins or glabrate, membranaceous, the 
lobes oblong to obovate-oval, acute or obtuse and usually abruptly short-pointed; 
inflorescences several-flowered, the pedicels densely puberulent, 2-3 cm. long; 
flowers 8-12 cm. broad, the sepals oblong-ovate to oval, 1-2 cm. long, rounded 
at the apex; capsule broadly obovoid-oval, 7-8 cm. long, grayish-tomentulose, 
5-valvate, umbilicate; seeds reniform, 4.5 mm. broad, densely white-lanate. 

Vernacular names reported in adjacent regions are "jicarillo," 
"berberia," "bombdn" (Honduras) ; "wild cotton" (British Honduras) ; 
"coquito" (Chiapas); "madera de pasta" (Yucatan); "chum," "choy," 
"chimu" (Yucatan, Maya) ; "pongolote," "palo de cuchara," "apompo" 
(Oaxaca and Veracruz) . The branches are brittle and can be easily 
broken from the tree. The wood is exceedingly soft and weak, and 
when green is a soft spongy mass, drying to a loose bundle of fibers. 
The branches root easily when placed in the ground, and are rarely 
planted in hedges. The large stamens are said to be used in Central 
America to adulterate or replace saffron, having considerable super- 
ficial resemblance to the stamens of that plant. The tree is employed 
commonly in domestic medicine of Guatemala in treating kidney and 



70 FIELDIANA: BOTANY, VOLUME 24 

liver diseases and to hasten childbirth. The tree is a conspicuous 
and handsome one when in flower, chiefly during the latter part of 
the dry season. The large flowers suggest bright yellow roses. The 
large capsules are soft and somewhat inflated, and can be crushed 
easily in the hand. The bark contains a tough fiber that is stated 
to be used sometimes for cordage. The name "tecomasuche" is of 
Nahuatl origin, signifying "trumpet-flower," a term of no very obvi- 
ous application. 

VIOLACEAE. Violet Family 

Reference: H. Melchior in Engler & Prantl, Nat. Pflanzenf. ed. 2. 
21: 329-377. 1925. 

Herbs or shrubs, sometimes trees, rarely scandent; leaves alternate or oppo- 
site, simple, entire or variously lobate or dentate; stipules small or foliaceous; 
flowers usually perfect, very irregular or regular, the inflorescence various, the flow- 
ers often solitary in the leaf axils; sepals 5, generally persistent, imbricate; petals 5, 
imbricate, hypogynous or somewhat perigynous, in some genera very unequal, the 
lowest petal larger than the others or different in form, often spurred at the base; 
stamens 5, hypogynous or somewhat perigynous; anthers erect, connivent about 
the ovary and sometimes united, sessile or subsessile, the connective often pro- 
duced beyond the cells into a membranous appendage, the cells opening by a 
longitudinal slit; ovary free, sessile, 1-celled, the placentae usually 3, parietal; 
style simple; ovules usually numerous on each placenta, anatropous; fruit capsu- 
lar and opening by as many valves as there are placentae, or sometimes baccate; 
testa of the seed membranaceous or coriaceous, the endosperm usually carnose 
and copious; embryo axial, usually straight, the cotyledons flat. 

Genera about 19, widely distributed in almost all tropical and 
temperate regions. Two other genera are known from southern 
Central America. 

Petals unequal, the lowest one different from the others, often calcarate at the base. 

Sepals produced at the base below the point of attachment; herbs Viola. 

Sepals not produced at the base. 

Plants scandent or shrubs; lower petal with a long basal spur longer than the 

blade of the petal Corynostylis. 

Plants not scandent, often herbaceous; lower petal not calcarate, sometimes 

short-saccate. 

Lowest petal conspicuously unguiculate; herbs or shrubs; flowers solitary 
in the leaf axils or in small, short-pedunculate lateral cymes. 

Hybanthus. 

Lowest petal sessile or nearly so; trees; flowers in large, long-pedunculate 
cymes grouped and subumbellate near the ends of the branches. 

Orthion. 

Petals subequal, not calcarate at the base; trees or shrubs Rinorea. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 71 

CORYNOSTYLIS Martius 

Woody vines; leaves alternate, petiolate, dentate or almost entire; flowers 
large, very irregular, axiliary, long-pedunculate, the pedicels 2-bracteolate; sepals 
small, subequal, not produced at the base; lower petal produced at the base into 
a long spur, the blade small, the other petals smaller; filaments very short, 2 or 4 
of the lowerjones'shortly villous, calcarate dorsally, the connective produced at 
the apex into'a membrane j'style clavate, the stigma anterior; fruit capsular, coria- 
ceous, large,*opening"by 3 valves,*these not elastic; seeds compressed and flat, 
suborbicular, r not*winged, r the testa'crustaceous, rugose; endosperm thin. 

Three or four species, in tropical America. Only the following is 
known in North America. 

Corynostylis arborea (L.) Blake in Standl. Contr. U. S. Nat. 
Herb. 23: 837. 1923. Viola arborea L. Syst. Nat. ed. 10. 1239. 1759. 
C. Hybanthus Mart. Nov. Gen. & Sp. 1: 26. 1823. 

Wet forest or thickets, at or little above sea level; Pete"n; Izabal. 
Southern Mexico; British Honduras; Honduras; Costa Rica; Pan- 
ama; West Indies; South America. 

A woody vine, sometimes 9 meters long, glabrous throughout or nearly so; 
leaves on rather short petioles, firm-membranaceous, oval to elliptic-ovate, 5- 
11 cm. long, acute or acuminate, crenulate or almost entire, prominently reticulate- 
veined; flowers large, white, solitary in the upper axils or forming short terminal 
racemes, the slender pedicels 3-4 cm. long, bracteate above the middle; spur of the 
corolla sometimes 2.5 cm. long, somewhat twisted, the blade of the spurred petal 
about 1 cm. long; capsule oval, 5.5 cm. long, 3.5 cm. broad, almost ligneous; 
seeds very numerous and densely packed, strongly compressed, irregular in out- 
line, brown, lustrous, about 12 mm. in greatest diameter. 

Called "monkey apple" in British Honduras. In South America 
the plant has been used as a remedy for fevers. 

HYBANTHUS Jacquin 

Reference: C. V. Morton, The genus Hybanthus in continental 
North America, Contr. U. S. Nat. Herb. 29: 74-82. 1944. 

Shrubs or herbs, sometimes with spinose branches; leaves alternate or rarely 
opposite; flowers mostly small and inconspicuous but sometimes rather large and 
showy, the peduncles axillary or in terminal racemes, solitary or fasciculate; sepals 
somewhat unequal, not produced at the base; lowest petal slightly or much larger 
than the others, unguiculate, the claw gibbous or subsaccate at the base; anthers 
subsessile or on distinct filaments, free or connate, the connective produced at the 
apex into a membrane; style incurved-clavate at the apex, the stigma anterior; 
capsule elastically 3-valvate; seeds ovoid-globose, the testa crustaceous. 

About 75 species, in the tropics of both hemispheres but most 
numerous in America. A few other species occur in southern Cen- 
tral America. 



72 FIELDIANA: BOTANY, VOLUME 24 

Flowers in cymes; shrub with often spinose branches H. yucatanensis. 

Flowers solitary or fasciculate in the leaf axils; plants herbaceous or woody, the 

branches never spinose. 
Calyx large, almost 1 cm. long, bearing numerous fleshy green appendages, 

densely long-pilose H. Calceolaria. 

Calyx much smaller, not appendaged, never long-pilose. 
Leaf blades mostly rounded or almost truncate at the base. 

Leaves, at least the upper ones, linear; stems glabrous. . . .H. oppositifolius. 

Leaves mostly oblong-ovate; stems puberulent H. Galeottii. 

Leaf blades all or chiefly acute to attenuate at the base. 
Plants annual; at least the lower leaves and branches opposite. 

H. attenuatus. 

Plants perennial, often woody below; leaves and branches alternate. 
Capsule about 4 mm. long; pedicels filiform; leaves mostly obtuse. 

H. Thiemei. 

Capsule 7-10 mm. long or even larger; pedicels not or scarcely filiform; 
leaves acute or acuminate. 

Plants herbaceous, or suffrutescent at the base, usually less than 40 cm. 
high; leaves mostly 3-5 cm. long, acute; branches densely puber- 
ulent or short-pilose H. verbenaceus. 

Plants woody, usually a meter high or even taller; leaves mostly 7- 
15 cm. long; branches glabrous or nearly so. 

Capsule 10-12 mm. long; pedicels minutely puberulent; leaves con- 
spicuously serrate H. elatus. 

Capsule about 7 mm. long; pedicels glabrous; leaves very obscurely 
crenate-serrulate H. sylvicola. 

Hybanthus attenuatus (Humb. & Bonpl.) G. K. Schulze, 
Notizbl. Bot. Gart. Berlin 12: 114. 1934. lonidium attenuatum 
Humb. & Bonpl. in Roem. & Schult. Syst. Veg. 5: 402. 1819. /. ri- 
parium HBK. Nov. Gen. & Sp. 5: 378. 1821. lonidium oppositi- 
folium and Hybanthus oppositifolius of authors, not Viola oppositifolia 
L. I. parietariaefolium DC. Prodr. 1: 308. 1824. 

In moist soil, in thickets or open ground, often a weed in waste 
or cultivated places, 1,800 meters or less; Izabal; Alta Verapaz; 
Zacapa; Jutiapa; Guatemala; Suchitepe"quez; Retalhuleu; Quezalte- 
nango; San Marcos; Huehuetenango; Escuintla. Mexico; El Salva- 
dor and Honduras to Panama; South America. 

An erect annual, commonly 30-50 cm. high but sometimes taller, often much 
branched, the stems puberulent or short-pilose; lower leaves and branches oppo- 
site, the upper ones alternate, short-petiolate, elliptic-lanceolate to ovate, 2-7 cm. 
long, acuminate or long-acuminate, acute at the base, serrate or crenate, incon- 
spicuously scaberulous or puberulent above, sparsely puberulent beneath or almost 
glabrous; flowers small and inconspicuous, axillary, white, the pedicels slender, 
often 2 cm. long; sepals lanceolate, 2.5 mm. long; lowest petal 8 mm. long; capsule 
3-4 mm. long; seeds ochraceous, subglobose, lustrous. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 73 

A common, often weedy species in the highlands of Central Amer- 
ica. The species is quite variable, especially in the leaves, which may 
be elliptic-lanceolate to ovate and from quite small to fairly large. 
Called "hierba del rosario" in El Salvador. 

Hybanthus Calceolaria (L.) G. K. Schulze, Notizbl. Bot. Gart. 
Berlin 12: 114. 1934. Viola Calceolaria L. Sp. PL ed. 2. 1327. 1763. 
V. Ipecacuanha L. Mant. PI. 484. 1771. H. Ipecacuanha Baill. Bot. 
Me~dic. 2: 841. 1884. 

British Honduras (Big Fall Pine Ridge, Lundell 4230). Mexico 
(Chiapas and Tabasco) ; South America. 

An erect perennial, usually 40 cm. high or less, simple or branched, herbaceous 
or somewhat frutescent below, densely or sparsely pilose throughout with long, 
pale, spreading or subappressed hairs; leaves alternate, short-petiolate or sessile, 
ovate to lanceolate or oblanceolate-oblong, mostly 2-4 cm. long, obtuse or sub- 
acute, narrowed to the base, serrate; peduncles axillary, solitary, shorter than the 
leaves; sepals about 1 cm. long, lanceolate, acute, densely villous, the margins 
pectinate-ciliate with green fleshy lobes; petals white or pale blue, the lowest one 
2-2.5 cm. long, villous dorsally; capsule ovoid, scarcely equaling the sepals; seeds 
blackish at maturity. 

The plant is well known in Brazil under the names "poaya," 
"poaya branca," and "purga do campo." There it has been used as 
a substitute for true ipecac, and has been exported in quantity to 
Europe for the same purpose. 

Hybanthus elatus (Turcz.) Morton, Contr. U. S. Nat. Herb. 
29: 80. 1944. lonidium elatum Turcz. Bull. Soc. Nat. Moscou 36, 
pt. 1: 556. 1863. Calceolaria glabra Dowell, Bull. Torr. Bot. Club 33: 
552, t. 20. 1906. C. brevis Dowell, I.e. H. glaber Standl. Journ. Wash. 
Acad. Sci. 17: 168. 1927. 

Dense, moist or wet, mixed forest, 900-2,000 meters; Escuintla; 
Santa Rosa; Chimaltenango; Alta Verapaz; Suchitepe"quez; Huehue- 
tenango; Quezaltenango; San Marcos. Southern Mexico. 

An erect shrub 1-3 meters high, often much branched, the branches rather 
stiff and slender, glabrous or sparsely puberulent; leaves alternate, short-petiolate, 
membranaceous, chiefly oblanceolate and 7-15 cm. long, long-acuminate, long- 
attenuate to the base, crenulate, glabrous or nearly so, sometimes finely puberu- 
lent; stipules linear-lanceolate, 1-3 mm. long; peduncles axillary, 5-10 mm. long; 
sepals linear-lanceolate, 4 mm. long, ciliate; petals white, the lowest one 1 cm. 
long; capsule 10-12 mm. long, ovoid, acute or acuminate, glabrous; seeds ochra- 
ceous or brown, subglobose, lustrous, smooth. 

Hybanthus Galeottii (Turcz.) Morton, Fieldiana, Bot. 29: 358. 
1961. lonidium Galeottii Turcz. Bull. Soc. Nat. Moscou 27 (2) : 339. 



74 FIELDIANA: BOTANY, VOLUME 24 

1854. I. occultum Polak. Linnaea 41: 548. 1877 (type from Cerro 
La Carpintera, Costa Rica). Calceolaria nigricans Dowell, Bull. Torr. 
Bot. Club 33: 554. 1906 (type from San Pedro Sula, Honduras). 
H. nigricans Standl. Journ. Wash. Acad. Sci. 17: 169. 1927. H. oc- 
cultus Standl. I.e. 18: 714. 1937. 

Moist forest, 800-1,200 meters; Alta Verapaz; Huehuetenango. 
Southern Mexico; Honduras; Nicaragua; Costa Rica. 

A slender shrub, sometimes a meter high or more, the branches puberulent; 
leaves alternate, oblong-ovate, 3-7 cm. long, acute or subacute, rounded or trun- 
cate at the base, often darkening when dried, serrulate, sparsely puberulent or 
almost glabrous; petioles slender, 4-12 mm. long; flowers white, solitary in the 
leaf axils, the peduncles slender, mostly 1.5-2 cm. long; sepals lanceolate, 4 mm. 
long, glabrous; lowest petal 9 mm. long; capsule ovoid, short-rostrate, glabrous. 

Hybanthus oppositifolius (L.) Taub. in Engler & Prantl, Pflan- 
zenfam. 3, pt. 6: 333. 1895. Viola oppositifolia L. Sp. PI. ed. 2. 1327. 
1763. lonidium angustifolium HBK. Nov. Gen. & Sp. 5: 377. 1821. 
H. angustifolius Standl. Journ. Wash. Acad. Sci. 17: 168. 1927. 

Reported from Pete"n and Alta Verapaz (Cubilgiiitz). British 
Honduras; southern Mexico; Honduras. 

Plants perennial, erect, herbaceous, mostly 50 cm. high or less, the stems very 
slender, terete, glabrous; leaves all opposite, the upper ones linear, the lower often 
linear-lanceolate, 3-7 cm. long, sessile, long-attenuate, rounded at the base, entire 
or very obscurely serrulate, green above and scaberulous or glabrate, paler beneath 
and glabrous; flowers on long slender pedicels in the upper axils, white; sepals gla- 
brous, lanceolate, subulate-attenuate; lowest petal about 12 mm. long; capsule 
6 mm. long, glabrous; seeds globose, black, very lustrous. 

Hybanthus sylvicola Standl. & Steyerm. Field Mus. Bot. 23: 
176. 1944 (type from Guatemala, Wilson 329). 
Alta Verapaz. Endemic. 

A shrub or small tree of 3-4.5 meters, glabrous throughout, the branches 
slender, pale green, subterete or obtusely trigonous, woody almost to the ends; 
stipules narrowly ovate, scarcely 1.5 mm. long, pale; leaves on petioles 3-4 mm. 
long, oblanceolate or oblong-oblanceolate, 8.5-10 cm. long, 2.5-3.5 cm. wide, acute 
or short-acuminate, gradually cuneate-attenuate to the base, obsoletely and re- 
motely crenate-serrate or almost entire, paler beneath; flowers subfasciculate from 
axillary buds, few, the peduncles almost filiform, 5-8 mm. long; sepals glabrous, 
very unequal, lanceolate or broadly lanceolate, acuminate; lowest petal 8 mm. 
long, its limb rhombic, the claw almost equaling the limb, the other petals almost 
equaling the claw; capsule ovoid-globose, 7 mm. long and broad, very obtuse, 
scarcely rostrate, glabrous. 

Hybanthus Thiemei (Donn.-Sm.) Morton, Contr. U. S. Nat. 
Herb. 29: 81. 1944. lonidium Thiemei Donn.-Sm. Bot. Gaz. 40: 1. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 75 

1905 (type from San Pedro Sula, Honduras). Hierba de pesar (Pete"n, 
fide Lundell). 

Mostly in moist open places, sometimes on open banks, 300 me- 
ters or less; Pete"n; Huehuetenango; Santa Rosa. Southern Mexico; 
El Salvador; Honduras; Costa Rica; Panama. 

Plants erect or suberect, perennial, commonly 6-12 cm. high, simple or sparsely 
branched, densely leafy, the stems puberulent; leaves alternate, on rather long, 
very slender petioles, ovate to oblong-elliptic, 2-6 cm. long, very obtuse to sub- 
acute, cuneately narrowed at the base, thin, glabrous or nearly so, paler beneath, 
crenate; peduncles filiform, 1.5-2.5 cm. long; sepals 2.5-3 mm. long; petals white, 
the lowest 1 cm. long; capsule ovoid, 4 mm. long; seeds minutely pitted, ochraceous. 

The Maya name of Yucatan is reported as "xcamuc olal." 

Hybanthus verbenaceus (HBK.) Loes. Bull. Herb. Boiss. II. 
3: 215. 1903. lonidium verbenaceum HBK. Nov. Gen. & Sp. 5: 379, 
t. Jf.97. 1821. Palomilla (Huehuetenango). 

Moist thickets or damp shaded banks, sometimes in pine-oak 
forest, 1,300-2,000 meters; Alta Verapaz; Jalapa; Jutiapa; Santa 
Rosa (type from Volcan de Jumaytepeque, Heyde & Lux 3943); 
Sacatepe'quez; Escuintla; Chimaltenango; Quiche"; Huehuetenango. 
El Salvador. 

Plants perennial, erect or decumbent, 30-50 cm. high, often much branched, 
the stems densely puberulent or short-pilose in two lines; leaves alternate, short- 
petiolate, broadly ovate to elliptic, mostly 2-5 cm. long, acute at each end, crenate, 
minutely hispidulous on both surfaces or finally glabrate, paler beneath; flowers 
axillary, white, the slender pedicels 10-12 mm. long; sepals lanceolate, acuminate, 
ciliate, 3 mm. long; lowest petal 6 mm. long; capsule ovoid, glabrous, 1 cm. long, 
acuminate; seeds subglobose, ochraceous or mottled with light and dark brown. 

Hybanthus yucatanensis Millsp. Field Mus. Bot. 1: 404. 1898. 
H. cymosus Bartlett, Proc. Am. Acad. 43: 56. 1907 (type from Gua- 
lan, Zacapa, C. C. Deam 385). 

In thickets, often in second growth, 600 meters or less; Pete"n; 
Alta Verapaz; Zacapa. Yucatan peninsula of Mexico. 

A shrub 1.5-2.5 meters high, the branches often short and spinose, conspicu- 
ously angulate, the angles wing-like, glabrous, the older branches whitish; leaves 
alternate, short-petiolate, ovate or oblong-oblanceolate, 2.5-6.5 cm. long, narrowed 
to an obtuse apex, cuneate or long-attenuate at the base, crenulate, glabrous; 
flowers small, greenish white, in few-many-flowered axillary cymes, these usually 
umbelliform, short-pedunculate, the pedicels mostly 3-4 mm. long, glabrous; 
sepals minute, obtuse; lowest petal 4 mm. long; capsule subglobose, 7 mm. long, 
glabrous. 

The Maya name of Yucatan is "sac-bacel-can," and the name "ta" 
also is reported. 



76 FIELDIANA: BOTANY, VOLUME 24 

ORTHION Standley & Steyermark 

Glabrous trees, the stipules short and broad, caducous; leaves alternate or 
congested at the ends of the branches, and subverticillate, chartaceous, narrow, 
short-petiolate or almost sessile, appressed-serrate or undulate; inflorescences axil- 
lary, mostly arising in the upper leaf axils and falsely umbellate, the flowers in 
lax, many-flowered, much-branched, corymbiform cymes, sessile or short-pedicel- 
late, white, the bracts minute; sepals small, subequal, rounded or oval, rounded 
to acute at the apex, not produced at the base; lowest petal slightly larger than the 
others, puberulent outside, sessile, sometimes slightly dilated at the base, retuse 
at the apex; stamens glabrous, the filaments complanate, broad, connate into a 
short tube; anthers oval, the broad connective produced at the apex into a short 
rounded appendage; style stout, subsigmoid-curved; capsule rather large, obtusely 
trigonous, elastically 3-valvate, thick-coriaceous, obtuse or rounded at the apex; 
placentae 3, the ovules numerous, the funicles short; seeds by abortion usually 6, 
sometimes 3, subglobose, smooth. 

Only the following not very distinctive species are known. 

Leaves sessile or nearly so, mostly 15-27 cm. long, narrowly rounded at the base. 

O. subsessile. 
Leaves petiolate, mostly 6-22 cm. long, acute at the base. 

Leaves lanceolate, long-attenuate at the apex O. malpighiifolium. 

Leaves oblanceolate, abruptly acuminate 0. oblanceolatum. 

Orthion malpighiifolium (Standl.) Standl. & Steyerm. Field 
Mus. Bot. 22: 250. 1940. Hybanthus malpighiifolius Standl. Carne- 
gie Inst. Wash. Publ. 461: 73. 1935. 

Known only from the type W. A. Schipp 1278, collected in forest, 
Camp 32 on the Guatemala (Pet^n) -British Honduras boundary, 
630 meters. 

A glabrous tree 10 meters high, the trunk 15 cm. in diameter, the branches 
slender, green when young; leaves on slender petioles 3-10 mm. long, narrowly 
lanceolate or oblong-lanceolate, 6-14 cm. long, 1.5-4.5 cm. wide, narrowly long- 
attenuate, acute or attenuate-decurrent at the base, remotely and inconspicuously 
appressed-serrate, the nerves and veins prominent and reticulate on both surfaces; 
cymes lax and many-flowered, long-pedunculate, 1.5-2.5 cm. long and broad, the 
peduncles angulate or complanate, 3-4.5 cm. long, the pedicels 2 mm. long or less; 
sepals broadly ovate, acute or obtuse, 1 mm. long; lowest petal longer than the 
others, 2 mm. long, puberulent outside, slightly dilated at the base; immature 
capsule globose, 4 mm. long, apiculate. 

Orthion oblanceolatum Lundell, Lloydia 4: 54. 1941. 

Moist or wet, mixed, lowland forest, 300 meters or lower; Pete"n 
(type from Guatemala, Mercedes Aguilar H. 497); Alta Verapaz; 
Izabal; Huehuetenango. Nicaragua. 

A glabrous tree 9-15 meters high, the young branches green, terete; leaves 
chartaceous, on petioles 4-9 mm. long, oblanceolate or oblong-oblanceolate, 7.5- 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 77 

23 cm. long, 2.5-7 cm. wide, rather abruptly acuminate, acute at the base, re- 
motely appressed-serrulate, prominently reticulate- veined; inflorescences crowded 
at the ends of the branches, long-pedunculate, as much as 13 cm. long, the pe- 
duncles and branches complanate, the pedicels in fruit to 6 mm. long, articulate 
below the middle; sepals unequal, broadly ovate or suborbicular, 2 mm. long, apic- 
ulate, minutely erose; petals white, 2.5 mm. long, rounded at the apex; capsule 
obtusely trigonous, suborbicular, 1.3 cm. long, emarginate; seeds 3-5, globose, 
4 mm. in diameter. 

This species possibly is not distinct from the preceding one. 

Orthion subsessile (Standl.) Steyerm. & Standl. Field Mus. Bot. 
22: 250. 1940. Hybanthus subsessilis Standl. Carnegie Inst. Wash. 
Publ. 461:72. 1935. 

Wet, mixed forest, 600 meters or less; Alta Verapaz; Izabal (type 
from Guatemala, Johnson 1071). British Honduras. 

A glabrous tree 6-18 meters high, the trunk as much as 45 cm. in diameter; 
stipules broadly rounded, mucronate, small, caducous; leaves large, alternate or 
subopposite, chartaceous, subsessile or on thick petioles 5 mm. long, narrowly 
oblong or oblanceolate-oblong, 12-34 cm. long, 4-11 cm. wide, acuminate or acute, 
sometimes obtuse, gradually narrowed to the narrowly rounded or obtuse, often 
unequal base, remotely undulate-crenate, the veins prominently reticulate on both 
surfaces; inflorescences cymose, long-pedunculate, subumbellate at the ends of 
the branches, the peduncles 7-11 cm. long, complanate, straight or slightly in- 
curved, lax and many-flowered, 5-6 cm. long and broad, the pedicels 2 mm. long 
or less; sepals rounded, pale, scarcely 1 mm. long; lowest petal 2 mm. long, puber- 
ulent outside, sessile, scarcely dilated at the base, white, the limb broad and deeply 
retuse; anthers oval, with a small rounded appendage at the apex; capsule obtusely 
trigonous, 1 cm. long, green, broadly rounded at the apex, usually 3-seeded; seeds 
subglobose, 5 mm. in diameter, ochraceous, smooth. 



RINOREA Aublet 

Reference: S. F. Blake, Revision of the American species of Ri- 
norea, Contr. U. S. Nat. Herb. 20: 491-518, it. 31-37. 1924. 

Trees or shrubs; leaves alternate or opposite, entire or serrate; flowers small, 
5-parted, solitary or usually racemose or paniculate, axillary or terminal; sepals 
subequal; petals subequal, sessile or short-unguiculate; filaments free or connate, 
dorsally appendaged or naked, the connective produced beyond the cells; placentae 
1-many-ovulate; style straight, the stigma terminal; fruit capsular, the 3 valves 
elastic or rigid; seeds few, subglobose, glabrous or pubescent, the testa crustaceous 
or coriaceous. 

About 250 species, in the tropics of both hemispheres. A few 
additional ones occur in southern Central America. 

Flowers in small panicles R. Hummelii. 

Flowers in racemes. 



78 FIELDIANA: BOTANY, VOLUME 24 

Leaves broadest above the middle, narrowed to the base, the base itself narrowly 
rounded; anthers not appendaged R. deflexiflora. 

Leaves broadest at the middle, acute at the base; anthers usually appendaged 
at the apex R. guatemalensis. 

Rinorea deflexiflora Bartlett, Proc. Am. Acad. 43: 56. 1907. 

Wet forest, at or near sea level; Izabal (type from Livingston, 
C. C. Deam 61). British Honduras. 

A shrub or tree, sometimes 6 meters high, the branches slender, sparsely pilos- 
ulous or glabrate; leaves opposite or ternate, on petioles 2-5 mm. long, obovate or 
obovate-oblong, 10-23 cm. long, 4-8 cm. wide, acuminate, often abruptly so, 
gradually attenuate to the narrowly rounded or subcordate base, remotely ser- 
rulate, firm-membranaceous, puberulent above on the costa, barbate beneath in 
the axils of the nerves; flowers small, racemose, creamy white, the racemes 4-6 cm. 
long, the pedicels 4-6 mm. long; sepals ovate or oblong-ovate, 1.8-2.5 mm. long, 
ciliolate; petals oblong-ovate, 4.2-4.8 mm. long, obtuse; anthers not appendaged; 
capsule 1.5-2 cm. long, glabrous. 

Called "wild coffee" in British Honduras. 

Rinorea guatemalensis (Wats.) Bartlett, Proc. Am. Acad. 43: 
56. 1907. Alsodeia guatemalensis Wats. Proc. Am. Acad. 21: 458. 
1886. Cuolcuol (Alta Verapaz). 

Mostly in wet, mixed forest, sometimes in wet thickets or second 
growth, often on limestone, 500 meters or less; Pete"n; Alta Verapaz; 
Izabal (type from Rio Chocon, Serena Watson 15). Mexico; British 
Honduras; El Salvador; Honduras. 

A shrub or small tree, 10 meters high or less, the bark very thin, flaking off 
irregularly, the branchlets densely puberulent at first; leaves opposite, on petioles 
4-9 mm. long, mostly elliptic, sometimes oval or elliptic-oblong, 7-15 cm. long, 
2.5-6 cm. wide, acute or short-acuminate, acute at the base, rather thick, sub- 
entire, sparsely strigose or pilosulous beneath along the costa; racemes 3-6.5 cm. 
long, densely pilosulous or puberulent, the pedicels 2-4 mm. long; sepals ovate or 
deltoid-ovate, 1-1.5 mm. long, acute or subacute, ciliolate; petals lance-ovate, 
5.5 mm. long, cream-colored; anthers bearing 1-2 cusps at the apex or unappend- 
aged; ovary densely pilosulous; capsule puberulent, 1.5-2 cm. long; seeds globose, 
6 mm. in diameter, fuscous, sparsely puberulent. 

Known in British Honduras as "cafecillo" or "wild coffee;" "Cos- 
tarrica," "frutillo" (Oaxaca). The sapwood is cream-colored, the 
heartwood thin, pinkish-brown. 

Rinorea Hummelii Sprague, Kew Bull. 307. 1921. 
Moist or wet, mixed forest or thickets, 300 meters or less; Pete"n; 
Izabal. Mexico; British Honduras (type from Salt Creek, Hummel). 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 79 

A shrub or small tree, 10 meters high or less, the young branchlets minutely 
pilosulous, the trunk up to 20 cm. in diameter; leaves opposite, on petioles 3-5 mm. 
long, obovate to elliptic or oblong-elliptic, 6-15 cm. long, 2.5-6 cm. wide, abruptly 
short-acuminate, acute at the base, obscurely serrulate, chartaceous, glabrous 
above except for the puberulent costa, glabrous beneath; flowers paniculate, 
cream-colored, the panicles racemiform, terminal and axillary, 1.5-5 cm. long, 
puberulent, the lower branches short and few-flowered, the pedicels 2 mm. long; 
sepals broadly ovate, 1-1.2 mm. long, short-cuspidate or apiculate, ciliolate; petals 
lanceolate, 4.5 mm. long, obtuse; ovary fulvous-velutinous; capsule minutely 
brownish-pilosulous or glabrate, 1.5 cm. long. 

Called "wild coffee" in British Honduras. 



VIOLA L. Violet 

Reference: Wilhelm Becker, Violae mexicanae et Centrali-Ameri- 
canae, Repert. Sp. Nov. 19: 392. 1924; 20: 1. 1924. 

Annual or perennial herbs; leaves alternate or often apparently all basal, the 
stipules usually foliaceous and conspicuous; peduncles axillary, 1-flowered; flowers 
often dimorphous, the earliest perfect and usually sterile, the later ones small, 
apetalous, and fertile; sepals subequal, produced at the base below the point of 
attachment; petals spreading, the lowest usually larger, calcarate or saccate at 
the base; anthers subsessile, the connective complanate, produced at the apex as 
a membranaceous appendage, the connective of the two lower anthers usually 
calcarate dorsally at the base; style usually capitate, clavate, or variously dilated, 
almost straight, the stigma terminal, or more or less recurved; capsule elastically 
3-valvate; seeds ovoid-globose, the testa crustaceous, often lustrous. 

A vast genus of perhaps 400 species, generally distributed, chiefly 
in temperate regions; in tropical America the species are relatively 
few and confined to mountain regions. One other species is known 
from southern Central America. 

Plants without rootstocks, the main stems elongate, sometimes scandent. 

Stipules pinnatifid; peduncles much longer than the leaves V. tricolor. 

Stipules not lobate; peduncles all or mostly shorter than the leaves . . V. scandens. 
Plants with usually thick rootstocks, never scandent, at first acaulescent but 
usually producing elongate creeping stolons. 

Plants cultivated, introduced species V. odorata. 

Plants native species, not in cultivation. 
Leaf blades cordate-ovate, most of them distinctly longer than broad. 

V. reptans. 
Leaf blades mostly rounded-reniform, as broad as long or broader. 

Leaves not at all pointed at the apex; stolons none or poorly developed. 

V. Nannei. 
Leaves usually conspicuously pointed at the apex; stolons usually well 

developed. 
Leaves glabrous or nearly so; stipules 1.5 cm. long, long-fimbriate. 

V. Seleriana. 

Leaves usually conspicuously hispidulous beneath; stipules 1 cm. long or 
shorter, sparsely short-fimbriate or almost entire. .V. guatemalensis. 



80 FIELDIANA: BOTANY, VOLUME 24 

Viola guatemalensis Becker, Repert. Sp. Nov. 20: 6. 1924. 
Violeta. 

Moist open banks or more often in dense moist forest, often in 
forests of pine and oak or of Abies or Cupressus, 1,800-4,000 meters; 
Jalapa; Guatemala; Sacatepe"quez (type from Volcan de Agua, 2,850 
meters, J. D. Smith 2165); Chimaltenango; Sacatepe"quez; Quiche"; 
Quezaltenango; San Marcos; Huehuetenango. Southern Mexico; 
Costa Rica; Panama. 

Plants acaulescent, arising from a thick or slender, fleshy rootstock, develop- 
ing stolons, these slender, elongate, 10-20 cm. long or more, the stems glabrous or 
nearly so, the stolons creeping and rooting at the nodes; stipules ovate or lanceo- 
late, 1 cm. long or shorter, acute, fimbriate or subentire, thin, pale green; leaves 
long-petiolate, rounded-reniform, thin, broadly rounded at the apex and usually 
abruptly short-pointed, deeply cordate at the base, 2-3.5 cm. long and equally 
broad or broader, conspicuously crenate, sparsely hispidulous, especially beneath; 
petioles equaling or exceeding the blades, hispidulous or sometimes quite glabrous; 
flowers usually longer than the leaves, pale bluish or lavender, 1 cm. long, the 
peduncles glabrous; sepals ovate or ovate-lanceolate, glabrous, the basal append- 
ages short and rounded; style subclavate toward the apex. 

This has been reported from Guatemala as V. Hookeriana HBK. 
The forma pubescentior Becker (op. cit. 7; type from Todos Santos, 
Huehuetenango, C. & E. Seler 2636) is apparently of little or no sys- 
tematic importance, having longer and more abundant pubescence 
on the leaf blades, petioles, and peduncles. The species of this group 
are poorly defined by Becker, and it is uncertain how many of them 
are really valid. 

Viola Nannei Polak. Linnaea 41 : 547. 1877. Corazon; violeta. 

Moist, open banks, alpine meadows, often in forest of Alnus, pine 
and oak, Juniper us, or Cupressus, 1,800-4,000 meters; Zacapa; Ja- 
lapa; Guatemala; Sacatepe"quez; Chimaltenango; Solola; Totonica- 
pan; Quiche"; Huehuetenango; Quezaltenango; San Marcos. Mexico 
(Chiapas) ; Costa Rica; Panama. 

Plants low and acaulescent, from a stout or rather slender rootstock, producing 
sometimes short and poorly developed stolons, or these wholly absent; stipules 
1 cm. long or shorter, lanceolate or ovate, acute, remotely long-fimbriate; leaf 
blades reniform, mostly 2-2.5 cm. long and equally broad, broadly rounded at the 
apex, deeply cordate at the base, crenate or subentire, hispidulous, at least near 
the base; flowers rather pale violet, 1 cm. long, the spur short, whitish; sepals 
ovate, hispidulous, the basal appendages short and rounded. 

This has been reported from Guatemala as V. Hookeriana HBK. 
V. Nannei var. glaberrima Becker (Repert. Sp. Nov. 20: 5. 1924; 
type collected between Ziha and Calel, Huehuetenango, 3,000 meters, 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 81 

C. & E. Seler 2942) is glabrous throughout. Apparently it is without 
systematic importance, since glabrous and hispidulous leaves may be 
found on different parts of the same plant. 

Viola odorata L. Sp. PI. 934. 1753. Violeta; sweet violet. 

Native of Europe, Asia, and Africa; now cultivated for ornament 
in all civilized parts of the earth; grown in almost all parts of Guate- 
mala, from sea level to the upper limits of cultivation. 

Plants low, perennial from a thick rootstock, producing long slender stolons; 
leaves on usually very long and slender petioles, cordate-ovate or reniform, crenate, 
deeply cordate at the base, glabrous or nearly so, rounded to subacute at the apex; 
stipules ovate-lanceolate, acuminate, fimbriate; flowers fragrant, deep violet, the 
spur straight, obtuse. 

Violets are one of the favorite flowers of Guatemala, and are 
grown almost anywhere in the country, but they thrive best at middle 
elevations. The flowers produced in Guatemala and Quezaltenango 
often are unusually large, with long stems, and they are sold in great 
bunches in the markets of large cities, as well as in other places. 
A few plants are to be found in almost any patio in the mountain 
regions. 

Viola reptans Robinson, Proc. Am. Acad. 27: 165. 1892. V. 
Schaffneriana Becker, Repert. Sp. Nov. 20: 2. 1924. 

Dry or moist open places or in forest, 1,800-2,500 meters; Chi- 
maltenango; Solola. Mexico. 

Plants acaulescent, from a rather thick, short rootstock, producing elongate 
stolons, these glabrous or sparsely or densely hispidulous; stipules lanceolate or 
linear-lanceolate, long-fimbriate; leaves long-petiolate, the blades broadly ovate, 
obtuse, shallowly cordate at the base, rather thick and firm, crenate, somewhat 
paler beneath, sparsely or densely hispidulous; flowers equaling or longer than the 
leaves, violaceous, the spur short and thick; sepals ovate-lanceolate, the basal 
appendages short, subquadrate. 

Viola scandens Willd. ex Roem. & Schult. Syst. 5: 391. 1819. 
V. Lindeniana Turcz. Bull. Soc. Nat. Moscou 27, Abth. 2: 336. 1854. 

Moist or rather dry thickets and mountain forest, 1,400-2,900 
meters; Baja Verapaz; El Progreso; Quiche"; Huehuetenango; San 
Marcos. Southern Mexico; El Salvador; Honduras; Costa Rica; 
northwestern South America. 

Plants scandent, herbaceous or suff rutescent below, forming dense tangles over 
bushes, often a meter long or even more, the stems glabrous; stipules oblong, re- 
motely dentate-ciliate; leaves slender-petiolate, triangular-ovate or oblong-ovate, 



82 FIELDIANA: BOTANY, VOLUME 24 

2-4 cm. long, acute, shallowly cordate at the base, crenate, glabrous; flowers about 
equaling the leaves, small, the sepals subulate-lanceolate; petals white, the lip 
striped with lavender, acuminate, the spur very short; anthers long-cuspidate at 
the apex; capsule globose, glabrous. 

Viola Seleriana Becker, Repert. Sp. Nov. 20: 6. 1924. 

Type, above Zacaltenango, Huehuetenango, C. & E. Seler 3132; 
San Marcos. 

Plants acaulescent, the rhizome rather thick, the stolons none or short and 
slender; stipules lanceolate, acuminate, 1.5 cm. long, brownish, long-fimbriate; 
leaves long-petiolate, not exceeding the flowers, the petioles glabrous, the blades 
broadly reniform, pointed at the apex, deeply cordate at the base, crenate, hispid- 
ulous, at least beneath near the base; flowers 1 cm. long, pale violet; sepals narrow, 
acute; spur of the lowest petal very short and rounded. 

This is evidently closely related to V. guatemalensis and V. Nan- 
nei, and perhaps all are forms of a single species, to be called V. Nannei, 
or more probably V. Hookeriana HBK. 

Viola tricolor L. Sp. PI. 935. 1753. Pensamiento; pansy. 

Native of Europe, now cultivated for ornament in all civilized 
parts of the earth; grown commonly in Guatemala in most settled 
regions. 

Plants annual or biennial, glabrous or pubescent, the stems angulate, often 
branched; stipules large and foliaceous, conspicuous, lyrate-lobate or deeply lacin- 
iate; upper leaves oval to lanceolate, 1.5-2.5 cm. long, the lower ones ovate, often 
cordate, crenate-serrate; flowers large and broad, variable in size and in color, 
yellow, purple, white, or brown, sometimes blue or almost black. 

Pansies are much grown at middle and rather high elevations, 
and at times even in the lowlands. They are common in small pri- 
vate gardens, and often are planted in beds in the parks. Quantities 
of the flowers are offered for sale in such markets as those of Guate- 
mala and Quezaltenango. 



FLACOURTIACEAE 

Trees or shrubs, sometimes armed with spines or thorns; leaves alternate, sim- 
ple, dentate or entire, generally with pellucid dots or lines; stipules small, decidu- 
ous, or none; flowers mostly small, sometimes large, regular, perfect or unisexual; 
sepals distinct or united below, free from the ovary or sometimes united below 
with it, imbricate or valvate; petals distinct, often none; stamens definite or 
numerous; ovary free or more or less inferior, mostly 1-celled (rarely 2-5-celled), 
the 2-6 placentae parietal; ovules numerous; style simple or divided; fruit bac- 
cate or berry-like, sometimes capsular; seeds usually few, often arillate; endosperm 
carnose, the embryo axial, the cotyledons broad, often cordate. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 83 

About 85 genera, widely dispersed in tropical regions. Several 
additional genera occur farther south in Central America. 

Petals present. 

Flowers in spikes or racemes. 

Leaves 3-5-nerved; flowers 4-parted; ovary superior Bartholomaea. 

Leaves penninerved; flowers 6-7-parted; ovary half inferior Homalium. 

Flowers neither spicate nor racemose, mostly paniculate or corymbose, some- 
times fasciculate or umbellate. 

Fruit echinate Lindackeria. 

Fruit unarmed. 

Fruit with broad, vertical wings; leaves mostly 30-50 cm. long. 

Carpotroche. 
Fruit not winged; leaves much smaller. 

Fruits many-seeded; inflorescences fasciculate or umbellate Prockia. 

Fruits usually 1-seeded; inflorescences paniculate, cymose or corymbose. 

Inflorescences cymose or corymbose; fruits glabrous Hasseltia. 

Inflorescences paniculate; fruits tomentose Pleuranthodendron. 

Petals none. 
Sepals valvate. 

Flowers in slender, usually much elongate racemes Lunania. 

Flowers fasciculate or umbellate Prockia. 

Sepals imbricate. 

Flowers unisexual or polygamous; staminodia none; plants often with spine- 
toothed leaves, or the trunks often armed with branched thorns. 
Leaves usually with spine-tipped teeth; fruit 4-5 cm. broad or larger. 

Olmediella. 

Leaves never with spine-tipped teeth; fruit less than 1 cm. broad . .Xylosma. 
Flowers perfect; leaves never spine- toothed; trunk never armed with branched 
thorns. 

Flowers without staminodia Laetia. 

Flowers with staminodia alternating with the stamens. 

Style none; stamens 20-40 Zuelania. 

Style present; stamens 6-16 Casearia. 



Banara dioica Benth. Journ. Linn. Soc. 5, suppl. 2: 94. 1861. 

Mexico (Veracruz) and Honduras. While this genus has not been 
found in Guatemala or British Honduras the fact that it is found on 
both sides of these countries indicates that it should be discovered 
there. 

BARTHOLOMAEA Standl. & Steyerm. 

Dioecious trees or shrubs, the branches slender, terete, flexuous; stipules mi- 
nute, subulate, deciduous; leaves alternate, small, short-petiolate, 3-5-nerved, 
sparsely pellucid-punctate, crenate-serrate or undulate-serrate, the pubescence of 
simple hairs; flowers mostly 4-parted, sometimes 3-parted, unisexual; inflorescence 



84 FIELDIANA: BOTANY, VOLUME 24 

spicate, axillary, sometimes branched from near the base, solitary; flowers solitary, 
bracteate at the base; sepals of the staminate flowers free, minute, suborbicular; 
petals of the staminate flowers similar to the sepals or sometimes 2-3 times as long; 
stamens 8-12, inserted on an inconspicuous, minute disc, filaments filiform, gla- 
brous, the anthers minute, didymous; style none, the stigmas usually 4, short, 
subulate, ovules 8, 2 on each of 4 parietal placentae; capsule small, glabrous, de- 
pressed-globose, usually 4-valvate to the middle; seeds 3-6, subglobose, covered 
with long, soft hairs, inserted along the middle of the valve near the apex, gla- 
brous within. 

The two following species are known. The genus is dedicated to 
Bartolome* de las Casas, Protector of the Indians, builder of the an- 
cient church of Coban, and one of the most admirable heroes of the 
Conquest. It was he who first introduced European civilization and 
Christianity to the Quecchi Indians of Verapaz, and was indeed the 
author of the very name Verapaz. 

Leaves pilosulous beneath; stamens 8; leaves membranaceous, dull B. mollis. 

Leaves glabrous beneath or only with tufts in axils of nerves; stamens 8-12; leaves 
coriaceous, lustrous B. sessiliflora. 

Bartholomaea mollis Standl. & Steyerm. Field Mus. Bot. 22: 
252. 1940. 

Wet, mixed forest at sea level, along the Rio Dulce, Izabal; en- 
demic; type collected between Livingston and 6 miles up the river, 
on the north side, Steyermark 39452. Figure 11. 

A shrub or small tree, the young branchlets densely spreading-pilosulous; peti- 
oles 4-5 mm. long; leaf blades elliptic-oblong to ovate, 4-8 cm. long, 2-3.7 cm. 
wide, acute or acuminate with obtuse tip, more or less asymmetric at the base, 
narrowly rounded to acute, remotely undulate-serrate or subentire, remotely and 
coarsely pellucid-punctate, very minutely puberulent above, slightly paler be- 
neath, pilose with rather short, spreading hairs, especially along the nerves and 
veins, 3-nerved from the base; staminate spikes almost sessile, very slender, 2.5- 
7 cm. long, many-flowered, the rachis densely pilosulous; bracts equaling the 
sepals; sepals suborbicular, 0.6-0.9 mm. long, rounded at the apex, minutely 
pilosulous outside; petals oval, glabrous, rounded at the apex, 1.2-1.5 mm. long; 
stamens 8, equaling or slightly longer than the petals; valves of the capsule 2.5 mm. 
long and 1.5-2 mm. broad, coriaceous; seeds 1-1.5 mm. long, covered with slender 
hairs 3-4 mm. long. 

Bartholomaea sessiliflora (Standl.) Standl. & Steyerm. Field 
Mus. Bot. 22: 252. 1940. Lunania sessiliflora Standl. Carnegie Inst. 
Wash. Publ. 461: 74. 1935. Bartholomaea paniculata Lundell, Wrightia 
2: 55. 1960 (type from Pete"n, Guatemala, Lundell 15383). 

Moist or wet lowland forests, 300-450 meters; Pete"n; Alta Vera- 
paz. British Honduras (type Schipp S606). 




FIG. 11. Bartholomaea mottis. A, Flowering stem; X %. B, Pistillate flower; 
X 10. C, Seed, showing attached hairs; X 10. D, Pistillate flower; X 10. 



85 



86 FIELDIANA: BOTANY, VOLUME 24 

A glabrous tree to 6 m. high, the trunk to 7.5 cm. in diameter, the slender 
branches sparsely lenticellate; petioles 4-5 mm. long; leaf blades broadly lanceo- 
late to ovate, 4.5-8.5 cm. long and 2.5-4 cm. broad, acuminate, oblique at the 
base and rounded on one side, acute on the other, closely serrate, lustrous above, 
fuscous when dry, dull beneath, 3-5-nerved at the base, with tufts of hair in the 
axils of the nerves below; staminate spikes 6-8 cm. long; flowers remote, scarcely 
1.2 mm. long; bracts lanceolate, acuminate, about 1 mm. long; sepals 4-5, broadly 
rounded, 1.2 mm. long; petals 4-5, suborbicular, about 1 mm. long; stamens 8-12, 
the filaments about 0.5 mm. long; pistillate flowers with 4 slightly imbricated se- 
pals, the sepals about 1.2 mm. long, disc somewhat fimbriated, fleshy; ovary globose, 
0.7 mm. long and 1 mm. broad, style obsolete, stigmas 4, spreading; ovules 8, 
2 on each of 4 parietal placentae; seeds about 1 mm. long, pilose. 



GARPOTROCHE Endlicher 

Reference: H. Pittier, Contr. U. S. Nat. Herb. 12: 177-181, t. 19. 
1909. 

Shrubs or small trees, unarmed; leaves usually very large, short-petiolate, 
membranaceous or thicker, serrate or dentate or subentire; stipules lanceolate or 
subulate, caducous; flowers mostly large, white, chiefly in short, axillary and ter- 
minal cymes or racemiform cymes, sometimes arising on the branches below the 
leaves, polygamo-dioecious; sepals 3; petals 6-9, strongly imbricate; stamens 
numerous, inserted on a small torus, the filaments short, the anthers linear; ovary 
with 6-7 placentae, the ovules numerous; styles 6-7, connate at the base, simple; 
fruit large, globose, baccate, bearing several or numerous broad vertical wings. 

A small genus of tropical America with perhaps 8-10 species. 
There are two more species described (and one undescribed) from 
Costa Rica and Panama. 

This genus has been considered by Macbride [Field Mus. Bot. 
13 (4) : 16. 1941] to be referable to Mayna. However, we believe it 
desirable, at least for the present, to maintain Carpotroche. A study 
of Carpotroche and allied genera of this difficult family is much needed. 

Carpotroche platyptera Pittier, Contr. U. S. Nat. Herb. 12: 
178, t. 19, f. 15, 16. 1909. Sucte; jaya. 

Dense, wet, mixed, lowland forest, at or little above sea level; 
Izabal. Honduras to Panama, along the Atlantic coast. 

A shrub or small tree, usually 2-5 meters high, with few branches, often sim- 
ple; leaves on petioles 4-5 cm. long, oblong-oblanceolate, mostly 30-50 cm. long 
and 10-14 cm. wide, usually narrowly cuspidate-acuminate, long-attenuate to the 
base, coarsely dentate, finely and softly pubescent beneath or in age often almost 
glabrous, glabrate on the upper surface; staminate inflorescence much reduced 
and few-flowered, the flowers appearing almost sessile in the leaf axils or on the 
trunk, the pedicels and calyx densely pubescent; sepals 2-3, concave, furfuraceous 
outside; petals about 4.5 mm. long, white, ovate or elliptic; stamens 15-19, the 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 87 

anthers 2.5 mm. long; pistillate flowers large, about 3 cm. broad, the petals about 8, 
elliptic-obovate, obtuse; ovary ovoid, pubescent; placentae generally 4, the styles 4, 
very short; fruit green or purplish, globose, about 5 cm. in diameter, with several 
broad vertical wings. 

The bark from plants of this genus is said to be rich in tannin. 
The wood is described as yellowish or pale brown, fine- textured, 
hard, tough, and strong, but apparently no use is made of it in Cen- 
tral America. 

CASEARIA Jacquin 

Trees or shrubs, commonly unarmed but sometimes with spinose branchlets; 
stipules mostly small and soon deciduous; leaves petiolate, entire or dentate, gen- 
erally with pellucid dots or lines; flowers perfect, fasciculate or umbellate, some- 
times solitary or racemose, greenish, whitish, or yellowish, small, the pedicels 
articulate, bracteate at the base; sepals 4-6, imbricate; petals none; stamens 
usually 6-15 (to 22 in C. spiralis), inserted in a single series on the tube of the 
calyx or at its base, alternating with staminodia; filaments free or united with the 
staminodia to form a ring; ovary free, ovoid or oblong, narrowed to a short style, 
the stigma capitate or with 3 stigmas; ovules numerous, on 3 parietal placentae; 
fruit capsular, fleshy or dry, 3-4-valvate, containing numerous seeds; seeds oblong 
or angulate, arillate; endosperm carnose, the embryo straight, the cotyledons ob- 
long to rounded, flat, the radicle cylindric. 

A genus found in the tropics of both hemispheres, chiefly in 
America and Africa. There are perhaps not more than thirty or 
forty species in the American tropics although many more have 
been described. A few others have been found in southern Central 
America and several more in Mexico. 

Flowers in stipitate or pedunculate heads or in cymes or panicles. 

Stamens 16-22 C. spiralis. 

Stamens less than 12. 

Flowers in small, dense, stipitate or pedunculate heads C. arborea. 

Flowers in cymes or panicles C. nitida. 

Flowers in sessile fascicles borne in leaf axils. 

Flowers small, about 2 mm. long; leaves entire or obscurely denticulate. 

Leaves rounded at the base, narrowly oblong, coloration of two sides con- 
trasting C. belizensis. 

Leaves mostly acute at the base, lanceolate or lance-oblong, coloration of two 
sides similar C. syhestris. 

Flowers larger, 3-12 mm. long; leaves entire or dentate. 

Leaves serrate, with close, dense, and very acute, conspicuous teeth. 

C. arguta. 
Leaves entire or sinuate dentate. 

Leaf blades mostly 4-8 cm. long, usually obtuse or narrowly rounded at the 

apex, obscurely crenate; plants usually spinose C. aculeata. 

Leaf blades mostly 10-18 cm. long, acute or acuminate; plants unarmed. 



88 FIELDIANA: BOTANY, VOLUME 24 

Sepals 8-12 mm. long. 

Ovary and young fruit glabrous or sparsely pubescent; leaves glabrous. 

C. Bartlettii. 

Ovary and young fruit densely pilose or tomentose; leaves pilose be- 
neath C. tacanensis. 

Sepals 2-6 mm. long. 

Sepals 3 mm. or less long; leaves mostly rounded at the base. 

C. belizensis. 

Sepals 3-4 mm. long; leaves mostly acute at the base, but sometimes 
obtuse C. javitensls var. myriantha. 

Casearia aculeata Jacq. Enum. PI. Carib. 21. 1760. C. hirta 
Swartz, Fl. Ind. Occ. 756. 1798. C. ramiflora Vahl var. spinosa 
Griseb. Fl. Brit. W. Ind. 24. 1859. Limoncillo; pullun (Pet&i, Maya). 

Dry or moist thickets, sometimes in open forest, 350 meters or 
lower; Pete"n; Alta Verapaz; Izabal; Santa Rosa; Escuintla; Suchite- 
pe"quez; Retalhuleu; San Marcos; Huehuetenango. Southern Mex- 
ico; British Honduras to Honduras and Panama; West Indies; South 
America. 

A shrub or tree, commonly 2-3 meters high but sometimes as much as 10 me- 
ters, with a trunk 10 cm. or more in diameter, the branches often or occasionally 
armed with stout spines or branched thorns, puberulent to glabrate; leaves short- 
petiolate, membranaceous or papyraceous, elliptic to obovate-elliptic, mostly 4- 
8 cm. long and very obtuse, broadly cuneate at the base, remotely serrate-dentate 
or almost entire, glabrous or nearly so, the lateral nerves 4-5 pairs; flowers fascicu- 
late in the leaf axils, the pedicels short, articulate at the middle or lower, 5-7 mm. 
long; sepals 4-5 mm. long, greenish white; stamens glabrous, the staminodia vil- 
lous; ovary villous; fruit subglobose, obtusely trigonous, 6-12 mm. long. 

Called "escambrdn" and "aguja de arrea" (Honduras); "wild 
lime," "bird berries," "limoncillo" and "ramoncillo" (British Hon- 
duras); "cedrdn" (Oaxaca). A common shrub in dry thickets or 
second growth of the Pacific plains, often forming dense tangles of 
spiny branches. 

Casearia arborea (L. Rich.) Urban, Symb. Antill. 4: 421. 1910. 
Samyda arborea L. Rich. Act. Soc. Hist. Nat. Paris 1: 109. 1792. 

Wet forest or thickets, at or near sea level; Izabal. British Hon- 
duras; Nicaragua; Costa Rica; Panama(?). Greater Antilles; Guianas 
and Brazil. 

A large shrub or tree, sometimes 9 meters high with a trunk 10 cm. in diam- 
eter, the branches slender, when young puberulent or glabrate; leaves on very 
short petioles, narrowly oblong or oblong-lanceolate, mostly 5-12 cm. long, nar- 
rowly long-acuminate, acute or very obtuse at the base and often somewhat 
unequal, distichous, closely serrulate-dentate, glabrous and lustrous above, paler 



STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 89 

beneath, sometimes pale-tomentulose; flowers subcapitate, white, the heads short- 
pedunculate or stipitate, the peduncles or stipes mostly 2-3 mm. long, the pedicels 
about equaling the calyx, puberulent; sepals 3.5-4.5 mm. long; staminodia pilose; 
stamens 10; ovary glabrous except at the apex; fruit ellipsoid, 4 mm. long. 

Casearia arguta HBK. Nov. Gen. & Sp. 5: 364. 1821. Manza- 
nilla; raspa-lengua; ixim-che (Alta Verapaz) ; guayabillo; camche (fide 
Aguilar) . 

Dry or moist thickets or open forest, often in second growth, 
2,000 meters or less, most common at low elevations; Alta Verapaz; 
Izabal; Jutiapa; Santa Rosa; Escuintla; Guatemala; Quiche"; Suchi- 
tepe"quez; Retalhuleu; Quezaltenango; San Marcos. Western and 
southern Mexico; British Honduras and El Salvador to Panama. 
South America. 

A shrub or small tree, unarmed, usually 2-5 meters tall but sometimes a tree 
of 12 meters with a trunk 30 cm. in diameter, the branches grayish or pale brown, 
the trunk often crooked, the crown spreading, the branches usually pubescent; 
leaves short-petiolate, membranaceous, oblong, lance-oblong, or oblong-elliptic, 
7-15 cm. long, acuminate or narrowly long-acuminate, rounded to acute at the 
base, closely and acutely serrate throughout, glabrous above or nearly so, glabrate 
beneath or densely pilose, even in age, sparsely punctate, usually barbate in the 
leaf axils; flowers greenish white or creamy yellow, densely fasciculate, the ped- 
icels 2-4 mm. long, densely pubescent; sepals 4-5 mm. long, densely pubescent; 
stamens generally 8, the staminodia white- villous; stigma entire; fruit subglobose, 
1-2 cm. in diameter at maturity; aril of the seeds red. 

The flowers have a honey-like odor. The inflorescences some- 
times are fasciated and deformed, presumably as the result of insect 
attacks. Called "cuculmico" in El Salvador; in British Honduras 
"hard moho" and "monkey plum." The wood is creamy white or 
pale yellow, the scant heartwood dark red or almost black. The ripe 
fruit is yellow. This species is especially common on the Pacific 
plains and foothills of Guatemala. 

Casearia Bartlettii Lundell, Lloydia 2: 104. 1939 (type from 
Guatemala, Bartlett 12558); C. elegans Standl. in Yuncker, Field 
Mus. Bot. 9: 311. 1940 (type from Honduras, Yuncker et al 8804); 
Samyda Bartlettii Lundell, Wrightia 2: 56. 1960. 

Wet, mixed forest at or near sea level; Pete"n; Izabal. Mexico 
(Michoacan, Tabasco); Honduras. 

A slender shrub or small tree, sometimes 7 meters high, the branchlets pilosu- 
lous or puberulent at first, soon glabrate; petioles mostly 4-10 mm. long; leaf 
blades membranaceous, oblong-oblanceolate to obovate-oblong, 10-19 cm. long, 
3-6.5 cm. wide, gradually or abruptly acuminate or long-acuminate, acute at the 
base or usually gradually long-attenuate, glabrous, serrate or serrulate or sub- 



90 FIELDIANA: BOTANY, VOLUME 24 

entire, sparsely pellucid-lineolate, green when dried, the lateral nerves about 8 
pairs; flowers axillary, solitary or in few-flowered fascicles, the very slender ped- 
icels 5-7 mm. long or even longer, densely short-pilosulous; sepals white, obovate- 
oblong, 7-12 mm. long, obtuse or rounded at the apex, densely and minutely 
puberulent outside; stamens 6-12, the filaments glabrous; staminodia densely 
pilose; ovary glabrous; fruit ovoid, about 2.5 cm. long and 1.5 cm. broad, apiculate, 
acutish at the base, prominently ridged. 

Casearia belizensis Standl. Field Mus. Bot. 12: 412. 1936. 

Wet forest, near sea level; Izabal. British Honduras (type, 
Schipp 1314). Nicaragua; Costa Rica. 

A tree, sometimes 12 meters high, the trunk 25 cm. in diameter, the young 
branchlets minutely puberulent, soon glabrate; leaves thick-chartaceous, on peti- 
oles 5-8 mm. long, narrowly oblong, 6-21 cm. long, 2.5-5 cm. wide, cuspidate- 
acuminate with a long narrow tip, obliquely rounded at the base or subcordate, 
inconspicuously and closely appressed-crenate, very densely pellucid-punctate, 
glabrous, lustrous above, paler and brownish beneath, the lateral nerves about 
10 pairs; flowers pink, numerous, fasciculate in the leaf axils or at defoliated 
nodes, the slender pedicels glabrous, 1 cm. long or less; sepals oval, glabrous, 
2 mm. long or slightly larger; stamens 8, glabrous; ovary glabrous; the style short 
and thick, entire. 

In British Honduras the tree is called "drunken bayman wood"; 
in Costa Rica it is called "espino amarillo," and a tree there has been 
reported to be 110 feet tall and 32 inches D.B.H. and to be used as 
ordinary lumber. 

Casearia javitensis HBK. var. myriantha (Turcz.) L. Wms 
Fieldiana, Bot. 29: 359. 1961. C. myriantha Turcz. Bull. Soc. Naf 
Mosc. 36: 609. 1863. C. Brighamii Watson, Proc. Amer. Acad. 21: 
459 1886 (type Izabal, S. Watson 18). C. laetioides Warb. in Engl. & 
Prantl, Pflanzenfam. 3, Abt. 6a: 52. 1893 (type from Izabal, Bernoulli 
896). Mierda de gallina (Pet&i); taixcaax, utaxcaax (Pet&i, Maya, 
fide Lundell) ; cakica-che (Alta Verapaz) ; guayabillo. 

Moist or wet thickets or forest, or often in dry situations, 1,200 
meters or less; Pete"n; Alta Verapaz; Izabal; Escuintla; Retalhuleu; 
Quezaltenango. Southern Mexico; British Honduras to Panama; 
northern South America. 

A shrub or tree, sometimes 11 meters high with a trunk 20 cm. in diameter, 
the bark smooth, pale, the rather stout branches glabrous or puberulent; leaves 
on short thick petioles, coriaceous, oblong to elliptic-oblong or lance-oblong, 8 
25 cm. long, acuminate or long-acuminate, usually acute at the base, very lus- 
trous, opaque, glabrous, rather inconspicuously appressed-serrate, the lateral nerves 
about 8 pairs; flowers white or greenish white, densely fasciculate, the pedicels 
often very numerous, 7 mm. long or less, densely puberulent; sepals 3-4 mm. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 91 

long, appressed-pilosulous with pale hairs; stamens 10-15, slightly longer than 
the sepals; stigma 3-fid; capsule sparsely pilose, about 1 cm. long, thin-walled, 
turning red and finally brown. 

Called "cafecillo" and "pochitoquillo" in Tabasco. This is dis- 
tinguished from all other local species by its coriaceous, lustrous 
leaves and pubescent calyces. Casearia belizensis is possibly only a 
somewhat pubescent phase of this species and perhaps should be con- 
sidered another variety of it. 

Casearia nitida Jacq. Enum. PL Carib. 21. 1760. Samyda nitida 
L. Syst. Nat. ed. 10, 1025. 1759. C. corymbosa HBK. Nov. Gen. & 
Sp. 5: 366. 1821. C. banquitana Krause, Beih. Bot. Centralbl. 32, 
Abt. 2: 345. 1914. C. laevis Standl. Contr. U. S. Nat. Herb. 23: 845. 
1923. C. mollifolia Standl. Field Mus. Bot. 9: 311. 1940. C. Gentlei 
Lundell, Field & Lab. 13: 9. 1945 (type from British Honduras, 
Gentle 4489). C. banquitana var. laevis Johnston, Sargentia 8: 211. 
1949. Cf. L. Wms. Fieldiana, Bot. 29: 359. 1961. Vara blanca; 
sakiche, ixim-te (Alta Verapaz) ; cafe de monte; cafe de montana; bale- 
lac de aguada; canjuro; iximche (British Honduras, Maya). 

Dry, wet or moist forests or thickets, often in second growth, 
abundant in many regions, especially along the Pacific plains and 
foothills, 1,300 meters or lower; Pete"n; Alta Verapaz; El Progreso; 
Izabal; Zacapa; Jutiapa; Santa Rosa; Escuintla; Guatemala; Saca- 
tepe"quez; Huehuetenango; Quezaltenango; San Marcos. Mexico to 
Panama; the West Indies; northern South America. 

A shrub or small tree, sometimes to 9 m. tall but usually less, the bark and 
branches whitish, the branchlets pilose or glabrous; leaves short-petiolate, mem- 
branaceous, elliptic or oblong-elliptic, lance-oblong or oblong, mostly 4-9 cm. long 
and 3-5 cm. broad, acute or acuminate, rarely obtuse, rounded to acute at the 
base, pellucid-punctate, serrulate to subentire, from entirely glabrous to densely 
velutinous-pilose below, sometimes pilose above; flowers in short corymbs or pan- 
icles, few- to several-flowered, the pedicels short or rather elongate, pilosulous to 
pilose; sepals 3-4 mm. long, usually densely puberulent to pilosulous; stamens 
usually 8; stigma entire; fruit ovoid or ellipsoid, 8 mm. long or more, becoming 
red or yellow. 

Known in El Salvador as "canjuro," "chilillo," and "come-cule- 
bra;" "comida de culebra" in Honduras; "paletillo" in British Hon- 
duras; "cafetillo" in Tabasco; "xmaben-che" in Yucatecan Maya. 
The name "iximche," Maya and signifying "maize- tree," is of inter- 
est. There is no obvious reason why the tree should be so called, 
but the name and the tree probably have some connection with the 
old Indian capital of Iximche, near Tecpan in the department of 
Chimaltenango. 



92 FIELDIANA: BOTANY, VOLUME 24 

The description above covers both the glabrous and the pubes- 
cent phases of this species. The pubescent form, which has received 
several names, varies from puberulent to densely pilose-pubescent 
and seems to occur within the range of the typical glabrous form 
from Mexico at least to Nicaragua. 

The name Casearia nitida Jacq. should not be written with a 
parenthetical authority. 

Casearia spiralis J. R. Johnston, Proc. Am. Acad. 40: 691. 1905. 
Zuelania belizensis Lundell, Wrightia 1 : 58. 1945 (type from British 
Honduras, Gentle 4649). 

Secondary forests, often in swampy places. British Honduras; 
Mexico; Venezuela. 

Trees to 15 m. tall, the branches dark with a few white, round to elongate len- 
ticels, glabrous; leaves lanceolate-ovate to oval, acute or acuminate, membrana- 
ceous, somewhat pellucid, obscurely denticulate or entire, glabrous, lateral nerves 
mostly 8-12 pairs, the blade 4-9 cm. long and 1.5-4 (5) cm. broad, the petiole 
short, slender, 0.5-1.2 cm. long; inflorescence a pedunculate or stipitate fascicle 
or perhaps condensed, paniculate, many-flowered, and borne on short pseudo- 
branches; sepals tepal-like, 6 in an outer and an inner series, imbricate, those of 
the outer series smaller, lanceolate and about 4-5 mm. long, those of the inner 
series larger, oval and 5-6 mm. long; petals none; stamens 16-22 with shorter 
alternating staminodia, the filaments pilose to nearly glabrous, the staminodia 
densely pilose-pubescent; ovary ovoid, pubescent at least when young. 

Unlike other species of Casearia in the larger number of stamens, 
and in the showy tepal-like sepals which were originally described as 
cyclic or spiralled. 

The disjunct range is of interest. We have not been able to find 
this species among West Indian material, although it should occur 
there. 

Casearia sylvestris Swartz, Fl. Ind. Occ. 2: 752. 1800. Samyda 
parviflora L. Syst. Nat. ed. 10. 1025. 1759, not Loefl. 1758. C. parvi- 
flora Willd. Sp. PL 2: 627. 1799. Coralillo (Suchitepe'quez) ; sacumba 
(Alta Verapaz). 

Moist or dry forest or thickets, often in second growth, 1,200 
meters or less, chiefly at low elevations; Pete"n; Alta Verapaz; Iza- 
bal; Santa Rosa; Escuintla; Retalhuleu; Suchitepe'quez; Solola; 
Huehuetenango. Mexico; British Honduras to Panama; West In- 
dies; South America. 

A shrub or tree, sometimes 9 meters high with a trunk 15 cm. in diameter 
but usually lower, the branches mostly long and slender, whitish or pale brownish; 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 93 

leaves on very short petioles, membranaceous or chartaceous, lanceolate or lance- 
oblong, mostly 6-10 cm. long and 2-3 cm. wide, narrowly long-acuminate, acute 
and oblique at the base, entire, glabrous or nearly so, densely pellucid-punctate; 
flowers small, white or greenish white, numerous, densely fasciculate in the leaf 
axils, the pedicels 2-4 mm. long, glabrous or pubescent, often almost obsolete; 
sepals 1.5-2 mm. long, glabrous or puberulent; stamens 10; stigma 3-lobate; fruit 
subglobose, red, 3-4 mm. in diameter, 2-6-seeded. 

Called "wild sage" in British Honduras; "sombra de armado," 
"sombra de conejo" (Honduras). The wood is said to be yellow, 
hard, heavy, and compact, and to be used in some regions for con- 
struction purposes when available in large enough pieces. 

The typical phase of this species is glabrous. A phase with pubes- 
cent pedicels and calyces is found through most of Central America. 

Casearia tacanensis Lundell, Lloydia 4: 54. 1941. 

At 1,000-2,000 meters; Chiapas; type from Volcan de Tacana, 
E. Matuda 2441; doubtless extending into San Marcos. 

A tree of 6-10 meters, the trunk as much as 15 cm. in diameter, the branchlets 
brownish-torn entose; leaves firm-membranaceous, on stout petioles 3-6 mm. long, 
lance-oblong or elliptic-oblong, 8.5-18 cm. long, 3-5 cm. broad, acuminate, acute 
at the base, almost glabrous on the upper surface, brown-pilose beneath, obscurely 
crenulate-serrate; flowers fasciculate, numerous, the pedicels 5-8 mm. long, artic- 
ulate near the middle, brown-tomentulose; calyx tomentulose, 8 mm. long or 
somewhat shorter, the 5 lobes linear-lanceolate, attenuate, reflexed at about the 
middle; stamens 10, the filaments sparsely pilose, joined into a tube and connate 
to the sepals; staminodia densely pilose; ovary densely brown-pilose; fruit globose, 
2 cm. or slightly more in diameter, densely, coarsely, and irregularly obtuse- 
tuberculate, densely brown-tomentose. 

This is perhaps the most unusual species of the genus. The large 
tuberculate fruits are not duplicated in any other species. 



Flacourtia Ramontchi I/Her., the Governor plum or Batoko plum, 
native of the Zambesi region of Africa, is planted in a few localities 
in Guatemala, as at Finca El Zapote, Escuintla. It is a shrub or 
small tree, with or without spines, the leaves oblong to elliptic or 
obovate, obtuse or acute, crenate-dentate, short-petiolate; sterile 
flowers in short racemes, the fertile ones solitary or geminate; fruit 
cherry-like, 2.5 cm. or less in diameter, purple, red, or blackish, con- 
taining 8-10 seeds. The fruit is edible, and on this account the plant 
has been introduced into cultivation in the West Indies and else- 
where in America, but apparently never has become popular. 



94 FIELDIANA: BOTANY, VOLUME 24 

HASSELTIA HBK. 

Shrubs or trees; leaves slender-petiolate, 3-nerved from the base, alternate, 
entire or serrate, usually with glands on the upper side at the base of the blade; 
stipules small, caducous; flowers small, white, cymose or corymbose, terminal, 
pedicellate, perfect; sepals 4-5, valvate, persistent; petals 4-5, persistent, resem- 
bling the sepals and alternate with them; stamens numerous, free, inserted upon 
small eglandular disk, the filaments filiform, the anthers small, subglobose, 2- 
celled, longitudinally dehiscent; ovary free, 2-celled, the ovules numerous, the 
style subulate, entire; seeds usually 1 in each cell, sometimes 2, pendulous, the 
testa coriaceous, the endosperm carnose; embryo straight, the cotyledons flat, 
foliaceous. 

About five species, ranging from Mexico to Peru in continental 
America. One other species, H. floribunda HBK., occurs in Cen- 
tral America from Honduras to Panama, and in South America. 

Hasseltia guatemalensis Warb. in Engl. & Prantl, Pflanzen- 
fam. 3, Abt. 6a: 32. 1893 (type from Santa Maria; probably Santa 
Maria de Jesus, Quezaltenango, Bernoulli & Carlo 2920). Canela 
de montana. 

Moist or wet, mixed forest, also in coffee plantations, 1,200-1,500 
meters; Suchitepe*quez ; Quezaltenango; San Marcos. Mexico (Chi- 
apas) ; Nicaragua. 

A tree of 9-20 meters, the trunk 30 cm. or less in diameter, the bark close, 
gray, slightly rough, glabrous throughout or nearly so; petioles 1-3.5 cm. long; 
leaf blades oblong-elliptic to elliptic or oblong-obovate, 18 cm. long and 8 cm. wide 
or smaller, very obtuse to short-acuminate, broadly acute to narrowly cuneate- 
acute at the base, with 2 small glands on the upper side at the base, subentire, 
3-plinerved, the costa emitting 3-4 pairs of lateral nerves above the basal nerves; 
inflorescence usually large and many-flowered, broadly corymbose and repeatedly 
dichotomous, the flowers aggregate in umbels, the pedicels mostly 1-1.5 cm. long, 
glabrous, the flowers white; sepals 3-4 mm. long, the outer ones almost glabrous 
outside, the petals tomentulose; fruit glabrous, subglobose, 1 cm. in diameter. 



HOMALIUM Jacquin 

References: S. F. Blake, The genus Homalium in America, Contr. 
U. S. Nat. Herb. 20: 221-235. 1919; L. Wms. Fieldiana, Bot. 29: 
362. 1961. 

Shrubs or large trees; leaves alternate, petiolate, glandular-crenate or serrate, 
rarely entire; stipules small; flowers small, in axillary panicles; calyx tube turbinate, 
connate with the base of the ovary, the lobes 6-7, persistent; petals 6-7, inserted 
in the throat of the calyx, narrow, persistent, plane; stamens collected in fascicles 
opposite the petals and alternating with glands, the filaments filiform, the anthers 
small, didymous; ovary semisuperior, the styles 2-5, filiform, the stigmas simple 
or capitellate; ovules few or numerous, affixed to 2-5 parietal placentae; capsule 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 95 

semisuperior, coriaceous, 2-5-valvate at the apex, few-seeded; seeds angulate or 
oblong, with crustaceous testa, the endosperm carnose; cotyledons foliaceous. 

Species 40 or more, in the tropics of both hemispheres. No other 
species are known from Central America. 

Stamens 5 or more in each fascicle; the filaments pilose, at least toward the base. 

H. trichostemon. 

Stamens 4 or fewer (usually 3) in each fascicle; filaments not pilose . .H. racemosum. 

Homalium racemosum Jacq. Enum. PL Carib. 24. 1760. 
H. hondurense Donn.-Sm. Bot. Gaz. 20: 4. 1895. H. nicaraguense 
Blake, Contr. U. S. Nat. Herb. 20: 225. 1918. H. riparium Standl. 
Carnegie Inst. Wash. Publ. 461: 74. 1935, not Gilg, 1908 (type, 
British Honduras, Schipp 1182). H. Schippii Standl. Tropical 
Woods 52: 27. 1937. 

Moist to wet forest, at or little above sea level; Pete"n, possibly 
also Alta Verapaz. British Honduras; Mexico through Central 
America; the West Indies; northern South America to Brazil and 
Colombia, perhaps Peru. 

A tree to 15m. high with a trunk 20 cm. or more in diameter, the branchlets 
glabrous; leaves on petioles 5-8 mm. long, oblong-lanceolate or oblong, mostly 
7-12 cm. long and 2.5-4.5 cm. wide, narrowly long-acuminate to acute, acute or 
obtuse at the base, crenate-serrate, glabrous, lustrous, the lateral nerves about 8 
pairs; flowers whitish, the racemes usually many-flowered, 16 cm. long or less, the 
rachis densely puberulent, the pedicels 1-8 mm. long, the bracts subulate, often 
persistent and conspicuous; calyx tube obconic, acute at the base, the sepals lance- 
oblong, 2.5-5 mm. long, obtuse, tomentulose, 1.2-2 mm. wide; petals white, 3- 
7 mm. long, oblong-elliptic, acute or subobtuse, densely tomentulose; stamens 3-4 
in each fascicle, the filaments glabrous; ovary densely hirsute. 

The flowers are said to have an unpleasant odor. 

Homalium trichostemon Blake, Contr. Gray Herb. 53: 60. 
1918. 

Common along rocky river banks, about 300 meters; Escuintla. 
Southern Mexico. 

A tree 9-12 meters high or larger, the trunk often 30 cm. or more in diameter, 
the bark thin, light gray, the crown broad or narrow; petioles 4-7 mm. long; leaf 
blades elliptic-oblong to elliptic or oval, 6-13 cm. long, 3-6 cm. wide, acuminate 
to obtuse, cuneate to rounded at the base, crenate-serrate, barbellate beneath in 
the nerve axils, otherwise glabrous, the lateral nerves 7-8 pairs; racemes simple, 
solitary or geminate, lax or dense, the rachis puberulent or pilosulous, the pedicels 
mostly 1-3 mm. long; calyx tube in fruit densely puberulent, the sepals 6-7, ovate- 
lanceolate, subobtuse, 4-4.5 mm. long, 1.5-1.8 mm. wide; petals ovate, obtuse, 
puberulent and ciliate, 4.8-5.5 mm. long; stamens in fascicles of 5-7, the filaments 
pilose to the middle, shorter than the petals. 



96 FIELDIANA: BOTANY, VOLUME 24 

Called "palo de piedra" in Oaxaca. The wood is described as 
light brown throughout. The tree is a characteristic one along rocky 
banks of streams on the plains between Escuintla and the coast. 

LAETIA Loefling 

Shrubs or trees; leaves petiolate, crenate, serrate, or subentire, pellucid-punc- 
tate; flowers perfect, axillary or terminal, fasciculate or corymbose; sepals 4-5, 
broad, somewhat petaloid, imbricate, reflexed in anthesis; petals none; stamens 
usually numerous, inserted on an eglandular disk, the anthers ovoid; ovary 1- 
celled, with 3 placentae, many-ovulate; style simple, the stigma capitate or obso- 
letely lobate; fruit baccate, sometimes resinous within, tardily dehiscent by valves; 
seeds arillate, with coriaceous testa; embryo straight, the cotyledons broad. 

Ten or more species, in tropical America. A single species is 
known from continental North America. 

Laetia Thamnia L. Amoen. Acad. 5: 413, 379. 1760. L. ameri- 
cana L. Syst. ed. 10. 1074. 1759, in part. Hueso de tortuga (Pete"n); 
ixbakelac, bakelac (Pete"n, Maya). 

Moist or wet forest or thickets, at or little above sea level; Pete"n; 
Izabal. Campeche to Chiapas, Yucatan, and Tabasco in Mexico; 
British Honduras; Honduras; Panama; West Indies; Colombia. 

A shrub or tree, sometimes 10 meters high with a trunk 10-40 cm. in diameter, 
the young branches glabrous; stipules minute, deltoid-ovate, deciduous; leaves 
short-petiolate, membranaceous, elliptic to oblong-lanceolate or oblanceolate- 
oblong, mostly 6-15 cm. long, acute or acuminate, obtuse or narrowly rounded 
at the base, entire or obscurely crenulate, pellucid-punctate; flowers fragrant, 
white or cream sometimes tinged with pink, in few-flowered pedunculate corymbs 
much shorter than the leaves, slender-pedicellate; sepals 4, mostly 4-6 mm. long, 
sparsely puberulent; filaments pubescent; fruit globose, fleshy, 1.5-4 cm. in diam- 
eter, 4-5-valvate, many-seeded, densely and closely tomentulose. 

LINDACKERIA Presl 

Shrubs or trees; leaves alternate, long-petiolate, penninerved; flowers rather 
small, polygamous, in small or large, axillary, usually many-flowered panicles or 
racemes; sepals 3, imbricate; petals 6-12, imbricate; stamens numerous, free or 
united, the filaments filiform, the anthers linear; ovary stipitate, 1-celled, muri- 
cate, with 3 parietal placentae, the ovules numerous; style elongate, simple, the 
stigma 3-lobate; fruit globose, coriaceous or subligneous, densely tuberculate or 
echinate, finally 3-2-valvate, few-seeded; seeds ovoid, with crustaceous testa, the 
cotyledons ovate, foliaceous, inflexed at the apex; endosperm copious. 

About 12 species, in tropical America and Africa. A single spe- 
cies is found in North America. The genus is closely related to 
Mayna and not very distinct from it. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 97 

Lindackeria laurina Presl, Rel. Haenk. 2: 89, t. 65. 1836. On- 
coba laurina Warb. in Engl. & Prantl, Pflanzenfam. 3, Abt. 6a: 19. 
1894. Mayna laurina Benth. Journ. Linn. Soc. Bot. 5: Suppl. 81. 
1861. 

Wet forest, at or little above sea level; Izabal. British Hon- 
duras; along the Atlantic coast to Panama. 

A large shrub or a tree, sometimes 15 meters high with a trunk 25 cm. in diam- 
eter, the crown spreading or pyramidal, the bark gray, very viscid on the younger 
parts and on the leaves, glabrous almost throughout; leaves on long slender peti- 
oles, membranaceous or thicker, lance-oblong or ovate-oblong, 12-30 cm. long, 
cuspidate-acuminate, obtuse at the base, entire, very lustrous, brownish beneath 
when dried; panicles long and narrow, pedunculate, usually shorter than the leaves, 
sparsely puberulent, the flowers pedicellate; sepals and petals white, the petals 
1 cm. long or less; stamens yellow; fruit globose, green, 1 cm. in diameter, covered 
with long slender flexible spines; seeds 1-4. 

In the fresh state the foliage is very viscid and adheres to the 
fingers and to clothing. The name "achiote" has been reported for 
this tree in Guatemala, but probably in error. 

The species is reported from Colombia and Venezuela but speci- 
mens we have seen, so determined, perhaps belong elsewhere. 

LUNANIA Hooker 

Trees or shrubs with slender branches; leaves alternate, petiolate, entire, 
3-5-nerved, minutely pellucid-punctate; stipules none; flowers small, in slender, 
axillary or terminal, simple or branched, often nutant racemes, the pedicels artic- 
ulate at the base, minutely several-bracteate; calyx subglobose, finally valvately 
cleft into 2-3 spreading membranaceous sepals; petals none; stamens 6-10, in- 
serted about a cupular disk; filaments short, the anthers ovoid or oblong, ex- 
trorsely dehiscent; ovary ovoid, 1-celled, attentuate to a short or elongate, 3-lobate 
style, the stigmas truncate; placentae 3, parietal, the ovules numerous, multiseri- 
ate; fruit capsular, coriaceous, globose, 3-valvate, few-seeded; seeds small, globose, 
usually deeply punctate, arillate, with endosperm. 

About 15 species, ranging from southern Mexico to northern 
South America. Two other Central American species are known, 
in Honduras and Costa Rica. 

Lunania mexicana T. S. Brandegee, Univ. Calif. Publ. Bot. 6: 
56. 1914. Palo chito; chile de montana. 

Dense, moist or wet, mixed, mountain forest, 750-2,000 meters; 
Chimaltenango; Suchitepe"quez; Solold; Quiche"; Quezaltenango; San 
Marcos. Mexico (Chiapas) ; Honduras. 

A large slender shrub or a tree, sometimes 11 meters high, the young branches 
densely and minutely puberulent; leaves short-petiolate, lance-oblong, mostly 7- 



98 FIELDIANA: BOTANY, VOLUME 24 

12 cm. long, cuspidate-acuminate with a long narrow tip, crenulate to almost 
entire, membranaceous, obtuse or rounded at the base and somewhat asymmetric, 
3-nerved from the base, the upper lateral nerves 3-4 pairs, minutely puberulent 
beneath on the nerves, barbate in the nerve axils; racemes simple or branched, 
5-10 cm. long, puberulent; flowers short-pedicellate, green, 2 mm. long, the sepals 
reflexed; stamens 7-9; ovary puberulent. 

A second and undescribed species of Lunania is represented by a 
sterile specimen from Huehuetenango: Steyermark 51160, from Paso 
del Boqueron, along Rio Trapichillo, below La Libertad, 1,200-1,300 
meters. In this the leaf blades are rounded and subcordate at the 
base, and the petioles are densely hirtellous (glabrous in L. mexicana) . 
There can be no doubt as to the reference of the specimen to this 
genus, but since it is sterile it probably should not be given a name. 

OLMEDIELLA Baillon 

Dioecious trees; leaves persistent, coriaceous, usually with coarse spine-tipped 
teeth, bearing 2 large glands on the upper side at the base of the blade, penni- 
nerved; flowers small, unisexual; sepals of staminate flowers 14-15, acute, imbri- 
cate; petals none; stamens very numerous, the filaments slender, the anthers small, 
ovate, longitudinally dehiscent; sepals of pistillate flowers 7-9; petals none; nec- 
taries forming 1-3 rows about the ovary, with staminodia between them; ovary 
superior, with 6-8 imperfect cells; style short, the stigmas 6-8, sulcate above; 
fruit baccate, depressed-globose. 

The genus consists of a single species. 

Olmediella Betschleriana (Goepp.) Loes. Notizbl. Bot. Gart. 
Berlin 4: 181. 1906. Ilex Betschleriana Goepp. Del. Sem. Vratisb. 
1852; Linnaea 26: 745. 1853-54. 0. ilicifolia Baill. Bull. Soc. Linn. 
Par. 1: 253. 1880. 0. Cesatiana Baill. loc. cit. Licopolia sincephala 
Rippa, Bull. Orto Bot. Napoli 2: 74. 1904. Croton Tuerckheimii 
Donn.-Sm. Bot. Gaz. 54: 242. 1912 (type from Tactic, Alta Verapaz, 
Tuerckheim II. 2163) . Manzanote; manzana; semito; roble de montana. 

Dense, wet or moist, mountain forest, 1,500-2,700 meters; Alta 
Verapaz; Zacapa; Chiquimula; El Progreso; Jalapa; Guatemala; 
Sacatepe"quez; Chimaltenango; Solola; Suchitepe'quez; Quezalte- 
nango; San Marcos; Huehuetenango. Mexico (Chiapas); El Salva- 
dor; Honduras. 

A glabrous tree, often 15 meters high or more, with a dense or open crown, the 
trunk low or tall, often 35 cm. or more in diameter; leaves coriaceous, on short or 
rather long petioles, mostly oblong or elliptic-oblong and 8-15 cm. long, acute to 
long-acuminate and spine-tipped, rounded or obtuse at the base, lustrous, slightly 
paler beneath, the lateral nerves 6-9 pairs, the margin cartilaginous-thickened, 
coarsely sinuate-dentate, with stiff spine-tipped teeth, or leaves on flowering 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 99 

branches often quite entire; inflorescences few-flowered and inconspicuous, the 
flowers green, the staminate in short racemes, the pedicels 7-14 mm. long; sepals 
cuspidate-ovate, 1.5-2 mm. long, the receptacle 4-6 mm. broad; stamens about 
60-85, the anthers 1 mm. long; fruit depressed-globose, somewhat resembling a 
small apple, green, almost smooth, 5-6 cm. broad, flattened at the apex. 

Called "cumbo de cerro" and "cajdn de burro" in Honduras. In 
some respects this is one of the most interesting trees of Guatemala, 
where it is known generally by the name "manzanote." It has had 
a varied taxonomic history, as the synonymy cited above well indi- 
cates. It has been in cultivation in Europe, particularly Italy, for 
more than 90 years, seeds probably having been sent there from 
Guatemala City by some Italian who admired the handsome holly- 
like foliage of the tree. In Europe, apparently, the tree seldom 
flowers or fruits, and botanists who saw the plants had great diffi- 
culty in guessing their proper relationship. The leaves are so much 
like those of holly that it is not surprising that the species was first 
described under the genus Ilex. The genus was guessed also as Sap- 
ium and Quercus, and it was placed by Baillon in the Moraceae. The 
early history of the tree has been brought together by Loesener (loc. 
cit.), who gave a most satisfactory account also of the true relation- 
ship of the genus, which is with the African genus Doryalis. The 
source of the European plants remained unknown until 1932, when 
the senior author was able to report definitely (Trop. Woods 32: 17) 
that Olmediella was a native of the Guatemalan mountains, on the 
basis of material furnished by don Jorge Garcia Salas, then of the 
Direction General de Agricultura. Manzanote trees grow abundantly 
in many of the Guatemalan mountains at middle or rather high ele- 
vations, and they are particularly plentiful on the slopes of the vol- 
canoes of Pacaya and Acatenango, as well as elsewhere in the central 
region. As seen in the forest, there is little to call attention to 
the tree, except that its spiny leaves are unusual. In the forest the 
tree is often exceedingly tall, of irregular branching, the crown 
high overhead. However, someone must have had some reason for 
introducing it into cultivation long ago, and it has now become one 
of the favorite park and street trees of the central region. Under 
cultivation it has a low trunk and a very dense, rounded crown of 
handsome, deep green leaves. It stands pruning well, and many of 
the street and park trees are trimmed closely, to produce dense glob- 
ular crowns that have a formal and somewhat artificial appearance, 
but are nevertheless very handsome. Almost every child of Guate- 
mala City sees manzanote trees in the course of the day, for they are 
planted in the parks and along the streets, the finest being along the 



100 FIELDIANA: BOTANY, VOLUME 24 

Paseo de la Reforma, the principal boulevard. There are fine indi- 
viduals also in the Parque Central of Antigua, in front of the ancient 
Palace of the Viceroys of Guatemala. The tree well deserves exten- 
sive cultivation in other regions that are cool and moist but not too 
cold. Probably it would succeed in some parts of California and in 
various parts of southern Europe, as well as in cooler regions of the 
West Indies and South America. The leaves are exceedingly vari- 
able in their margins, the great majority of them being coarsely 
spine-toothed so as to resemble closely the common American or 
European hollies, but on fertile branches they often are quite entire 
and so distinct in appearance that one with only a herbarium ac- 
quaintance with specimens would feel sure two distinct species were 
represented. 

PLEURANTHODENDRON L. Wms. 

Small trees. Leaves alternate, petiolate, biglandular at the junction of the 
blade and the petiole or glands obscure; tri-quinqueplinerved at the base, the 
margin entire to serrate, the teeth sometimes gland-tipped. Inflorescence often a 
diffuse panicle or panicles, originating in the axils of the upper leaves or terminal; 
the bracteoles early deciduous and the scar often glandular. Flowers perfect; 
sepals 4-6, valvate; petals 4-6, alternate with the sepals, soon falling; stamens 
numerous; anthers small, bilocular, the cells opening longitudinally. Ovary supe- 
rior, sessile, tomentose, 1-celled, placentae 3-4. Fruit a subcoriaceous capsule. 

The following is the only species of the genus. 

Pleuranthodendron mexicana (Gray) L. Wms. Fieldiana, Bot. 
29: 365. 1961. Banara mexicana Gray, Proc. Am. Acad. 5: 174. 1862 
(type from Mexico). Hasseltia pyramidalis Hemsl. Diag. PI. Mex. 4. 
1878 (type from Mexico). Hasseltia mexicana Standl. Contr. U. S. 
Nat. Herb. 23: 840. 1923. Chichimi; quina (Izabal); madrdn (Gua- 
temala) . 

Wet to dry forests, most common at lower elevations, rarely to 
1,500 m.; Pete"n; Alta Verapaz; Izabal; Escuintla; Guatemala; Chi- 
maltenango; San Marcos. Mexico (San Luis Potosi and Veracruz), 
through Central America to Colombia, Peru and Brazil. Figure 12. 

A large shrub or usually a tree, sometimes 15 meters high with a trunk 35 cm. 
in diameter, the crown usually spreading, the branchlets minutely puberulent at 
first, soon glabrate; leaves on slender, short or elongate petioles, oblong-lanceolate 
to oblong-elliptic or elliptic, mostly 7-15 cm. long, abruptly short-acuminate, ob- 
tuse or acute at the base, with 2 large glands on the upper side of the blade at its 
base, glabrous or nearly so, crenate-serrate, 3-nerved from the base; flowers white 
or cream, in small or large panicles, the branches of the panicle densely and mi- 
nutely puberulent; sepals densely tomentulose, 4 mm. long; fruit subglobose, 




ftfl 



FIG. 12. Pleuranthodendron mexicana. A, Branch with inflorescence; X 1 A. 
B, Flower and bud; X 3. C, Flower opened out; X 3. D, Immature ovary; X 6. 
E, Fruits; natural size. F, Under surface of leaf, showing detail of veins and nerves; 
natural size. G, Upper surface at base of leaf, showing glands; natural size. 

101 



102 FIELDIANA: BOTANY, VOLUME 24 

scarcely dehiscent, covered with a dense, minute, gray or white tomentum, sub- 
globose, 5-6 mm. long. 

This tree is often plentiful in lowland forests, forming a consid- 
erable part of the forest understory. It is called "guatuso" in Hon- 
duras and "pochitoquitto" in Veracruz. The bark is dark brown and 
thin; wood yellowish throughout, rather hard, easy to work and saw. 

This species has been reported as both Hasseltia dioica (Benth.) 
Sleumer and Hasseltiopsis dioica (Benth.) Sleumer, an error for which 
Sleumer is responsible. He assumed that Banara dioica Benth. and 
Banara mexicana Gray were synonymous, when, in fact, they do not 
belong to the same genus; the first belongs to Banara and the second 
to Pleuranthodendron. 

PROCKIA L. 

Shrubs or small trees; stipules often large and foliaceous; petiole usually bear- 
ing 2 glands at the apex; leaves membranaceous, serrate, 3-5 (-7) -nerved at the 
base, penninerved above base; inflorescence fasciculate or short-racemose, terminal; 
sepals 3-4, valvate, persistent; petals as many as the sepals or fewer, smaller than 
the sepals, persistent, sometimes none; stamens numerous, free, inserted on a small 
disk, the anthers small, subglobose; ovary 3-5-celled, many-ovulate, the style sub- 
ulate, entire, fruit baccate, indehiscent, many-seeded; seeds small, obovoid, sur- 
rounded by pulp, with crustaceous testa; endosperm carnose; embryo straight, the 
cotyledons ovate, the radicle straight. 

About 10 species, in tropical America. One additional species 
has been described from Costa Rica and one from Mexico. 

Prockia crucis L. Syst. Nat. ed. 10. 1074. 1759. Cajetillo. 

Moist or dry forest or thickets, sometimes in second growth, 2,400 
meters or less; Izabal; Pete"n; Alta Verapaz; Jalapa; Jutiapa; Santa 
Rosa; Escuintla; Guatemala; Solola; Suchitepe"quez ; Quiche"; Hue- 
huetenango. Western and southern Mexico; British Honduras to 
Panama; West Indies; South America. 

Usually a shrub of 2-4 meters, sometimes a tree of 6 meters, with slender 
branches; stipules large, green, semicordate, dentate; leaves slender-petiolate, 
membranaceous, broadly ovate to elliptic or rounded-ovate, mostly 3-12 cm. long, 
acute or acuminate, rounded or shallowly cordate at the base, coarsely or finely 
serrate, pilose or glabrate; pedicels slender, fasciculate or in short racemes; sepals 
3-4, reflexed in anthesis, about 5 mm. long, broadly ovate, tomentose within; 
petals usually reduced or none; stamens very numerous, yellow, glabrous; fruit 
3-5-celled, subglobose, 6 mm. broad, black at maturity, sparsely pilose. 

The wood is brittle and medium hard. Called "quesadilla" in 
Honduras and "cacho de novillo" in El Salvador. The species has a 
wide range and is variable, but attempts thus far made to segregate 
some of the forms as species have not been successful. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 103 
XYLOSMA Forster 

Dioecious or sometimes polygamodioecious shrubs or trees, usually armed with 
spines, the trunks often bearing large, stout, branched spines or thorns; leaves 
dentate or sometimes entire; flowers very small, whitish or green, usually unisex- 
ual; inflorescence glomerate, fasciculate or short-racemose, in leaf axils or on short 
pseudo-branches; sepals 4-5, usually small and scale-like, imbricate, often ciliate; 
petals none; stamens several to many, often surrounded with a glandular disc or 
interspersed with glandular staminodes; anthers short, versatile; ovary inserted 
upon a glandular, annular disk, the placentae 2, or rarely 3-6, parietal, the ovules 
2 or few; style entire or usually divided, the stigmas dilated or not, rarely subsessile 
and peltate-lobate; fruit baccate, indehiscent, small, 2-8-seeded; seeds obovoid, 
the testa smooth, crustaceous, the cotyledons broad. 

Xylosma is a difficult genus of, usually, dioecious trees and shrubs. 
There are some 40 species recorded from Mexico, Central America 
(including Panama), and the West Indies. Perhaps less than a third 
of those ascribed to Central America are valid species. The genus 
in America is much in need of revision and no satisfactory treatment 
of the species for Guatemala may be written until this is done. The 
species are often exasperatingly variable, but this may be due in part, 
at least, to inadequate understanding of them. Staminate, pistillate 
and polygamodioecious plants apparently may differ slightly from 
one another, in addition to the obvious difference in the flowers. 
The leaves on branches without spines seem to differ somewhat from 
those having coarse, heavy spines and again from those having slen- 
der, needle-like spines. The presence or absence of spines is of little 
or no taxonomic importance except as these may affect the leaves. 
Leaves of two forms are found almost constantly in one species 
(X. anisophyllum) and occur occasionally in others. The texture of 
leaves varies, perhaps partly in response to the ecological situation. 
Pubescence, when found, is often not constant and in many cases 
not critical in classification. 

The bark of some species of Xylosma, is reported to be rich in 
tannin but probably is not used in Guatemala for tanning. The 
wood is yellowish, pinkish or brownish, fine-grained and moderately 
hard but is little used except as fuel. Some species of the genus have 
been used as hedges, for which they are highly effective even though 
unsightly and difficult to control. 

It was found difficult to construct a workable key to even the 
eight species to which those in Guatemala have been reduced, prob- 
ably because too many species are still maintained. The key below, 
as well as the treatment of the species, should be considered to be 
provisional. 



104 FIELDIANA: BOTANY, VOLUME 24 

Leaves velutinous to pilose, at least on the lower surface. 

Leaves evidently 3-nerved at the base X. trinervium. 

Leaves not evidently 3-nerved at the base X. velutinum. 

Leaves glabrous (or essentially so). 

Flowers borne in short racemes X. quichense. 

Flowers borne in fascicles. 

Leaves, or most of them, borne in fascicles, usually at the ends of very short 

pseudo-branches X. anisophyllum. 

Leaves not in fascicles, scattered along the branches. 

Leaf-blades lanceolate or oblong-lanceolate, usually abruptly caudate- 
acuminate, mostly 12-15 cm. long X. chloranthum. 

Leaves mostly not as above, mostly 3-9 cm. long. 
Leaves sessile or essentially so, the petiole usually 2 mm. or shorter, 
veins about 6 pairs and the nerves between them very obscure. 

X. sessile. 
Leaves with petioles more than 2 mm. long, both veins and nerves 

prominent. 
Leaves usually rounded or obtuse at the apex, usually very obtuse or 

even rounded at the base X. celastrinum. 

Leaves mostly acute or even acuminate at the apex, the base cuneate 
to rounded X. flexuosum. 

Xylosma anisophyllum Standl. Field Mus. Bot. 11: 135. 1932. 

Moist or wet thickets or savannas, rarely in pine-oak forest, 1,900 
meters or usually at low elevations; Pete"n; Alta Verapaz; Guate- 
mala. Mexico (Campeche, the type from Tuxpena); British Hon- 
duras; Panama. 

A much-branched, stiff shrub of 1.5-2.5 meters, or sometimes a small tree, 
glabrous throughout, the branches mostly unarmed but frequently bearing stout 
spines 1-1.5 cm. long; leaves borne in fascicles or usually so, coriaceous, on stout 
petioles 2-4 mm. long, very variable upon a single plant, narrowly oblanceolate- 
oblong to rhombic-lanceolate, obovate, or broadly elliptic-ovate, as much as 9 cm. 
long and 3-5 cm. wide, obtuse or rounded at the apex, cuneate-attenuate at the 
base or sometimes obtuse, remotely crenate-serrate with appressed teeth or sub- 
entire, lustrous above, the veins scarcely prominulous, the veins prominulous and 
closely reticulate beneath; flowers numerous, in the leaf axils or at defoliate nodes, 
fasciculate, the slender pedicels glabrous, 5-10 mm. long, or often very short; sepals 
oblong or lance-oblong, obtuse, 1 mm. long, ciliate, spreading, deciduous; fruit 
glabrous, subglobose, 5-6 mm. long; seeds about 5, ferruginous. 

This species seems reasonably distinct by reason of the fascicu- 
lated leaves which are borne on the same pseudo-branch as the fas- 
cicle of flowers. The disjunct range is of interest. 

Xylosma celastrinum (HBK.) Standl. & Steyerm. Field Mus. 
Bot. 23: 176. 1944. Flacourtia celastrina HBK. Nov. Gen. & Sp. 7: 
239. 1825. Putzil (Huehuetenango) . 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 105 

Moist or dry, brushy plains or hillsides, sometimes on limestone, 
150-1,650 meters; Jalapa; Santa Rosa; Huehuetenango. Southern 
and western Mexico, the type from Acapulco. British Honduras 
and Honduras. 

A shrub or small tree of 2-6 meters, the stiff branches armed with stout, 
straight spines 1-2 cm. long, the whole plant glabrous throughout or nearly so; 
leaves on short petioles mostly 2 mm. long or shorter, elliptic to rhombic-ovate or 
cuneate-obovate, mostly 3-5 cm. long and 1.5-2.5 cm. broad, rounded to obtuse 
or sometimes subacute at the apex, usually rounded or very obtuse at the base, 
coarsely crenate or crenate-serrate, chartaceous, often lustrous, the veins promin- 
ulous and reticulate on both surfaces; flowers fasciculate, short-pedicellate, the 
pedicels glabrous; sepals small, green, oblong or ovate, glabrous, white-ciliate; 
style very short, the stigmas short and broad; fruit subglobose, 5-6 mm. in diam- 
eter, glabrous, red or at maturity black; seeds 2-4. 

Called "aguja, de arrea" in El Salvador. This species is closely 
allied to X. flexuosum (HBK.) Hemsl. and perhaps not really dis- 
tinct from it. Some of the forms segregated here also are very close 
to X. anisophyllum Standl. 

Xylosma chloranthum Donn.-Sm. Bot. Gaz. 57: 415. 1914 
(type from Guatemala, Tuerckheim 4111). X. sylvicola Standl. Field 
Mus. Bot. 4: 316. 1929 (type from Honduras, Standley 52917). Santa 
Rosa; Alta Verapaz; Pete*n. British Honduras; Honduras. 

A shrub or small tree, glabrous throughout, the trunk armed with large, 
branched spines; leaves on slender petioles 5-7 mm. long, pergamentaceous, lan- 
ceolate or lance-oblong, mostly 12-15 cm. long and 3-4 cm. broad, narrowly 
caudate-attenuate or acuminate, acute or acuminate at the base, crenate-serrate, 
concolorous, lustrous on both surfaces, especially above; flowers numerous, fascicu- 
late, the pedicels 2-2.5 mm. long or longer; pistillate sepals 4, green, 2 mm. long, 
glabrous; ovary 4-celled, glabrous, the placentae 2-ovulate. 

Very closely allied to the X. flexuosum complex but perhaps 
distinct. 

Xylosma flexuosum (HBK.) Hemsl. Biol. Cent. Am. Bot. 1: 
57. 1879. Flacourtia flexuosa HBK. Nov. Gen. & Sp. 7: 239. 1825. 
Xylosma Paliurus (Clos) Briq. Ann. Cons. Jard. Ge"neve 4: 221. 1900. 
Xylosma elliptica (Clos) Hemsl. Biol. Cent. Am. Bot. 1: 57. 1879. 
Alfiler; pico de burridn; pico de gurridn; alfilerillo; escoba blanca de 
montana. 

Moist or dry, often rocky thickets or mixed forest, chiefly on 
mountain sides or sometimes on lowland plains, occasionally in pine- 
oak forests. Often invading overgrazed lands. Lowlands up to about 
2,400 meters, in all or almost all departments. Mexico to Nicaragua 
and possibly to Panama and on into South America. 



106 FIELDIANA: BOTANY, VOLUME 24 

Shrubs or small trees sometimes to 6 m. tall but generally less, glabrous or 
sometimes the branches and leaves pubescent or puberulent, the branches stiff 
and stout or sometimes slender and flexuous, unarmed or armed with slender or 
with stout and coarse spines, the trunks commonly with large and much-branched 
spines or thorns; leaves mostly short-petiolate, thick-membranaceous to subcoria- 
ceous, rhombic-ovate to elliptic-oblong or elliptic-ovate, rarely obovate, usually 
broadest at or below the middle, commonly acute or acuminate, the base cuneate 
to rounded, serrate to subentire, often lustrous, the veins usually prominent and 
reticulate; flowers fasciculate in the leaf axils, on glabrous or puberulent pedicels 
1.5-5 mm. long; sepals oblong or narrower, obtuse or acute, spreading, glabrous 
or pubescent dorsally, ciliate or eciliate; fruits subglobose, 5-6 mm. in diameter 
or sometimes larger, bright red when mature. 

Called "aguja de arrea" and "pepenance" in El Salvador and 
"aguja de ara" in Honduras. This species as reported here is pos- 
sibly an aggregate. Considering the present state of knowledge of 
this genus there seems no point in attempting to separate subspecific 
taxa. In the sense that the species is described here it may include 
as well Xylosma Hemsleyanum Standl., X. Seemannii Tr. & PI., 
X. panamensis Turcz., and possibly X. prunifolium (HBK.) Griseb. 
from South America. Species closely allied but kept apart in this 
flora are X. sessile Standl. & Steyerm., X. chloranthum Donn.-Sm., 
X. celastrinum (HBK.) Standl. & Steyerm., and X. velutinum (Tul.) Tr. 
&Karst. 

Hedges of this species, or a related one, are reported planted in 
the Jocotan region and in Jalapa, where they are called "aguja, de 
arrea." The wood is valued there because it burns slowly and steadily. 

Xylosma quichense Donn.-Sm. Bot. Gaz. 18: 197. 1893. X. qui- 
chense var. subalpinum Donn.-Sm. op. cit. 198 (type collected near 
El Jute, Quiche", Heyde & Lux 3064). 

Moist or wet, mixed or oak forest, 1,600-3,000 meters; endemic; 
Baja Verapaz; El Progreso; Jalapa; Chimaltenango; Quiche" (type 
from San Miguel Uspantan, Heyde & Lux 3066); Huehuetenango; 
San Marcos; Solola. 

A shrub or often a tree of 2-8 meters, rarely 12-15 meters high, the branches 
unarmed or bearing rather long, slender, stiff spines, pubescent or almost wholly 
glabrous, the trunk usually armed with very large, stout, much-branched, brown 
spines; leaves short-petiolate, thick-membranaceous or subchartaceous, oblong- 
elliptic to ovate-oblong, mostly 6-12 cm. long, acuminate or rather abruptly so, 
rounded or obtuse at the base, serrate, the teeth often close, elongate, and incurved 
or salient, the veins usually prominent and reticulate, often lustrous; flowers race- 
mose, the racemes solitary or fasciculate, short and few-flowered, sometimes 
branched; sepals 4-6, at an thesis broadly ovate, becoming elliptic-oblong, green, 
as much as 4 mm. long, glabrous, not ciliate; stamens 20-35, not exceeding the 
sepals; fruit globose, glabrous, 5-6 mm. in diameter, often bright red. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 107 

The species is allied to the main Central American complex 
around Xylosma flexuosum but is easily distinguished by the broad 
sepals and racemose inflorescence. 

Xylosma sessile Standl. & Steyerm. Field Mus. Bot. 23: 177. 
1944. 

Moist or wet forest or thickets, often or perhaps usually on lime- 
stone, 1,500 meters or less; endemic; Alta Verapaz (type from Coban, 
Tuerckheim 11.1617; collected also in the region of Chelae); Izabal 
(Los Amates). 

A glabrous shrub, the branches ferruginous or brown, unarmed or bearing 
stout spines 1-2 cm. long; leaves almost sessile, on petioles 2 mm. long or usually 
shorter, coriaceous, elliptic-oblong or lance-oblong, 6.5-8.5 cm. long, 2-3 cm. 
broad, acute or short-acuminate, cuneate-acute at the base, remotely appressed- 
serrate or almost entire, often lustrous above, the nerves prominulous, the veins 
obsolete, slightly paler beneath, usually brownish when dried, the lateral nerves 
about 6 pairs, prominulous, the veins obsolete or obscure; flowers densely fascicu- 
late in the leaf axils, the slender pedicels 5-7 mm. long, glabrous; sepals oblong- 
ovate, 1.5 mm. long, obtuse, glabrous, ciliate; stamens numerous, the filaments 
glabrous, 2.5-3 mm. long; fruit subglobose, 5-7 mm. in diameter, glabrous; style 
very short, the stigmas short, ovate. 

This species is a part of the Central American complex which 
centers around X. flexuosum and perhaps should be referred there. 

Xylosma trinervium Standl. & Steyerm. Field Mus. Bot. 23: 
178. 1944. 

Known only from the type, Huehuetenango, along Rio Cuilco 
between Cuilco and San Juan, 1,200-1,300 meters, Steyermark 50876. 

A slender shrub 1.5-3 meters high, the branches tortuous, terete, brown, when 
young densely pilosulous with short spreading whitish hairs, unarmed; leaves 
small, thick-membranaceous, petioles 3-4 mm. long, elliptic or lance-elliptic, wid- 
est near the middle, 4-5.5 cm. long, 2-2.8 cm. broad, acute or acuminate, cuneate- 
acute or subobtuse at the base, conspicuously 3-nerved from the very base, coarsely 
crenate or crenate-serrulate, densely velutinous-pilose on both surfaces with spread- 
ing grayish hairs, the lateral nerves about 4 pairs, ascending at a very narrow angle; 
flowers unisexual, the staminate in short racemes, the racemes subterminal, soli- 
tary, subsessile, densely few-flowered, about 1 cm. long, the pedicels short, pilosu- 
lous; sepals oval or rounded, green, slightly more than 1 mm. long, rounded at the 
apex, spreading, long-ciliate, sparsely pubescent or glabrous dorsally; stamens 7-8, 
slightly longer than the sepals, glabrous; pistillate flowers unknown. 

Xylosma velutinum (Tulasne) Triana & Karst. ex Karst. Fl. 
Colomb. 1: 123, t. 97. 1858. Flacourtia velutina Tulasne, Ann. Sci. 
Nat. ser. Ill, 7: 295. 1847. Myroxylon velutinum Warb. in Engler & 



108 FIELDIANA: BOTANY, VOLUME 24 

Prantl, Pflanzenfam. 3, Abt. 6a: 41. 1893. Xylosma characanthum 
Standl. Field Mus. Bot. 11: 136. 1932 (type from British Honduras, 
Bartlett 12004). Corona; copalin; tira-zapato; puntezapat; aguja rabia; 
aguja de ara. 

Mostly in dry, rocky thickets or on forested hillsides, rarely in 
moist or wet places, occasionally forming large and dense thickets; 
Santa Rosa; Escuintla; Chimaltenango; Guatemala; Retalhuleu; 
Quezaltenango; Huehuetenango. Mexico; El Salvador; Honduras; 
Nicaragua; Costa Rica; Colombia; Ecuador. 

A shrub or tree, sometimes 10 m. tall, the trunk often armed with large, much- 
branched spines or sometimes unarmed; the branches slender and flexuous to thick 
and stout, unarmed or armed with numerous stout to slender spines, when young 
densely pilosulous with pale spreading hairs or almost glabrous; leaves short- 
petiolate, rather thin to coriaceous, elliptic to rounded or obovate, mostly 3.5- 
6 cm. long and 2-4 cm. broad, acuminate to very obtuse at the apex, cuneate to 
rounded at the base, conspicuously crenate-serrate to almost entire, densely velu- 
tinous-pilosulous on both surfaces or glabrous above and sparsely pilose to villous 
beneath; flowers fasciculate in the leaf axils, the pedicels usually short, pilosulous; 
fruit globose, 5-6 mm. in diameter, often bright red, at maturity black, usually 
glabrous; seeds 2-4. 

An exceedingly variable species, similar in many of its forms to 
those of X. flexuosum, to which it is closely related. 



ZUELANIA A. Richard 

Trees or large shrubs; leaves stipulate, pellucid-punctate, alternate, entire or 
serrate, pubescent, penninerved; flowers small, perfect, densely fasciculate, the 
pedicels articulate, bracteate at the base; calyx 4-5-lobate, the lobes imbricate; 
petals none; stamens 20-40, alternating with fleshy staminodia; ovary free, 1-celled, 
many-ovulate, with 3 parietal placentae; stigma sessile or subsessile, peltate; fruit 
large, capsular, fleshy, berry-like, at length opening by valves; seeds numerous, 
arillate, the endosperm carnose. 

Three species, the others West Indian. 
The genus is hardly distinct from Casearia. 

Zuelania Guidonia (Swartz) Britton & Millsp. Bahama Fl. 295. 
1920. Laetia Guidonia Swartz, Prodr. Veg. Ind. Occ. 83. 1788. 
Z. Roussoviae Pittier, Contr. U. S. Nat. Herb. 18: 163, t. 79. 1916 
(type from Panama). Palacio (Izabal); quacap (Alta Verapaz); 
tamay (Pete"n, Maya) ; trementino (Pete"n) . 

Moist or wet thickets or in open forest, often in pine forest; 
Pete"n; Izabal; Alta Verapaz. Veracruz and Oaxaca to Yucatan; 
British Honduras along the Atlantic coast to Panama; West Indies. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 109 

Sometimes only a large shrub but usually a tree, often 10-25 meters high, the 
trunk 30-60 cm. in diameter, the crown rounded or spreading, the bark light or 
dark grayish to very dark brown, moderately smooth, the inner bark deep pink- 
ish, yielding a sticky transparent sap when cut; young branchlets densely brownish- 
hirsute; leaves deciduous, often clustered at the ends of the branches, on petioles 
about 1 cm. long, broadly to narrowly oblong, 15-25 cm. long, acute or acuminate, 
obtuse to shallowly cordate at the base, inconspicuously serrate, densely veluti- 
nous-pilose beneath, glabrate above in age and often lustrous; flowers often ap- 
pearing before the leaves, cream-colored, in large, very dense clusters at the ends 
of the branchlets or in lateral clusters, the pedicels mostly 10-15 mm. long; sepals 
7 mm. long, sericeous outside, broadly rounded at the apex; stamens mostly 20- 
25 but sometimes more, the filaments glabrous or sparsely pilose; ovary densely 
pilose; fruit globose, shallowly 3-sulcate, 3.5 cm. in diameter or larger, yellowish 
green, with soft flesh, juicy; seeds obovoid, angulate, 4 mm. long. 

Called "water-wood" in British Honduras; "resina," "sangre de 
playa" (Honduras); "volador" (Yucatan); "manzano," "manzanillo," 
"campanillo," "tepecacao" (Oaxaca, Veracruz). The wood is yellow 
throughout, moderately hard, tough, fine-textured, not durable. It 
is little used, except in some regions for construction purposes. 

TURNERACEAE 

Herbs, shrubs, or trees, glabrous or pubescent, the pubescence of simple or 
branched hairs; leaves alternate, petiolate, simple, usually serrate, often 2-gland- 
ular at the base; stipules small or none; flowers regular, perfect, small or large, 
axillary, solitary or few, sessile or pedunculate, rarely racemose, the peduncles free 
or connate with the petiole, often articulate, frequently 2-bracteolate; calyx tu- 
bular, 5-fid, deciduous, the lobes imbricate; petals 5, most often yellow, inserted in 
the throat of the calyx, membranaceous, contorted in bud, deciduous, naked or 
sometimes bearing a fimbriate scale at the apex of the claw; stamens 5, inserted 
at the base, middle, or throat of the calyx tube, rarely hypogynous, the filaments 
free, complanate-subulate; anthers oblong, the cells introrsely dehiscent; ovary free, 
ovoid or elongate, 1-celled, the styles 3, terminal, filiform, simple or 2-fid; stigmas 
flabellately multifid, rarely simply dilated; ovules numerous, 2-seriate on 3 parietal 
placentae, ascending, anatropous; capsule 1-celled, 3-valvate at the apex or for its 
whole length, the valves bearing the placentae along their middle; seeds oblong- 
cylindric, slightly curved, arillate, the testa subcrustaceous, foveolate; endosperm 
abundant; embryo terete, axile, orthotropous, the cotyledons plano-convex, the 
radicle terete. 

Eight genera, in tropical or warm regions of both hemispheres. 
Only the following are represented in North America. 

Flowers large, mostly 6-7 cm. long; petals bearing a fimbriate scale at the apex of 

the claw; stigmas entire; large trees Erblichia. 

Flowers much smaller; petals naked; stigmas fimbriate; herbs or low shrubs. 
Receptacle of the flower bearing a corona; pubescence stellate; low herbs. 

Piriqueta. 
Receptacle without a corona; pubescence of simple hairs; herbs or shrubs. 

Turner a. 



110 FIELDIANA: BOTANY, VOLUME 24 

ERBLICHIA Seemann 

Reference: Standley & Steyermark, Notes on the American spe- 
cies of Erblichia, Field Mus. Bot. 22: 351-357. 1940. 

Large trees; leaves alternate, petiolate, serrate; stipules minute; flowers large, 
yellow or orange-yellow, the peduncles articulate below the middle, 2-bracteolate, 
the bractlets subfoliaceous, serrate; sepals lanceolate, colored; petals hypogynous, 
obovate-spatulate, unguiculate, the claw bearing a fimbriate appendage at its apex; 
stamens 5, hypogynous, the anthers linear-oblong; ovary oblong, the styles 3, the 
stigmas dilated and subclavate; capsule oblong, pubescent or glabrous, 3-valvate, 
many-seeded; seeds ovate-oblong, straight. 

Erblichia was united to Piriqueta by Urban, and in this he was 
followed by Gilg in his account of the family in Naturliche Pflanzen- 
familien. There seems to be adequate reason to maintain the genus. 
The wood in this genus is pale brownish, of medium density but 
hard and tough; the texture is fine, the grain straight to irregular. 
So far as we know no use is made of it. We have not seen the 
African species that have been associated with Erblichia and do not 
know what their status may be. 

Leaves glabrous or essentially so below E. odorata. 

Leaves velutinous below E. odorata var. mollis. 

Erblichia odorata Seem. Bot. Voy. Herald 130, t. 27. 1854 (type 
from Panama) ; Piriqueta xylocarpa Sprague & Riley, Kew Bull. 1923: 
373. 1923 (type from British Honduras); E. xylocarpa Standl. & 
Steyerm. Field Mus. Bot. 22: 353. 1940; E. Standleyi Steyerm. 
Field Mus. Bot. 22: 353. 1940 (type from Honduras). Condp; cortez; 
candillaria. 

Occasional in wet forest to about 900 m.; Alta Verapaz; Pete"n; 
Izabal. Mexico, Central America (except Nicaragua), and Panama. 

A tall tree to 40 m., but usually much less, with a trunk 50 cm. in diameter; 
leaves short-petiolate, lance-oblong or elliptic-oblong, mostly 7-14 cm. long, acute 
or acuminate at each end, appressed-serrate or crenate, glabrous and lustrous 
above, glabrous beneath or nearly so, often with appressed hairs along the costa 
above; sepals lanceolate or linear-lanceolate, 5-5.6 cm. long, long-acuminate or 
caudate, usually yellow-green with salmon or pale orange margins, somewhat 
appressed-pilosulous outside; petals yellow, bright orange or orange-salmon, cune- 
ate-oblong, cuspidate or caudate, 6-8 cm. long, 3.5-4.5 cm. broad; anthers 4-6 mm. 
long; ovary glabrous to densely velutinous with short ascending stiff yellow- 
brown hairs. 

Called "flor de fuego" in El Salvador; "butterfly tree" in British 
Honduras. When in flower it is an outstanding tree, covered with 
brilliant flowers, and often towering high above the surrounding 
trees. It blooms usually in March and April. 




FIG. 13. Erblichia odorala var. mollis. Branch, showing open flower and buds. 



Ill 



112 FIELDIANA: BOTANY, VOLUME 24 

Erblichia odorata var. mollis (Standl. & Steyerm.) L. Wms. 
Fieldiana: Bot. 29: 368. 1961. E. xylocarpa var. mollis Standl. & 
Steyerm. Field Mus. Bot. 22: 355. 1940. Candellaria de montana; 
polo de mora. 

Moist highland forests, 1,200-1,500 meters; Solola; Suchitepe- 
quez; Quezaltenango; San Marcos. Mexico. Figure 13. 

A tall tree differing from the typical variety principally in the 
velutinous pubescence of the under surface of the leaves. 

PIRIQUETA Aublet 

Usually annual herbs, abundantly stellate-pubescent, some of the hairs often 
simple; leaves alternate, dentate or entire, rarely pinnatifid, often 2-glandular at 
the base; flowers yellow, axillary or in greatly reduced panicles; sepals 5, imbri- 
cate in bud; petals usually yellow, sometimes purple; filaments glabrous or pubes- 
cent; anthers emarginate at the base, truncate or emarginate at the apex, rarely 
mucronate; stigmas 3- and 2-cleft; capsule loculicidally 3-valvate; seeds numer- 
ous, curved or almost straight, the testa roughened. 

About 20 species, mostly American, a few in Asia. Probably 
only one occurs in Central America. 

Piriqueta cistoides (L.) Mey. ex Steud. Nom. Bot. ed. 2. 2: 344. 
1841. Turnera cistoides L. Sp. PL ed. 2. 387. 1762. 

Mostly in savannas, sometimes on open rocky hillsides, 1,400 
meters or less; Pete'n; Alta Verapaz; Chiquimula; Jalapa; Jutiapa; 
Huehuetenango. Mexico; British Honduras to Panama. West In- 
dies; South America. 

An erect annual or sometimes longer lived, simple or branched, the stems 
puberulent and also hirsute with long stiff spreading brownish hairs; leaves linear- 
lanceolate to ovate-lanceolate, short-petiolate or the upper ones sessile, 3-9 cm. 
long, acute or obtuse, obtuse or acute at the base, undulate-dentate or crenate or 
almost entire, usually densely stellate-pubescent on both surfaces; flowers axillary, 
solitary, the peduncles 3 cm. long or shorter; sepals lanceolate, acute, 4-9 mm. 
long; petals obovate, orange or yellow to almost white; filaments glabrous; cap- 
sule ellipsoid or globose, 4.5-9 mm. long, densely short-pilose; seeds oblong- 
obovoid, almost straight or curved, 1.5-1.8 mm. long, foveolate-striate. 

A typical savanna plant of Central America, but sometimes found 
in other situations. It is not at all showy, for the flowers are small 
and few, although they are brightly colored. 

TURNERA L. 

Herbs or low shrubs, the pubescence of simple hairs; leaves dentate; flowers 
usually yellow, solitary, axillary, often showy, the peduncles often adnate to the 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 113 

petioles; petals obovate or spatulate; stamens inserted below the petals, sometimes 
hypogynous; styles 3, filiform, the stigmas cleft few or many times; capsule 3-val- 
vate to the base, usually many-seeded; seeds obovoid to oblong or cylindric, curved. 

Species about 60, all except two in tropical America, the others 
African. One species is known from Costa Rica and Panama. 

Peduncles free from the petioles; shrubs, woody almost throughout. 

Seeds globose-obovoid, little longer than broad T. odorata. 

Seeds cylindric, twice as long as broad or longer T. diffusa. 

Peduncles, at least most of them, adnate to the petioles, the flowers appearing to 
arise from the petiole; annual or perennial herbs, herbaceous or sometimes 
woody below. 

Plants annual ; flowers in dense leaf y-bracted clusters at the ends of the branches. 

T. Pumilea. 
Plants usually perennial; flowers mostly in the axils of scattered leaves. 

T. ulmifolia, 

Turnera diffusa Willd. ex Schult. Syst. 6: 679. 1820. T. aphro- 
disiaca Ward, Virginia Med. Monthly 49. 1876. T. diffusa var. 
aphrodisiaca Urban, Jahrb. Bot. Gart. Berlin 2: 127. 1883. Mejorana 
(Huehuetenango) . 

Mostly in dry rocky open places, sometimes on limestone or along 
cliffs or in pine-oak forest, 500-1,950 meters; Chiquimula; Baja Vera- 
paz; Quich^ ; Huehuetenango. Mexico; British Honduras; Honduras; 
Nicaragua; Costa Rica; Panama; West Indies; South America. 

A shrub, often densely branched, a meter high or less, usually densely pilose 
or pubescent throughout; leaves very variable in shape and size, oblanceolate to 
spatulate or oblong-ovate, mostly 1-2 cm. long, coarsely serrate or crenate, the 
nerves impressed above, often densely soft-pilose beneath, cuneate at the base; 
peduncles very short, the flowers often almost sessile; calyx 5-dentate; petals 
spatulate, yellow, 4-8 mm. long, much exceeding the calyx teeth; capsule sub- 
globose, 1.5-2 mm. in diameter. 

Known in Mexico as "damiana"; "misibcoc, chac-mixib" (Yuca- 
tan, Maya) . The plant is uncommon in Guatemala. In many parts 
of Mexico it is a common weedy plant, and it is well known there 
because of its reputed medicinal properties. It is particularly famed 
as an aphrodisiac, although it has not been proved that the plant 
has such properties to any great degree. It has an aromatic odor and 
a rather agreeable taste, and contains a volatile oil. The dried leaves 
are used in Mexico sometimes as a substitute for Chinese tea, for 
flavoring liquors and in domestic medicine for various purposes, 
chiefly as a tonic or for treating colds, intestinal infections and vene- 
real diseases. The dried leaves are sold sometimes in Guatemalan 
markets for medicinal purposes. Quantities of the plant have been 



114 FIELDIANA: BOTANY, VOLUME 24 

exported at various times to the United States and Europe for use in 
medicine, although it has never been an official drug, at least in the 
United States. The plant grows in Guatemala, so far as known, 
only at high elevations, but it is to be expected in the lowlands of 
Pete"n. In British Honduras it is found at or little above sea level. 
The species is a variable one in foliage and pubescence. 

Turnera odorata L. Rich. Act. Soc. Hist. Nat. Paris 1: 107. 
1792. 

In pine ridge, at or little above sea level. British Honduras; 
South America. 

A branched shrub, woody throughout, 1-2 meters high, the young branches 
densely short-pilose; leaves short-petiolate, ovate or broadly ovate, mostly 1.5- 
7 cm. long and 1-3 cm. broad, acute or obtuse, very obtuse to cuneate at the base, 
coarsely crenate or crenate-serrate, densely short-pilose on both surfaces, some- 
times tomentose beneath; flowers scattered in the leaf axils, often dimorphous, 
the peduncles 4 mm. long or shorter; calyx usually 8-11 mm. long, pubescent or 
glabrate, the lobes lanceolate or lance-linear; petals yellow, 6-8 mm. long; fruit 
globose, 4-5.5 mm. long, pubescent; seeds globose-obovoid, 1.5 mm. long, longi- 
tudinally striate. 

The shrub appears to be common in the pinelands of British 
Honduras, but it is not known in other parts of the North American 
mainland. 

Turnera Pumilea L. Syst. ed. 10. 965. 1759. 

Open, usually dry and rocky, grassy or brushy slopes, sometimes 
in pine forest, 400-1,200 meters; Zacapa; Chiquimula; Jalapa; Juti- 
apa; Huehuetenango. British Honduras; Honduras; Nicaragua; 
Mexico; West Indies; South America. 

An erect or decumbent annual or perennial, the stems 10-25 cm. long, usually 
branched from the base and often also above, hirsute; leaves short-petiolate, obo- 
vate or oblanceolate to ovate or ovate-lanceolate, mostly 2-5 cm. long, acute or 
subacute, cuneate at the base, coarsely serrate, hirsute with long spreading hairs; 
flowers crowded in many-flowered dense clusters at the ends of the branches, the 
clusters very densely hirsute; calyx 6-7 mm. long; petals slightly longer than the 
sepals, yellow; seeds obovoid-oblong, curved, the testa deeply foveolate. 

In Mexico the perennial phase of this species has been described 
as Turnera Palmeri S. Wats. 

Turnera ulmifolia L. Sp. PI. 271. 1753. Tapalayote (Baja Vera- 
paz) ; escoba, malva cimarrona (Pete"n, fide Lundell) . 

Savannas or stony fields or hillsides, often on open banks or a 
weed in waste ground, sometimes on limestone, 1,200 meters or less; 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 115 

Izabal; Pete"n; Alta Verapaz; Baja Verapaz; El Progreso; Zacapa; 
Chiquimula; Jutiapa; Escuintla; Guatemala; Huehuetenango. Mex- 
ico; British Honduras to El Salvador and Panama; West Indies; 
South America. 

An erect herb or shrub, usually 75 cm. high or less, densely pilose throughout 
with pale, spreading or sometimes appressed hairs; leaves petiolate, lanceolate to 
oblong-ovate, mostly 3-10 cm. long, acute or acuminate, usually attenuate to the 
base and conspicuously 2-glandular, serrate or crenate-serrate; peduncles united 
with the petiole, the flower thus appearing to be sessile on the petiole; calyx deeply 
5-lobate, the segments lanceolate, acute to long-acuminate; petals deep or bright 
yellow, 1-3 cm. long; capsule 6-10 mm. long, 3-valvate above or almost to the base, 
densely pilose; seeds short-cylindric, ochraceous, densely foveolate. 

Known in El Salvador by the names "flor de San Lorenzo," "esco- 
billo," and "escobillo bianco;" "clavel de oro," "calendula" (Yucatan). 
The plant is showy when in full flower, but in general it is rather 
weedy. Through its wide range it exhibits a great deal of variation, 
but the Guatemalan material is fairly uniform. 



PASSIFLORACEAE. Passion-flower Family 

References: E. P. Killip, Passifloraceae of the Mayan region, 
Carnegie Inst. Wash. Publ. 461: 301-328, t. 1,2. 1936; The Amer- 
ican species of Passifloraceae, Field Mus. Bot. 19: 1-613. 1938. 

Scandent herbs or shrubs, with tendrils, or rarely erect shrubs or trees; leaves 
alternate, simple, entire or lobate, very rarely compound, the petiole usually bearing 
glands; stipules 2, filiform to ovate; peduncles usually geniculate; bracts and 
bractlets 3, small and distant from the flower, or large, foliaceous, and close to 
the flower and forming a kind of involucre; tendrils axillary, not branched; flowers 
axillary, perfect or rarely unisexual, regular, solitary, cymose-racemose or cymose- 
paniculate; calyx tube receptacle almost flat to saucer-shaped or campanulate, 
bearing in the center a gynophore; sepals 5, inserted in the throat of the receptacle, 
imbricate, often with a small horn-like process dorsally near the apex, usually 
colored inside; petals none or as many as the sepals and alternate with them, free, 
imbricate, withering, persistent; corona present, inserted in the throat or tube of 
the receptacle, 1 -several-seriate, divided into radiate or erect filaments, sometimes 
membranous or wanting; stamens mostly 5; ovary superior, free, stipitate, 1-celled; 
style simple or with 3-4 branches or the styles usually 3; ovules generally numerous, 
pendulous, attached to 3 or 5 parietal placentae; fruit more or less baccate, or 
sometimes capsular and opening by 3 apical valves; seeds numerous, usually ovoid, 
compressed, covered with a fleshy aril or surrounded by pulp; endosperm carnose; 
embryo rather large, the cotyledons often foliaceous; radicle straight. 

About a dozen genera, in both hemispheres. Most of the mem- 
bers of the family belong to the genus Passiflora. Three other genera 
are known from tropical America, but only one of them, Tetrastylis, 
extends to Central America (Costa Rica). 



116 FIELDIANA: BOTANY, VOLUME 24 

PASSIFLORA L. Passion-flower 

Herbaceous or woody vines or rarely erect shrubs or small trees; leaves alternate, 
petiolate, simple or compound, often lobate; inflorescences axillary, simple or 
compound; bracts small and remote from the flower or large and surrounding the 
flower as an involucre; flowers perfect; calyx tube patelliform to campanulate 
or cylindric; sepals fleshy or membranaceous, often corniculate or arista te dorsally 
just below the apex; petals 5, membranous, sometimes wanting; corona of 1-several 
series of distinct or more or less united filaments, rarely tubular; operculum borne 
within or below the corona, membranous, flat or plicate, entire, lacerate, or fila- 
mentose, rarely wanting; stamens 5, the filaments united to form a tube, this 
closely adnate to the gynophore, the free filaments erect at first, at length reflexed, 
attached dorsally to the anther near its middle; anthers linear, ovate, or oblong, 
2-celled; ovary globose, ovoid, or fusiform, borne on a gynophore, this usually 
elongate; styles 3, distinct or united at the base, cylindric or clavate; stigmas 
capitate, orbicular, or reniform; fruit indehiscent, containing mucilaginous pulp; 
seeds more or less compressed, reticulate, punctulate, or transversely grooved, 
borne on 3 parietal placentae. 

More than 350 species are known from tropical America, and a 
few others occur in Asia and Australia. Two species extend into 
temperate regions of the United States. The plants of this genus 
are easily recognized by their distinctive flowers, of rather compli- 
cated and curious structure. The early Spanish explorers, as well 
as some of the scientists of their day, were greatly impressed by a 
fancied resemblance between the design of the flower and the em- 
blems associated with the Crucifixion; hence the Spanish name 
"pasionaria" applied to the plants, and the Latin term "Passiflora" 
given to the genus. Many of the species are notable for their hand- 
some and showy flowers. The fruits of all species probably are edible. 
Some of the larger ones have excellent quality and flavor, but the 
smaller fruits usually are insipid and seldom are eaten. 

KEY TO PASSIFLORA 

Bracts of the peduncles at least 8 mm. wide, forming an involucre at the base of 
the flower, entire or pinnatisect; flowers usually more than 4 cm. broad when 
expanded, often highly colored. 
Bracts 2-4 times pinnatisect, the segments filiform, gland-tipped. 

Leaves not lobate, coriaceous or subcoriaceous, the nerves impressed on the 

upper surface P. Urbaniana. 

Leaves lobate, usually membranaceous, the nerves not impressed. 
Petals linear, more than twice as long as the outermost filaments of the 

corona; gynophore at least 2 cm. long P. Palmeri. 

Petals oblong or oblong-lanceolate, less than twice as long as the outermost 
filaments of the corona; gynophore less than 2 cm. long. . .P. foetida. 
Bracts entire. 

Leaves deeply lobate, the lobes extending to the middle of the blade or lower. 
Lobes of the leaves obtuse. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 117 

Leaves 3-lobate P. subpeltata. 

Leaves 5-lobate P. caerulea. 

Lobes of the leaves very acute or acuminate. 

Stipules conspicuous, semiovate P. Oerstedii var. choconiana. 

Stipules setaceous or linear, inconspicuous. 

Bracts united for much of their length P. platyloba. 

Bracts free. 

Lobes of the leaves merely acute; bracts 2-2.5 cm. long . . .P. edulis. 
Lobes of the leaves cuspidate-acuminate; bracts 5 cm. long. 

P. prolata. 

Leaves not at all lobate or only very shallowly so. 
Leaves peltate. 

Bracts usually purplish; leaves orbicular in outline, obscurely 3-lobate at 

the apex P. membranacea. 

Bracts green; leaves broadly ovate in outline. 

Stipules crenate; bracts more than 3 cm. long, entire P. Cookii. 

Stipules setiferous-dentate; bracts less than 3 cm. long, denticulate 
toward the base P. Hahnii. 

Leaves not peltate. 

Stems acutely 4-angulate, winged on the angles P. quadrangularis. 

Stems neither angulate nor winged. 

Leaf blades as broad as long or much broader, lunately bilobed or shal- 
lowly 3-lobate at the broad apex. 
Leaves densely velutinous-pubescent, shallowly 3-lobate. 

P. porphyretica. 
Leaves glabrous or nearly so, lunately bilobate P. pulchella. 

Leaf blades much longer than broad, not at all lobate, acute or acu- 
minate. 
Leaves densely pubescent beneath. 

Stipules linear-subulate; leaves usually serrulate. . .P. serratifolia. 

Stipules semiovate; leaves entire P. Oerstedii. 

Leaves glabrous or practically so. 

Leaf blades obtuse or rounded at the base P. ambigua. 

Leaf blades conspicuously cordate at the base. 

Glands of the petiole liguliform or filiform, at least 3 mm. long. 

P. ligularis. 
Glands of the petiole saucer-shaped P. Nelsonii. 

Bracts of the peduncles often absent, when present not involucrate, usually seta- 
ceous and scattered along the peduncle, not forming an involucre; flowers 
rarely more than 4 cm. broad when expanded, often much smaller, not highly 
colored. 

Petioles bearing conspicuous glands. 
Leaves peltate. 

Leaf blades transversely oblong-elliptic, much broader than long, green 
beneath; peduncles not more than 1.5 cm. long P. coriacea. 

Leaf blades subrotund, about as long as broad, glaucous beneath; peduncles 
more than 1.5 cm. long P. clypeophylla. 



118 FIELDIANA: BOTANY, VOLUME 24 

Leaves not peltate. 

Leaves not lobate, entire P. obovata. 

Leaves shallowly or deeply lobate, often also dentate. 
Stipules linear or filiform. 

Terminal lobe of the leaf acute or acuminate P. suberosa. 

Terminal lobe of the leaf rounded or very obtuse P. holosericea. 

Stipules semiovate or ovate. 

Petals none; leaves glabrous, the lobes long-acuminate. . .P. trinifolia. 
Petals present; leaves usually pubescent, the lobes mostly obtuse or 

somewhat acute. 
Bracts more than 3 mm. broad, laciniate or irregularly lobate. 

Bracts laciniate; fruit globose P. adenopoda. 

Bracts irregularly few-lobate; fruit narrowly obpyriform. 

P. dolichocarpa. 
Bracts narrow, entire. 

Glands borne near the middle of the petiole P. sicyoides. 

Glands borne near the apex of the petiole or at least in its upper 
third P. morifolia. 

Petioles without glands. 
Leaves glabrous or very minutely pubescent; bracts usually present. 

Bracts none; peduncles very slender, widely spreading, more than 4 cm. 

long P- filipes. 

Bracts present though often deciduous in age; peduncles stouter, usually 
less than 4 cm. long. 

Peduncles 3-flowered; leaves 2-lobate to the middle P. allantophylla. 

Peduncles 1-flowered. 

Leaves truncate or usually somewhat angulately and very shallowly 

3-lobate ac the apex P. Helleri. 

Leaves deeply or shallowly 2-lobate at the apex. 

Sepals 7-8 mm. long; corona filaments in a single series. 

P. ornithoura. 

Sepals 9-20 mm. long; corona filaments in 2 series. 

Leaves 2-lobate to below the middle, the lobes erect or nearly so. 

P. macrostemma. 

Leaves very shallowly 2-lobate or, if deeply 2-lobate, the lobes 

widely spreading P- biflora. 

Leaves usually densely and conspicuously pubescent with short or rather long 

hairs; bracts none. 
Leaves not at all lobate, acute or acuminate. 

Corona filaments linear-clavate; leaves attenuate-acuminate, finely pilos- 

ulous beneath P- cobanensis. 

Corona filaments subulate; leaves acute or acuminate, densely villpus- 

pubescent beneath P. brevipes. 

Leaves more or less lobate, usually conspicuously so. 

Peduncles bearing more than 1 flower P. sexflora. 

Peduncles 1-flowered. 

Stems and leaves hirsute with brownish hairs; leaf blades rounded at 
the base. . P. costaricensis. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 119 

Stems and leaves with short grayish hairs, or tomentulose. 
Leaf blades much longer than broad, deeply cordate at the base, the 

short lobes suberect P. Rovirosae. 

Leaf blades mostly broader than long, very shallowly or usually not 

at all cordate at the base, the lobes divergent. 
Peduncles solitary; outer corona filaments 12-15 mm. long. 

P. capsularis. 

Peduncles geminate; outer and inner corona filaments about 2 mm. 
long P. Pavonis. 

Passiflora adenopoda DC. Prodr. 3: 330. 1828. 

Dense, wet, mixed forest, ranging from about 2,700 meters almost 
down to sea level; Pete*n; Alta Verapaz; Santa Rosa; Quezaltenango; 
San Marcos. Southern Mexico; British Honduras; Costa Rica and 
Panama; Colombia to Peru. 

A large herbaceous vine, the stems hispidulous or glabrate; stipules semi- 
orbicular, 1 cm. long and 1.5 cm. broad, entire or dentate; petioles 3-5 cm. long, 
bearing 2 opposite orbicular glands 2-4 mm. in diameter about 8 mm. from the 
base of the blade; leaf blades 7-12 cm. long, 7-15 cm. broad, 3-5-lobate, the lobes 
ovate, abruptly acuminate, cordate at the base, entire or remotely denticulate, 
hispidulous on both surfaces or glabrate; peduncles solitary or geminate, 2-2.5 cm. 
long; bracts 3, inserted at the middle of the peduncle, lanceolate or oblong, 7-10 
mm. long, 4-6 mm. broad, lacerate; flowers 2-7 cm. broad, the sepals oblong- 
lanceolate, 2-4 cm. long, 1 cm. broad, obtuse, terminating in a horn-like appendage 

1 cm. long, greenish white or yellowish; petals linear-lanceolate, 10-12 mm. long; 
corona filaments uniseriate, filiform, 15-18 mm. long, white with purple bands; 
ovary densely brown- tomentose; fruit globose, 2-2.5 cm. in diameter; seeds ob- 
cordate, 6 mm. long, 4 mm. broad, straw-colored, reticulate. 

Passiflora allantophylla Masters in Donn.-Sm. Bot. Gaz. 16: 
7. 1891. 

Known only from the type, Santa Rosa, Baja Verapaz, 1,500 
meters, Tuerckheim 1425. 

Plants glabrous throughout; stipules setaceous, 1.5-2 mm. long, deciduous; 
petioles 5-10 mm. long, eglandular; leaf blades 2-2.5 cm. long, 4-5 cm. broad, 
2-lobate to the middle, the lobes rounded, mucronate, somewhat divergent, the 
blades entire, rounded at the base, bearing 1-2 pairs of ocellae on the lower surface; 
peduncles slender, equaling or slightly exceeding the petioles, 3-flowered, the 
pedicels very slender, about equaling the peduncles; bracts setaceous, deciduous; 
flowers 10-12 mm. broad, white; sepals oblong, 5-7 mm. long, 2-2.5 mm. wide, 
obtuse; petals hyaline, slightly shorter than the sepals; corona filaments uniseriate, 

2 mm. long, capitellate; ovary subglobose, glabrous. 

Passiflora ambigua Hemsl. Bot. Mag. 128: t. 7822. 1902. 

Wet, mixed forest or thickets, 500 meters or less; Alta Verapaz; 
Izabal. Southern Mexico; British Honduras to Panama. 



120 FIELDIANA: BOTANY, VOLUME 24 

A large, often somewhat woody vine, glabrous throughout, the stems terete: 
stipules filiform, 5-8 mm. long, deciduous; petioles 2-3 cm. long, bearing at or 
below the middle 2 sessile flattened glands; leaf blades oblong-lanceolate or ovate- 
lanceolate, 10-20 cm. long, 5-9 cm. broad, short-acuminate, rounded or cuneate 
at the base, penninerved, coriaceous, lustrous, entire; peduncles 4-7 cm. long; 
bracts ovate, 3-6 cm. long, 3-4 cm. broad, entire, eglandular; flowers 8-12 cm. 
broad, the calyx tube cylindric-campanulate, 1 cm. long; sepals narrowly linear- 
oblong, 4-5 cm. long, 1.5-1.8 cm. broad, obtuse, short-corniculate dorsally just 
below the apex, fleshy, white outside, pale pink and purple puncticulate within; 
petals linear-lanceolate, 3-4 cm. long, 9-10 mm. broad, white, dotted with rose- 
purple; corona filaments about 5-seriate, the outermost 1-1.5 cm. long, red, banded 
with white; gynophore slender, 1.5-2 cm. long; ovary narrowly ovoid, densely 
brown-tomentose; fruit ovoid, 10-12 cm. long, 4-4.5 cm. broad, greenish yellow; 
seeds ovate-oblong, 6-7 mm. long, 5-6 mm. broad, reticulate. 

Called "granadilla" and "granadilla de monte" in Honduras. The 
name "granadilla" is given commonly in Guatemala and Central 
America to most species of Passiflora. La Granadilla is a caserio of 
the Department of Quezaltenango, Guatemala. 

Passiflora biflora Lam. Encycl. 3: 36. 1789. P. lunata var. 
costata Masters in Mart. Fl. Bras. 13, pt. 1: 552. 1872. P. Brighami 
Wats. Proc. Amer. Acad. 21: 473. 1887 (type from Rio Chocon, Iza- 
bal, S. Watson 97). P. transversa Masters in Donn.-Sm. Bot. Gaz. 
16: 7. 1891 (type from Masagua, Escuintla, J. D. Smith 2099). 
Granadilla; murcielago; ala de murctilago; xiczotz (Pete"n, Maya). 

Dense, mixed, wet to rather dry forest or thickets, 2,500 meters 
or less, usually at 1,000 meters or lower; Pete*n; Alta Verapaz; Izabal; 
Escuintla; Retalhuleu; Quezaltenango; San Marcos; probably in all 
the lowland departments. British Honduras to El Salvador and 
Panama. Bahamas; Colombia and Venezuela. 

A small or large vine, herbaceous or perhaps at times suff rutescent, the stems 
glabrate; stipules linear-subulate, 1.5-3 mm. long; petioles 5-10 mm. long or some- 
times longer, eglandular; leaf blades very variable in outline, transversely linear 
or oblong to rounded-obdeltoid, 10 cm. long and broad or smaller, sometimes 
2-lobate, the lobes lanceolate to obovate, acute to rounded at the apex, the blades 
rounded or subcordate at the base, 3-nerved, reticulate- veined, glabrous above, 
glabrous or minutely puberulent beneath, ocellate with about 4 pairs of ocellae, 
coriaceous or subcoriaceous or often membranaceous; peduncles geminate, com- 
monly 10-12 mm. long, articulate slightly above the middle; bracts setaceous, 
2 mm. long; flowers 2.5-3.5 cm. broad; sepals ovate-lanceolate, 9-12 mm. long, 
obtuse, green outside, puberulent or glabrate, white and glabrous within; petals 
8 mm. long, 5 mm. broad, white; corona filaments biseriate, the outer ones yellow, 
7 mm. long, the inner filiform, 5 mm. long; ovary subglobose or ovoid, terete 
or angulate, glabrate to densely tomentose; fruit globose, 1-2 cm. in diameter, 
glabrous to densely puberulent; seeds obovoid, 2.5-3 mm. long, with 6-7 trans- 
verse ridges. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 121 

Called "media-luna" in Honduras and sometimes "calzoncillo" in 
El Salvador. 

Passiflora brevipes Killip, Carnegie Inst. Wash. Publ. 461: 312, 
t. 2. 1936. 

Moist or wet forest or thickets, 150-700 meters; Alta Verapaz 
(Cerro Chinaja, Steyermark 45674) . Type from Jacinto Hills, British 
Honduras, W. A. Schipp 1304. 

A large vine, densely and finely velutinous-pilosulous throughout, almost 
tomentulose on the leaves, the stems triangular; stipules linear-lanceolate, 5-7 mm. 
long, coriaceous, persistent; petioles 8-10 mm. long, eglandular; leaf blades ovate- 
lanceolate or oblong-ovate, 5-8 cm. long, 2.5-4 cm. broad, acute or acuminate, 
shallowly cordate at the base, entire, 3-nerved, conspicuously reticulate-veined, 
subcoriaceous; peduncles geminate on the main stem or on short, axillary, almost 
leafless branches 4-5 cm. long, the pedicels 4-5 mm. long, ebracteate; sepals 
linear, 8-9 mm. long, the flowers 1-1.5 cm. broad, pale green; petals narrowly 
linear, 3-4 mm. broad; corona filaments uniseriate, subulate, 2.5-3 mm. long; 
ovary narrowly ovoid; fruit ovoid, 4 cm. long, 2 cm. broad, attenuate at the base, 
short-stipitate, 6-angulate; seeds ovate, 4-5 mm. long, 2 mm. broad, with 6-7 
smooth transverse ridges, black, lustrous. 

Passiflora caerulea L. Sp. PL 959. 1753. 

Native from Brazil to Argentina; often grown in other regions for 
ornament or for its fruit; rarely cultivated in Guatemala. 

Plants glabrous and often glaucous; stipules semiovate and subreniform, 
1-2 cm. long, remotely dentate or subentire; petioles 1.5-4 cm. long, bearing 
2-4 or rarely 6 stipitate glands; leaf blades generally palmately 5-lobate about 
two-thirds their length, the lobes linear-oblong to ovate-oblong, obtuse or emar- 
ginate, entire, 2-4-glandular in the sinuses, the blades cordate at the base, mem- 
branaceous; peduncles 3-7 cm. long, eglandular; bracts broadly ovate to ovate- 
oblong, 1.5-2.5 cm. long, 1-1.5 cm. broad, rounded at the apex, pale green; flowers 
as much as 10 cm. broad, the calyx tube cupular; sepals lance-oblong or oblong, 
1.5-2 cm. long, obtuse, subcoriaceous, green outside, white or pinkish within, 
faintly carinate dorsally, the keel terminating in a slender awn as much as 5 mm. 
long; petals oblong, 1.5-2.5 cm. long, obtuse, white or pinkish; corona filaments 
4-seriate, equaling or shorter than the petals, the outer ones blue at the apex, 
white at the middle, purple at the base; ovary pruinose; fruit ovoid or subglobose, 
about 6 cm. long and 4 cm. broad, orange or yellow; seeds obcordate or cuneate, 
5 mm. long, 3.5-4 mm. broad, coarsely reticulate. 

Passiflora capsularis L. Sp. PL 957. 1753. 

Moist or wet forest or thickets, 500-2,200 meters; Chiquimula; 
Jalapa; Santa Rosa. British Honduras to El Salvador and Costa 
Rica; Greater Antilles; Colombia to Paraguay. Figure 14. 

A small or rather large, herbaceous vine, the stems 3-5-angulate, glabrate 
or pubescent; stipules linear-subulate, 5-7 mm. long; petioles 1-3 cm. long, eglan- 



122 FIELDIANA: BOTANY, VOLUME 24 

dular; leaf blades 2-7 cm. long along the costa, 4-10 cm. broad, shallowly bilobate, 
the lobes lanceolate or broader, acute or obtuse, the sinus acute or broad and 
shallow, the blades shallowly cordate, 3-nerved, pilosulous or glabrate above, 
paler beneath and densely pubescent; peduncles solitary, 1-6 cm. long, articulate 
about 1 cm. from the apex; bracts none; flowers 2-6 cm. broad, greenish white or 
pale yellow-green; sepals linear-lanceolate, 1-3 cm. long, acute, pilose dorsally; 
petals narrowly oblong-lanceolate or subspatulate, 6-15 mm. long, obtuse; corona 
filaments 1-2-seriate, the outer ones filiform, 12-15 mm. long, united at the base 
to form a thin transparent membrane, violet, the inner ones barely 3 mm. long; 
ovary obovoid or narrowly obovoid, minutely puberulent, 6-angulate; fruit ellipsoid 
or fusiform, sharply 6-angulate, 5-6 cm. long, 1.5-2 cm. broad; seeds ovate, 3.5 mm. 
long, transversely sulcate, with 5-6 smooth ridges. 

Called "calzoncillo" in El Salvador. 

Passiflora clypeophylla Masters in Donn.-Sm. Bot. Gaz. 16: 
6. 1891. 

Known only from the type, Alta Verapaz, Barranco de Rubelcruz, 
750 meters, Tuerckheim 1625. 

Plants glabrous throughout; stipules narrowly linear, 5 mm. long, coriaceous; 
petioles 2-3 cm. long, bearing at the middle 2 sessile glands 1 mm. in diameter; 
leaf blades subrotund or obscurely trilobate, 7-8 cm. long, 8-10 cm. broad, peltate, 
5-7-nerved, reticulate-veined, entire or emarginate at the apex of the nerves, 
membranaceous, deep green above, glaucous beneath; peduncles solitary or gem- 
inate, 1.5-2.5 cm. long, articulate at the middle; bracts early deciduous; flowers 
1-1.5 cm. broad; sepals oblong, 7 mm. long, obtuse, yellowish green; petals none; 
corona filaments biseriate, the outer ones filiform, 6 mm. long, purple at the 
base, yellow toward the apex, reflexed, the inner ones linear-clavate, 2 mm. long, 
erect; ovary globose. 

Passiflora cobanensis Killip, Journ. Wash. Acad. Sci. 14: 111. 
1924. 

Wet forest or thickets, on limestone, 500-950 meters; endemic; 
Alta Verapaz (type collected between Chama and Coban, Harry 
Johnson 411; collected also between Chirriact and Semococh). 

Stems slender, finely pubescent; stipules lanceolate, subfalcate, 8-9 mm. long, 
acuminate, 5-7-nerved; petioles 8-16 mm. long, eglandular, pubescent; leaf blades 
ovate-lanceolate or ovate-oblong, 7-10 cm. long, 2.5-3.5 cm. broad, attenuate- 
acuminate, rounded at the base, 3-nerved, reticulate-veined, without ocellae, 
serrulate, membranaceous, glabrous above, finely hirtellous beneath; peduncles 
1.5 cm. long; bracts none; flowers 2.2 cm. broad or larger; sepals linear-lanceolate, 
1 cm. long or longer, acute, pubescent outside; petals lanceolate, 5 mm. long, 
obtuse; corona filaments uniseriate, linear-clavate, 2.5 mm. long; ovary obovoid, 
subangulate, densely tomentulose. 

Passiflora Cookii Killip, Journ. Wash. Acad. Sci. 12: 256. 1922. 
Cul (Quezaltenango) . 




FIG. 14. Passiflora capsularis. A, Habit of plant, with flowers and buds; 
X 1 A- B, Fruit; X 1. 



123 



124 FIELDIANA: BOTANY, VOLUME 24 

Moist or wet, mixed forest or thickets, 1,000-2,700 meters, or 
perhaps even lower; Alta Verapaz (type from Sepacuite", 0. F. Cook 
& R. F. Griggs 593); Jalapa; Chimaltenango; Suchitepe'quez; Que- 
zaltenango; San Marcos. Mexico (Veracruz). 

A small or large, glabrous vine, herbaceous throughout or nearly so, the stems 
glaucous; stipules reniform, 1.5 cm. long, crenate; petioles 3-4 cm. long, eglandular; 
leaf blades ovate-orbicular, 7-8 cm. long, 6-7 cm. broad, very obscurely trilobate, 
the middle lobe broadly deltoid, obtuse, mucronate, the blades peltate above the 
rounded or truncate base, membranaceous, 5-plinerved, deep green above, glaucous 
beneath; peduncles about 8 cm. long; bracts 2, cordate, about 5 cm. long and 
4 cm. broad, usually green, entire, membranaceous; flowers 3.5-4.5 cm. broad, 
white; sepals ovate-lanceolate, 1.5 cm. long, obtuse; petals ovate-lanceolate, 
1.5 cm. long, obtuse, white, spotted with purple; corona filaments biseriate, the 
outer ones 1 cm. long, dilated at the apex, the inner scarcely 3 mm. long, capitate; 
ovary subglobose. 

This species is not very definitely separated from P. membranacea. 

Passiflora coriacea Juss. Ann. Mus. Hist. Nat. 6: 109, t. 39, f. 2. 
1805. Hoja de murcielago; ala de murcielago; granadilla de monte; 
media-luna; xicozotz (Pete"n, Maya). 

Dry to wet thickets, sometimes in pine forest or mixed forest, 
1,000 meters or usually lower; Pete"n; Alta Verapaz; Izabal; Es- 
cuintla; Suchitepe'quez; Retalhuleu; probably in all the lowland 
departments. Mexico; British Honduras to El Salvador and Pan- 
ama; South America. 

Usually a small herbaceous vine; stipules narrowly linear; petioles 2-4 cm. 
long, biglandular near the base, rarely glandular at the apex or 4-glandular, the 
glands sessile; leaf blades transversely oblong-elliptic, as much as 7 cm. long 
along the costa and 7-25 cm. broad, coriaceous to membranaceous, glabrous, 
usually truncate or subtruncate along the upper margin, peltate, the lateral lobes 
generally acute or subacute; upper inflorescence a terminal raceme devoid of 
leaves, bearing stipules and bract-like biglandular appendages (the petioles of 
abortive leaves), the lower flowers solitary or geminate in the leaf axils; flowers 
2.5-3.5 cm. broad, yellowish green; sepals oblong-lanceolate; petals none; corona 
filaments biseriate, the outer ones filiform, the inner linear; ovary ovoid, glabrous; 
fruit globose, 1-2 cm. in diameter. 

The vine is well known in most parts of Central America where 
it goes by the name "murcielago" or some variant of that word. The 
unique leaves have a slight resemblance to the wing of a bat. The spe- 
cies is perhaps the most easily recognized of all those occurring in 
Central America because of the distinctive peltate leaves, much 
longer than broad, and sharp-pointed at the lateral tips. The Passi- 
flora species are much used in Guatemala and other parts of Central 
America in domestic medicine, particularly for infections of the kid- 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 125 

neys and in general as a diuretic. This species is one of those most 
commonly used, and bunches of the dried stems and leaves often 
are on sale in the markets remote from places where the plants grow, 
as at Quezaltenango. 

Passiflora costaricensis Killip, Journ. Wash. Acad. Sci. 12: 257. 
1922. 

Wet forest, 350 meters; Alta Verapaz (Cubilgiiitz, Tuerckheim 
7877). Atlantic coast of Honduras; Costa Rica. 

Stems triangular, hirsute, at least on the angles, with long, spreading, light 
brown hairs; stipules subulate, 6-8 mm. long; petioles 1.5-2 cm. long, densely 
hirsute, eglandular; leaf blades oblong to ovate or rounded-ovate in general outline, 
9-13 cm. long, 7-11 cm. broad, bilobate, the lobes short, deltoid, acute or acuminate, 
one-third the length of the blade or shorter, suberect, the terminal sinus lunate 
or almost semicircular, the blades rounded at the base, 3-nerved, membranaceous, 
hirsute, especially beneath; peduncles solitary, 1.5 cm. long, articulate at the middle; 
bracts none; flowers 4.5-5 cm. broad; sepals linear-lanceolate, 2 cm. long, obtuse, 
hirsute dorsally, glabrous within, the margins hyaline; petals linear-oblong, 8 mm. 
long, obtuse, hyaline; corona filaments uniseriate, narrowly ligulate, equaling the 
petals; ovary minutely puberulent; fruit ellipsoid, 7-8 cm. long, 1-1.5 cm. broad, 
long-attenuate at each end; seeds slightly compressed, narrowly oblong, 3 mm. 
long, 1.5 mm. broad, black, lustrous, transversely sulcate, with 6-7 smooth 
ridges. 

Passiflora dolichocarpa Killip, Journ. Wash. Acad. Sci. 20: 374. 
1930. 

In forest, 1,500-2,500 meters; endemic; Jalapa (between Jalapa 
and Montana Miramundo) ; Chimaltenango (type from Chichavac, 
Tecpan, Jose Garcia Salas 584). 

Stems herbaceous, somewhat quadrangular, hispidulous; stipules semiovate, 
6-7 mm. long, finely hispidulous; petioles about 2 cm. long, biglandular below the 
middle or sometimes toward the apex, the glands clavate, sessile, 1.5 mm. long; 
leaf blades subhastate, 6-10 cm. long, 4.5-7 cm. broad, trilobate, the middle lobe 
lanceolate to deltoid, the basal lobes reduced, acute or short-acuminate, the blades 
sinuate-dentate, more or less cordate at the base, membranaceous, finely his- 
pidulous on both surfaces; peduncles geminate, 3.5-4 cm. long, articulate near 
the apex; bracts ovate-lanceolate, 4-5 mm. long, oblique at the base, sessile, 
irregularly lobate, hispidulous, borne on the uppermost third of the peduncle; 
flowers 2.5 cm. broad, white; sepals oblong, 10-13 mm. long, slightly cucullate 
at the apex, hispidulous outside, white, longitudinally streaked with red or purple 
within; petals oblong, 8-10 mm. long, obtuse; corona filaments uniseriate, narrowly 
linear, 6-7 mm. long, white banded with red or purple; ovary ovoid, tapering at 
the apex, stipitate, glabrous; fruit narrowly ovoid-clavate, 6 cm. long (including 
a stipe 1.5 cm. long), 1.5 cm. broad; seeds broadly obovate, 3-4 mm. long, com- 
pressed, reticulate. 



126 FIELDIANA: BOTANY, VOLUME 24 

Passiflora edulis Sims, Bot. Mag. 45: 1. 1989. 1818. Granadilla; 
granadilla silvestre; granadina. 

Native from Brazil to northern Argentina; often grown elsewhere 
for its fruit or for ornament; rarely planted in Guatemala, and per- 
haps naturalized occasionally. Figure 15. 

Plants herbaceous, glabrous throughout or rarely pilosulous; stipules linear- 
subulate, 1 cm. long, entire or minutely glandular-serrulate; petioles 4 cm. long or 
shorter, biglandular at the apex, the glands sessile or short-stipitate; leaf blades 
5-11 cm. long and 4-10 cm. broad, trilobate to below the middle, the lobes 2-4 
cm. broad, acute or acuminate, the blades rounded or shallowly cordate at the 
base, serrate, rather thick; peduncles stout, 6 cm. long or shorter; bracts ovate, 
2-2.5 cm. long, 1-1.5 cm. broad, obtuse or acute, sharply serrate, pectinate- 
dentate, or almost lacerate, often glandular on the margins; flowers as much as 
7 cm. broad; sepals oblong, 3-3.5 cm. long, 1 cm. broad, corniculate, green outside, 
white within; petals oblong, 2.5-3 cm. long, obtuse, white; corona filaments 4-5- 
seriate, those of the 2 outer series filiform or narrowly liguliform, 1.5-2.5 cm. long, 
white, purple at the base; ovary ovoid or globose, sericeous-tomentose or glabrous; 
fruit ovoid or globose, 4-5 cm. long, yellow, greenish yellow, or purplish; seeds oval, 
5-6 mm. long, 3-4 mm. broad, minutely reticulate. 

This is a rare plant in cultivation in Central America. The name 
"granadilla silvestre" supplied by Jesus Morales Ruano for a plant 
collected in Guatemala would indicate that the vine was growing 
wild, but it seems more probable that the specimens were taken 
from a garden. 

Passiflora filipes Benth. PL Hartweg. 118. 1843. 

Moist or wet, mixed forest, 1,800 meters or less; Izabal; Chiqui- 
mula; Huehuetenango; Quezaltenango. Western Texas; Mexico; 
Honduras and El Salvador; Nicaragua; Venezuela. 

Plants very slender, glabrous throughout, herbaceous, the stems terete; 
stipules linear-lanceolate, 2-4 mm. long, falcate, acuminate; petioles 1-2 cm. long, 
eglandular; leaf blades 1-4 cm. long, 2-6 cm. broad, very shallowly trilobate at 
the apex, rounded or truncate at the base, not dentate, glaucous beneath, mem- 
branaceous; peduncles very slender, almost filiform, 4-6 cm. long, widely spreading; 
bracts none; flowers 8-15 mm. broad, yellowish green or greenish white; sepals 
lanceolate or linear-lanceolate, 6-9 mm. long, subacute; petals narrowly linear, 
3-4 mm. long; corona filaments biseriate, filiform, the outer ones equaling the 
petals, the inner ones shorter; ovary globose, glabrous; fruit globose, 5-7 mm. 
in diameter; seeds slightly compressed, obovate or obcordate, 4 mm. long, 2.5-3 mm. 
broad, transversely sulcate, with 3-4 interrupted ridges. 

Called "sandillita de p&jaro" in El Salvador. 
Passiflora foetida L. Sp. PI. 959. 1753. 

A small or rather large, ill-scented, herbaceous or suffrutescent vine, glabrous 
or with highly variable indument; stipules semiannular, deeply cleft into filiform 




FIG. 15. Passiflora edulis. Habit, flower and fruits; reduced. 



127 



128 FIELDIANA: BOTANY, VOLUME 24 

gland-tipped divisions; petioles 6 cm. long or less, eglandular; leaf blades cordate 
at the base, membranaceous, 3-5-lobate, very variable in the form of the lobes; 
peduncles solitary, 6 cm. long or less; bracts forming an involucre, 2-4 times 
pinnatifid or pinnatisect, the ultimate divisions filiform, gland-tipped; flowers 
2-5 cm. broad, white to pink, lilac, or purplish; sepals ovate-oblong or ovate- 
lanceolate, corniculate dorsally just below the apex; petals oblong, oblong-lanceolate, 
or oblong-spatulate, slightly shorter than the sepals; corona filaments several- 
seriate, those of the 2 outer series filiform, about 1 cm. long, the others capillary, 
1-2 mm. long; fruit globose or subglobose. 

The species in its various forms is distributed throughout the 
American tropics and has become established in some parts of the 
Old World. It is highly variable, and some of the forms are strik- 
ingly different, but most or many of them are connected by inter- 
grading forms, and it has been considered more logical and practical 
to consider all as representing variants of a single, easily recognizable 
species. In his monograph of the genus Killip recognizes no fewer 
than 38 such varieties. Of these the following have been collected 
in Guatemala and British Honduras: 

KEY TO THE VARIETIES OF PASSIFLORA FOETIDA 

Stems glabrous or sometimes with a few scattered hairs. 
Leaves glabrous. 

Basal lobes of the leaves much longer than wide P. foetida var. ciliata. 

Basal lobes of the leaves slightly if at all longer than wide, or wanting. 
Leaves evidently hastate-lobate, the basal lobes well developed. 

P. foetida var. nicaraguensis. 
Leaves scarcely at all lobate, the basal lobes much reduced or none. 

P. foetida var. subintegra. 

Leaves pubescent on one or both surfaces P. foetida var. mayarum. 

Stems densely or rather sparsely pubescent. 

Ovary hirsute P. foetida var. gossypiifolia. 

Ovary glabrous. 

Lower surface of the leaves hirsute with short, mostly appressed hairs. 

P. foetida var. hastata. 
Lower surface of the leaves softly lanate or lanuginous, usually hirsute on the 

nerves with long spreading hairs. 
Ultimate segments of the bracts closely interlaced; styles pilose. 

P. foetida var. hirsutissima. 
Ultimate segments of the bracts not closely interlaced; styles glabrous. 

P. foetida var. lanuginosa. 

Passiflora foetida var. ciliata (Dryand.) Masters, Trans. Linn. 
Soc. 27: 631. 1871. P. ciliata Dryand. in Ait. Hort. Kew. 3: 310. 
1789. 

Pete"n (San Clemen te, H. H. Bartlett 12832). Mexico (Yucatan 
peninsula) ; Jamaica. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 129 

Plants glabrous throughout except for a few gland-tipped hairs on the petioles 
and leaf margins; leaves subhastate, subtruncate or cordate at the base, finely 
denticulate, the lobes narrowly oblanceolate or oblong-lanceolate, acute or acumi- 
nate, usually much longer than broad; bracts 2-3-pinnatisect, 3-4 cm. long, the 
segments not closely interlaced; flowers 4-5 cm. broad, light blue or pale pink; 
ripe fruit 2.5-3.5 cm. in diameter, bright red. 

The Maya name of Yucatan is recorded as "pochcac." The plant 
is reported there to have narcotic and sedative properties, producing 
deep and restful sleep. 

Passiflora foetida var. gossypiifolia (Desv.) Masters, Trans. 
Linn. Soc. 27: 631. 1871. P. gossypiifolia Desv. in Hamilt. Prodr. 
Fl. West Ind. 48. 1825. Granadilla; granadilla de culebra; sandia de 
raton; granadillita amarga. 

Dry to wet thickets, sometimes on banks or in open fields, 2,000 
meters or less; Pete"n; Alta Verapaz; Izabal; Zacapa; Jalapa; Jutiapa; 
Santa Rosa; Escuintla; Guatemala; Sacatepe"quez; Chimaltenango; 
Quiche"; Retalhuleu; Huehuetenango. Western Texas; Mexico; 
British Honduras to El Salvador and Costa Rica; West Indies; 
South America. 

Plants villous or hirsutulous throughout, often viscid, the indument grayish 
or yellowish, the hairs mostly 1.5 mm. long or shorter; leaf blades averaging about 
5 cm. long and broad, undulate or crenate-serrulate, the basal lobes usually semi- 
circular in outline, sometimes abruptly narrowed to an obtuse apex; bracts 2-3 cm. 
long, 2-3-pinnatisect, the ultimate segments not closely interlaced; ovary sparsely 
or densely hirsute, the hairs persistent in age; fruit 2-2.5 cm. in diameter, yellow 
or greenish yellow, spotted with red. 

"Pooch, pochil" (Yucatan, Maya); "pepe" (Oaxaca). This plant, 
like all forms of P. foetida, has a strong and disagreeable odor when 
crushed, the odor doubtless arising from the glands borne so pro- 
fusely on most parts of the vine. In most varieties the plants are 
extremely viscid and unpleasant to handle. Normally they climb 
over bushes or small trees but when growing in the open they often 
spread widely over the ground. 

Passiflora foetida var. hastata (Bertol.) Masters, Trans. Linn. 
Soc. 27: 631. 1871. P. hastata Bertol. Fl. Guat. 427. 1840. Dysos- 
mia hastata M. Roem. Fam. Nat. Syn. 2: 149. 1846. Granadilla. 

Moist or rather dry thickets, 300 meters or less; Alta Verapaz; 
Izabal; Escuintla (type from Escuintla, Vel&squez); Retalhuleu. 
Southern Mexico; British Honduras. 

Plants not or scarcely viscid, densely hirsute throughout with long yellowish 
hairs; leaves hastate, the pubescence chiefly of appressed or subappressed hairs, 



130 FIELDIANA: BOTANY, VOLUME 24 

the middle lobes lanceolate or oblong-lanceolate, acute or acuminate, the lateral 
lobes rounded or acute; bracts 3-3.5 cm. long after anthesis; ovary glabrous; 
fruit 2.5-3 cm. in diameter; flowers greenish white or pale purplish green. 

Passiflora foetida var. hirsutissima Killip, Carnegie Inst. 
Wash. Publ. 461: 326. 1936. 

Moist or wet thickets, 300-1,500 meters; endemic; Alta Verapaz 
(type from Sepacuit, Mary W. Owen 9; collected also near Rubel- 
tein and Cubilguitz). 

Plants densely hirsute or lanuginous-hirsute throughout, most of the hairs 
more than 2 mm. long, stiff, yellowish; leaves subhastate or hastate, the middle 
lobes ovate or oblong-ovate, acute or acuminate; bracts 2 cm. long at anthesis, 
3-pinnatisect, the segments closely and densely interlaced, densely long-hirsute; 
flowers 2.5 cm. broad, pale pink, the sepals and petals densely spotted with deep 
pink within; ovary glabrous. 

Passiflora foetida var. lanuginosa Killip, Carnegie Inst. Wash. 
Publ. 461:325. 1936. 

Moist or wet thickets or sometimes in open fields or on sand 
dunes, 200 meters or less; Pete"n; San Marcos. Southern Mexico; 
British Honduras. 

A small or rather large, herbaceous vine, the stems hirsute with spreading 
yellowish hairs 2 mm. long or more; leaves hastate or subhastate, hirsutulous or 
hirsute above, lanuginous beneath, hirsute on the nerves and veins, the middle 
lobes oblong or oblong-ovate, acute or acuminate; bracts 1.5-5 cm. long, the 
ultimate segments usually elongate, not closely interlaced; ovary glabrous; fruit 
2-3 cm. in diameter, yellowish or reddish. 

Passiflora foetida var. mayarum Killip, Carnegie Inst. Wash. 
Publ. 461:327. 1936. 

Wet to dry thickets, sometimes on gravel bars along streams, 
400 meters or less; Retalhuleu; Huehuetenango. British Honduras 
(type collected along Belize-Sibun road, Belize District, P. H. 
Gentle 6). 

Plants slender, the stems glabrous or with a few scattered long spreading hairs; 
leaves hastate or subhastate, hirtellous above with subappressed hairs, densely 
and softly pilosulous beneath, the middle lobe oblong to ovate, acute or acuminate, 
the lateral lobes broadly ovate or suborbicular, rounded or rarely subacute at the 
apex; bracts as much as 4 cm. long, the segments not closely interlaced; sepals and 
petals cream-colored, or the flowers purple and white; ovary glabrous; fruit 2-2.5 
cm. in diameter, red. 

Passiflora foetida var. nicaraguensis Killip, Carnegie Inst. 
Wash. Publ. 461: 328. 1936. P. hastata var. nicaraguensis Killip in 
Standl. Field Mus. Bot. 10: 293. 1933. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 131 

Moist or wet thickets, 250 meters or less; Pete"n; Izabal; Santa 
Rosa; Escuintla; Suchitepe"quez; Retalhuleu. Southern Mexico; 
British Honduras; Honduras; Nicaragua. 

Plants glabrous throughout or nearly so; leaves hastate, ciliate, the lateral 
lobes usually elongate, rounded or subacute at the apex, the middle lobe usually 
ovate-elliptic, acute or obtuse; bracts 3-4 cm. long in fruit, the segments not 
closely interlaced; flowers purplish white or greenish purple; fruit 2.5-3 cm. in 
diameter, glabrous. 

This variety and var. ciliata are much alike and scarcely worthy 
of separation. 

Passiflora foetida var. subintegra Killip, Carnegie Inst. Wash. 
Publ. 461:328. 1936. 

Known only from the type, All Pines, British Honduras, W. A. 
Schipp 648. 

Plants glabrous throughout, the stems very slender; leaves lanceolate or usually 
somewhat hastate, 4-6 cm. long, 1-2 cm. broad, acuminate, undulate, the basal 
lobes very short and rounded; bracts 2 cm. long at anthesis, the ultimate segments 
not closely interlaced; flowers dark rose; fruit scarlet. 

Among the Guatemalan names reported for Passiflora foetida, the 
variety uncertain, are "granadilla de culebra, pasidn, tujo" (Pete"n). 
From other regions: "melon de raton" (British Honduras) ; "granadilla 
color ada, sandia de culebra" (El Salvador); "tuuboc" (Yucatan, Maya); 
"jujito peludo" (Tabasco). The fruit in all forms of the species is 
edible but not very good. 

Passiflora Hahnii (Fourn.) Masters in Mart. Fl. Bras. 13, pt. 1 : 
569. 1872. Disemma Hahnii Fourn. Rev. Hort. 41: t. 4$0. 1869. 
P. guatemalensis Wats. Proc. Amer. Acad. 22: 473. 1887 (type from 
Rio Chocon, Izabal, S. Watson 81) . 

Wet forest or thickets, often or usually on limestone, 800 meters 
or less; Petn; Izabal. Southern Mexico; British Honduras to Costa 
Rica; Colombia. 

A slender glabrous vine, herbaceous, the stems wiry, terete, angulate above, 
striate; stipules reniform, 1 cm. long, setiferous-dentate; petioles 1.5-3 cm. long, 
eglandular; leaf blades ovate-orbicular, 5-8 cm. long, 3.5-7 cm. broad, entire or 
usually shallowly and inconspicuously trilobate, obtuse or acute and mucronate 
at the apex, peltate a short distance above the base, 3-5-nerved, membranaceous; 
peduncles solitary, 7 cm. long or shorter; bracts 2, rounded-cordate, 2.5-3 cm. long, 
1.5-2 cm. broad, denticulate toward the base, the teeth terminated by short setae; 
flowers 4-6 cm. broad; sepals and petals similar, oblong, 2-3 cm. long, 7-9 mm. 
broad, obtuse, spreading, white or cream-colored; corona filaments biseriate, yellow, 
the outer as much as 15 mm. long, spreading, dilated at the apex, the inner ones 
5 mm. long or less, clavate, erect; ovary glaucous, subglobose or ovoid, glabrous; 



132 FIELDIANA: BOTANY, VOLUME 24 

fruit globose, 3-3.5 cm. in diameter; seeds ovate-oblong, slightly compressed, 
5 mm. long, 4 mm. broad, reticulate. 

Passiflora Helleri Peyr. Linnaea 30: 54. 1859. 

Moist or wet, usually dense, mixed forest, 1,400 meters or less; 
Alta Verapaz; Baja Verapaz; Izabal; Suchitepe"quez; Quezaltenango. 
Southern Mexico. 

A small or rather large, herbaceous vine, the stems almost glabrous or finely 
pubescent; stipules linear-subulate, 3 mm. long; petioles 2-3 cm. long, eglandular; 
leaf blades ovate-oblong to orbicular in outline, 3.5-8 cm. long, 3-7 cm. broad, 
trilobate at the apex, the lobes usually very short, acute to almost truncate, mu- 
cronulate, the middle lobes usually longest, the blades rounded or subcuneate at 
the base, 3-nerved, reticulate-veined, ocellate beneath, thick-membranaceous, 
almost glabrous or minutely puberulent beneath on the nerves; peduncles 2-3.5 cm. 
long, articulate just below the base of the flower; bracts setaceous, 1.5-2.5 mm. 
long, deciduous; flowers 3-4 cm. broad; sepals oblong-lanceolate, 13-15 mm. long, 
obtuse, green outside, greenish white within; petals narrowly oblong, 1 cm. long, 
4 mm. broad, white tinged with pink; corona filaments uniseriate, 5-7 mm. long, 
green and purple-dotted along the margin outside, purple within along the margin; 
ovary ovoid, minutely pilose; fruit globose, glabrate, 1.5 cm. in diameter. 

This has been reported from Guatemala as P. trisetosa DC., a 
Mexican species. 

Passiflora holosericea L. Sp. PL 958. 1753. 

Moist or dry thickets, 700 meters or less, mostly on the plains; 
Zacapa; Chiquimula; Santa Rosa; Escuintla; Retalhuleu. Mexico; 
Honduras; El Salvador; Cuba; northern South America. 

A small or often large vine, herbaceous or somewhat woody, very densely 
and softly velutinous-pilose or tomentose throughout, the stems terete, the lowest 
ones often covered with thick corky bark; stipules filiform, 6 mm. long; petioles 
1-2.5 cm. long, bearing near the middle 2 dark brown, sessile glands 2 mm. in 
diameter; leaf blades 5-15 cm. long, 4-14 cm. broad, rather thick, shallowly 
trilobate, entire, cordate at the base, 3-nerved, reticulate-veined, densely velutinous- 
pubescent above, tomentose beneath, the lobes rounded, mucronate, the middle 
ones much longer than the others; peduncles solitary or geminate in the leaf 
axils, 2-4-flowered or those of the lower axils rarely 1-flowered, the flowers pedi- 
cellate, the pedicels bearing 2-3 subulate bractlets 2 mm. long; flowers 3-4 cm. 
broad; sepals ovate-lanceolate, 13-15 mm. long, obtuse, densely pubescent outside, 
glabrous and white within, sparsely spotted with red; petals oblanceolate or spat- 
ulate, 10-13 mm. long, 6 mm. broad, white, streaked with brown and mottled 
with red; corona filaments biseriate, the outer ones lanceolate, 7 mm. long, yellow 
at the apex, purple toward the base, the inner ones capillary, 4-5 mm. long, clavate; 
ovary obovoid, densely pilose; fruit globose, 1.5 cm. in diameter, glabrous or softly 
pubescent; seeds obovate-obcordate, 3 mm. long, 2 mm. broad, coarsely reticulate. 

A common plant in the coastal thickets of the Pacific but scarce 
elsewhere in Guatemala. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 133 

Passiflora ligularis Juss. Ann. Mus. Hist. Nat. 6: 113, t. 40. 
1805. Granadilla; cranix (Quiche"). 

Moist or wet, usually dense, mixed forest or in thickets, 900- 
2,500 meters; Alta Verapaz; Baja Verapaz; Jalapa; Sacatepe"quez; 
Suchitepe"quez; Huehuetenango; Quezaltenango; San Marcos; much 
cultivated in the mountain regions, at middle or rather high eleva- 
tions. Central and southern Mexico; El Salvador; Honduras; Costa 
Rica; widely distributed in South America. 

A large, coarse vine, glabrous throughout, the stems stout, terete; stipules 
ovate-lanceolate or oblong-lanceolate, 1-2.5 cm. long, acute or acuminate, narrowed 
at the base, entire or serrulate; petioles 4-10 cm. long, bearing 4-6 scattered, 
liguliform or filiform glands 3-10 mm. long; leaf blades broadly ovate, 8-19 cm. 
long, 6-14 cm. broad, abruptly acuminate, deeply cordate at the base, entire, 
membranaceous; peduncles solitary or geminate, 2-4 cm. long; bracts 2-3.5 cm. 
long, 1-1.5 cm. broad, connate one-fifth to one-third their length, the free portions 
ovate or ovate-lanceolate, acute, entire; flowers 6-9 cm. broad; calyx tube short- 
campanulate; sepals ovate-oblong, 2.5-3.5 cm. long, acute, green outside, white 
within; petals oblong, about 3 cm. long, 8-10 mm. broad, white or pinkish white; 
corona filaments 5-7-seriate, the filaments of the two outer rows as long as the 
petals, radiate, terete, blue at the apex, banded with white and reddish purple be- 
low, the inner filaments barely 2 mm. long; fruit ovoid or ellipsoid, 6-8 cm. long and 
4-5 cm. broad or slightly larger, the pericarp yellowish, purplish, or dark reddish, 
thick and tough, the pulp about the seeds white; seeds narrowly obcordate, 6 mm. 
long, 4 mm. broad, minutely tridentate at the apex, the sides irregularly reticulate. 

This species is the granadilla of Central America, being the only 
species grown commonly there for its edible fruit. The term "grana- 
dilla" is derived, of course, from granada, the Spanish term for the 
pomegranate, that being doubtless the only Mediterranean fruit with 
which the early explorers could associate this strange new product 
of the Indies. The fruit of P. ligularis generally is considered supe- 
rior in flavor to that of such species as P. maliformis and P. laurifolia, 
which are much grown in some other parts of tropical America. The 
vines are grown very commonly in the mountains of Guatemala as 
well as in other countries of Central America, and the fruits are pro- 
duced in vast quantities and carried long distances for sale in regions 
where the plants are not cultivated. Granadillas are eaten raw. 
They are much liked by most persons because of their agreeable 
flavor, and they are easy to eat; the tough rind is opened and the 
pulp and seeds removed with a spoon. The seeds are crisp and 
crunchy, and often are chewed and eaten with the abundant pulp. 
Just where P. ligularis is native in Central America is rather uncer- 
tain, for many of the specimens are taken from cultivated plants and 
collectors seldom note on their labels whether the plants were found 
wild or in gardens. There can be no doubt, however, that the species 



134 FIELDIANA: BOTANY, VOLUME 24 

is native in the mountains of Guatemala or else it has become 
widely naturalized in places where it appears native. The juice of 
the fruit is often used for flavoring beverages and ices. 

Passiflora macros tern ma Killip, Field Mus. Bot. 23: 217. 1947. 

Moist slopes and thickets (type from Guatemala, Steyermark 
31519). Endemic; Chiquimula; Zacapa. 

Herbaceous vine, glabrous throughout; stem subangular; stipules falcate- 
subulate, 4 mm. long, subpersistent; petioles 1-1.5 cm. long, glandless; leaf blades 
membranous, sublustrous on both surfaces, rounded or subtruncate at the base, 
ocellate beneath, bilobed one-half to two-thirds their length, 1.5-4 cm. long along 
the midnerve, 4-9 cm. along the lateral nerves, 3-7 cm. between the apices of 
the lobes, the lobes erect or very slightly divaricate, 1.2-3 cm. broad, rounded, 
the sinus rounded or subtruncate; peduncles in pairs, 2-3 cm. long, divaricate; 
bracts setaceous, 2.5-3 mm. long, borne just below the base of the flower; flowers 
3-4 cm. broad when expanded, the calyx tube broadly campanulate; sepals lance- 
olate, about 2 cm. long, 1 cm. broad at the base, obtuse, ecorniculate, greenish 
purple without, deep lavender within; petals oblong, about 1.5 cm. long, 4-5 mm. 
broad, obtuse, membranous, deep lavender; corona 1-ranked, the filaments numer- 
ous, filiform, 6-7 mm. long, erect; operculum membranous, erect, 4-5 mm. high, 
very slightly plicate, purple, green at the base; nectar ring none; limen about 
3 mm. high, the base closely surrounding the gynophore, the margin spreading 
outward, entire; gynophore about 1 cm. long, rather stout; ovary subglobose. 

Passiflora membranacea Benth. PI. Hartweg. 83. 1841 (based 
on material from Cuesta de Argueta, Solola and Volcan de Agua, 
Sacatepe"quez, Hartweg, without number). Cieca membranacea M. 
Roem. Fam. Nat. Syn. 2: 140. 1846. Granadilla; granadilla silvestre. 

Dense, moist or wet, mixed forest, occasionally in oak forest, 
sometimes in exposed places in thickets, often on cliffs or in white 
sand, 1,350-3,000 meters, chiefly at the higher elevations; El Pro- 
greso; Sacatepe"quez; Chimaltenango; Sol old; Suchitepe'quez; Que- 
zaltenango; San Marcos. Southern Mexico; Honduras; El Salvador; 
Costa Rica. 

A small or often very large, slender, herbaceous vine, glabrous throughout; 
stipules cordate-reniform, 1-1.5 cm. long, 1.5-3 cm. broad, obscurely crenulate; 
petioles 2-4 cm. long, eglandular; leaf blades orbicular or nearly so in outline, 
5-10 cm. long and broad, obscurely and very shallowly trilobate at the apex, 
peltate a short distance above the base, 3-nerved, membranaceous, usually tinged 
with red or purple when young, glaucescent beneath; peduncles solitary, 9-15 cm. 
long; bracts 2-3, cordate-ovate or cordate-reniform, 3-5 cm. long, 2-4 cm. broad, 
obtuse or rounded at the apex, red-purple, rose-colored, or reddish green, thin; 
flowers white or cream-colored; calyx tube broadly campanulate, 2 cm. broad at 
the base; sepals oblong-lanceolate, 3.5-4 cm. long, 1 cm. broad, obtuse, erect; 
petals oblanceolate, 3.5-4 cm. long; corona filaments biseriate, the outer ones 8-10 
mm. long, filiform, deep red, the inner ones capillary, barely 2 mm. long, capitellate; 






STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 135 

gynophore about 4 cm. long; ovary narrowly ellipsoid, trisulcate; fruit ovoid or 
ellipsoid, 4-8 cm. long, green or yellow, the flesh light green, bitter; seeds black, 
oblong-ovate, 4.5 mm. long, 3 mm. broad, deeply punctate. 

This vine is abundant in many places in the mountains of the 
Occidente, often almost completely covering shrubs or good-sized 
trees, or forming festoons over cliffs. It is the handsomest and showi- 
est of Guatemalan Passifloras, not because of the flowers but on 
account of the large and delicately colored bracts. The vines often 
creep closely along the trunks of cypress (Cupressus) trees, giving a 
strange effect of having been planted there. 

Passiflora morifolia Masters in Mart. Fl. Bras. 13, pt. 1: 555. 
1872. P. Heydei Killip, Journ. Wash. Acad. Sci. 12: 258. 1922 (type 
from Casillas, Santa Rosa, Heyde & Lux 3772). Granadilla de monte; 
granadilla de Castillo,; granadilla. 

Moist or rather dry thickets, sometimes on dry open grassy slopes, 
1,300-2,100 meters; Jalapa; Santa Rosa; Guatemala; Huehuete- 
nango; San Marcos. Mexico (?); Peru to Argentina. 

A rather stout, herbaceous vine, the stems somewhat 4-angulate, hispidulous 
or'glabrate; stipules semiovate, 6 mm. long, long-acuminate, minutely hispidulous; 
petioles 6 cm. long or less, hispidulous or glandular-pubescent, biglandular, the 
glands borne on the upper part of the petiole, thick-stipitate, 1.5 mm. long; leaf 
blades membranaceous, 4-11 cm. long, 5-15 cm. broad, trilobate to below the 
middle, deeply cordate at the base, 3-nerved, repand-dentate or subentire, dark 
green above and hispidulous with minute uncinate hairs, paler and minutely 
pilosulous beneath, the lobes acute, the middle lobe ovate or ovate-lanceolate, 
usually narrowed below; peduncles solitary or geminate, 1-2 cm. long; bracts 
setaceous, 2.5-3 mm. long, inserted about 1 cm. below the base of the flower; 
flowers 2-3 cm. broad; sepals linear-oblong, 1-1.5 cm. long, obtuse, densely his- 
pidulous to glabrate outside, green, glabrous and white within, mottled with red, 
the apex terminating in a horn 3.5 mm. long; petals linear-lanceolate, 6-8 mm. 
long, obtuse, white; corona filaments uniseriate, filiform, 5-6 mm. long, white 
banded with blue or violet; ovary subglobose, densely pubescent, glaucous; fruit 
globose, 2 cm. in diameter, bluish black and glaucous, hispidulous; seeds very 
slightly compressed, obcordate-obovoid, 4 mm. long, coarsely reticulate. 

One collection that we refer here was named P. pilosa Ruiz & 
Pavon by Killip, and another was indicated as a new species, near 
P. exsudans Zucc. There is no doubt that the species of this alliance 
have been divided too finely and that the characters depended upon 
for separating them, especially the position of the petiolar glands, 
are inconstant and unreliable. 

Passiflora Nelsonii Mast. & Rose, Contr. U. S. Nat. Herb. 5: 
142, t. 17, 1897. 



136 FIELDIANA: BOTANY, VOLUME 24 

Moist or wet, mixed forest, 1,000-1,800 meters; Alta Verapaz; 
Huehuetenango. Southern Mexico, the type from Tumbala, Chiapas. 

A large, herbaceous vine, glabrous throughout or nearly so, the stems terete; 
stipules oblanceolate, 10-12 mm. long, acuminate, narrowed at the base, deciduous; 
petioles 2-3 cm. long, bearing 2 pairs of sessile saucer-shaped glands near the apex; 
leaf blades broadly ovate, 9-17 cm. long, 8-11 cm. broad, acuminate, rounded or 
cordate at the base, entire, penninerved, thick-membranaceous; peduncles solitary, 
3 cm. long or less; bracts broadly ovate, 5-6 cm. long, 3.5-4.5 cm. broad, acuminate, 
free almost to the base; flowers 5-7 cm. broad; sepals oblong-lanceolate, 3 cm. 
long, 1 cm. broad, corniculate just below the apex; petals equaling and slightly 
narrower than the sepals; corona filaments several-seriate, the outermost filaments 
subulate, 1-1.5 cm. long, the inner ones 1-2 mm. long; gynophore stout, 7-9 mm. 
long; ovary narrowly obovoid; fruit subglobose. 



Passiflora normalis L., a synonym of P. perfoliata L., a Jamaican 
species, was reported by Bertoloni as collected in Guatemala by 
Velasquez, with the local name "camacarnata." It is quite uncertain 
what the plant so listed may have been, and the vernacular name 
has not been noted as in use in Guatemala, although a similar 
name is used in El Salvador for P. pulchella. 

Passiflora obovata Killip, Carnegie Inst. Wash. Publ. 461: 308, 
t. 1. 1936. 

Known only from the type, Pete"n, Camp 35 on the British Hon- 
duras boundary, 850 meters, W. A. Schipp 713. 

A large glabrous vine as much as 18 meters long, the stem 5 cm. in diameter, 
subangulate; stipules soon deciduous; petioles 2 cm. long, biglandular just above 
the middle, the glands oblong, scar-like, 1 mm. long; leaf blades obovate or oblong- 
obovate, 9-12 cm. long, 5-6.5 cm. broad, rather abruptly acuminate, slightly 
narrowed at the base, entire, obscurely 5-plinerved, chartaceous, lustrous; peduncles 
geminate, 2.5-3 cm. long, articulate above the middle; bracts minute, triangular- 
ovate, acute, closely appressed to the peduncle and inserted near its base; flowers 
4 cm. broad, greenish white; calyx patelliform, the sepals oblong, 1.5 cm. long, 8 mm. 
broad, obtuse; petals linear-oblong, 13 mm. long, 4 mm. broad; corona filaments 
biseriate, the outer ones ligulate, about equaling the petals, the inner capillary, 
2 mm. long, minutely capitellate; ovary globose; ovules apparently reticulate. 

Passiflora Oerstedii Masters in Mart. Fl. Bras. 13, pt. 1: 562. 
1872. 

Moist or wet thickets, 1,200-1,400 meters; Suchitepe"quez; Que- 
zaltenango. Southern Mexico; Costa Rica; Colombia and Venezuela. 

A small or large, herbaceous vine, the stems slender, terete or nearly so; 
stipules semiovate, 1-4 cm. long, 5-15 mm. broad, acuminate and usually long- 
cuspidate, rounded at the base; petioles 1-4 cm. long, bearing 4-6 or rarely 2 
scattered or geminate, stipitate glands; leaf blades ovate-lanceolate to narrowly 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 137 

oblong-lanceolate, 6-13 cm. long, 3-9 cm. broad, obtuse to long-acuminate, rounded 
or usually subcordate at the base, subpeltate, 5-7-nerved, entire or nearly so, 
membranaceous, glabrous above, glaucescent beneath and usually thinly pilose 
or villosulous with somewhat matted hairs; peduncles 2-4 cm. long, glabrous; 
bracts ovate-lanceolate, 1-1.5 cm. long, 5-8 mm. broad, acute, rounded or cordate 
at the base, serrulate, reticulate-veined, inserted 5-10 mm. below the base of the 
flower, deciduous; flowers 4-6 cm. broad; calyx tube campanulate, 7-8 mm. long, 
the sepals ovate-lanceolate, 2-3 cm. long, 5-12 mm. broad, cucullate at the apex,' 
carinate, the keel terminating in a short awn, white within; petals linear, 1-1.5 cm! 
long, 3-5 mm. broad, obtuse, white or pinkish; corolla filaments several-seriate! 
purple, those of the 2 outer series filiform, 1.5-2 cm. long, erect, the innermost 
1.5-3 mm. long; ovary ovoid, glabrous, glaucous; fruit ovoid, 4-6 cm. long, 2-3 
cm. broad; seeds narrowly obovate, 4-5 mm. long, 2-3 mm. broad, shallowly 
tridentate, coarsely reticulate. 

Much more common in Guatemala than the typical form of the 
species is the following variety: 

Passiflora Oerstedii var. choconiana (Wats.) Killip, Carnegie 
Inst. Wash. Publ. 461: 321. 1936. P. choconiana Wats. Proc. Amer 
Acad. 22: 474. 1887 (type from Rio Chocon, Izabal, S. Watson 212). 

Moist or wet, mixed forest or thickets, 1,500 meters or less; Pete"n; 
Alta Verapaz; Baja Verapaz; Izabal; Santa Rosa; San Marcos! 
Southern Mexico; British Honduras; Atlantic coast of Honduras; 
Costa Rica. 

A small or large, herbaceous vine; differing from the typical form of the species 
only in having the leaves deeply trilobate. 

Called "cacapache" and "granada" in Veracruz. Schipp reports 
the flowers as blue. 

Passiflora ornithoura Masters in Donn.-Sm. Bot. Gaz. 16: 8. 
1891. P. dichthyophylla Masters, loc. cit. (type from Duenas, Saca- 
tepe*quez, J. D. Smith 2136). Granadilla de montana; calzoncillo. 

Wet to dry thickets or forest, 2,000 meters or less; Zacapa; Chi- 
quimula; Jalapa; Santa Rosa; Escuintla; Guatemala; Sacatepe'quez; 
Chimaltenango; Solola; Huehuetenango. El Salvador; Costa Rica! 

Plants glabrous, the stems compressed, striate; stipules setaceous, 3 mm 
long, deciduous; petioles 1.5-2.5 cm. long, eglandular; leaves bilobate usually 
to below the middle, often almost to the base, cuneate to rounded at the base, 
3-nerved, membranaceous, entire, deep green above and often with a broad silvery 
stripe along the costa, paler beneath or glaucescent, the lobes linear to ovate, 
3-10 cm. long, 5-20 mm. broad, acute or obtuse, mucronulate, usually ascending,' 
sometimes spreading; peduncles geminate, 1-2 cm. long; bracts setaceous, 1.5 mm' 
long; flowers 1.5 cm. broad; sepals oblong, 7-8 mm. long, obtuse; petals linear,' 
5 mm. long, hyaline, white; corona filaments uniseriate, linear-clavate, barely 



138 FIELDIANA: BOTANY, VOLUME 24 

2 mm. long; fruit globose, 6-8 mm. in diameter, purple-black; seeds strongly 
compressed, broadly ovate, 2 mm. long, 2 mm. broad, transversely sulcate with 
about 6 conspicuously rugulose ridges. 

This has been reported from Guatemala as P. mexicana Juss. 

Passiflora Palmeri Rose, Contr. U. S. Nat. Herb. 1: 131, 1. 14. 
1892. 

Plants usually low and densely branched shrubs, or more elongate and scandent, 
very viscid, the branches densely pilose; stipules semiannular, deeply cleft into 
filiform gland-tipped divisions; petioles 1-1.5 cm. long, pilose, bearing numerous 
coarse gland-tipped hairs; leaf blades 1-3 cm. long, 2-4 cm. broad, trilobate, cordate 
at the base, 3-5-nerved, serrulate, densely glandular-ciliate, densely brown- 
hirsute-tomentose, the lobes orbicular, rounded at the apex; peduncles 4-5 cm. 
long; bracts 2-4 cm. long, pilose, deeply 2-pinnatisect into capillary gland-tipped 
segments; flowers 5-7 cm. broad; sepals linear, 3.5 cm. long, 3-nerved, pilose, 
white within, slender-awned dorsally just below the apex; petals linear, 2-3 cm. 
long, white, 1-nerved; corona filaments several-seriate, those of the two outer 
series erect, capillary, 2.5 mm. long; gynophore slender, 2 cm. long; fruit globose, 
2.5-3 cm. in diameter, densely pilose; seeds oblong or narrowly cuneate, 6 mm. long, 
2 mm. broad, tridentate at the apex, reticulate. 

The typical form of the species is known only from Baja Califor- 
nia, Mexico. The species is represented in Guatemala by the follow- 
ing variety: 

Passiflora Palmeri var. sublanceolata Killip, Carnegie Inst. 
Wash. Publ. 461:322. 1936. 

Pete"n (between Uaxactun and San Clemente, H. H. Bartlett 
12788). Mexico (Yucatan peninsula). 

Leaf blades lanceolate in general outline, 3-5 cm. long, 2-3.5 cm. broad, the 
basal lobes much reduced; bracts 2-2.5 cm. long, 8-10 mm. broad; flowers said 
to be "bright scarlet" or rose-red. 

Passiflora Pavonis Masters, Journ. Bot. 21: 35. 1883. 

Moist or wet thickets or forests near streams or in sandy oak 
forest, 2,400-2,600 meters; Huehuetenango; Quezaltenango; San 
Marcos. Central and southern Mexico. 

A large, herbaceous vine, the stems angulate, glabrous or sparsely pubescent; 
stipules narrowly falcate, 3 mm. long, acuminate, deciduous; petioles 5-15 mm. 
long, densely pubescent, eglandular; leaf blades 1-5.5 cm. long, 2-7.5 cm. broad, 
broad and shallowly trilobate at the apex, rounded at the base, 3-5-nerved, entire, 
green and glabrous above, paler beneath and thinly or densely pilose, the lobes 
subequal, short and broad, rounded or truncate at the apex, mucronate; peduncles 
geminate, 1-2.5 cm. long, puberulent; bracts none; flowers 1.5-2 cm. broad, yel- 
lowish green or greenish white; calyx tube setulose, ventricose at the base; sepals 
linear-lanceolate, 10-12 mm. long, 2 mm. broad, acuminate, obscurely 3-nerved, 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 139 

minutely puberulent; petals 3-4 mm. long, acute; corona filaments biseriate, capil- 
lary, the outer ones one-third to one-half as long as the petals, reflexed; ovary 
setulose; fruit globose, 6-8 mm. in diameter, glabrate; seeds broadly ovate, com- 
pressed, 2.5 mm. long, 2 mm. broad, transversely rugose, with 6 rounded, conspic- 
uously rugulose ridges. 

Passiflora platyloba Killip, Journ. Wash. Acad. Sci. 12: 260. 
1922. Granadilla. 

Moist or dry, often rocky thickets, on plains or hillsides, 150- 
1,400 meters; Chiquimula; Jutiapa; Santa Rosa; Retalhuleu. El 
Salvador; Nicaragua; Costa Rica. 

A small, herbaceous vine, glabrous except on the bracts, the stems stout, terete, 
striate; stipules coriaceous, linear, 10-12 mm. long, 3-nerved, remotely serrulate, 
deciduous; petioles 6-7 cm. long, bearing above the base 2 sessile flattened glands 
2 mm. broad; leaf blades 10-14 cm. long, 12-18 cm. broad, trilobate to the middle, 
deeply cordate at the base, finely serrulate, biglandular in the sinuses, 3-5-nerved, 
membranaceous, the middle lobes broadly ovate, abruptly acuminate, 4-8 cm. 
broad, the lateral lobes almost equaling the middle one; peduncles solitary, 6-7 cm. 
long; bracts ovate, entire, 5-7 cm. long, 3-5 cm. broad, membranaceous, inserted 
1 cm. below the apex of the peduncle, completely enveloping the flower, united for 
about 2 cm., acute or obtuse, densely puberulent on both surfaces; flowers purple, 
4-5 cm. broad; calyx tube campanulate, 1 cm. long; sepals oblong-lanceolate, 18- 
20 mm. long, 8 mm. broad, obtuse, strongly carinate, the keel terminating in an 
awn 5-6 mm. long; petals linear-lanceolate, 15-17 mm. long, 5 mm. broad, thin, 
obtuse; corona several-seriate, the outermost filaments filiform, 7 mm. long, those 
of the second series stout, liguliform, 15 mm. long, white banded with purple; 
gynophore glabrous; ovary ellipsoid, glabrous; fruit 3-3.5 cm. long, with a hard 
pericarp. 

Called "granadilla dcida" and "granadilla monies" in El Salvador, 
where the very acid fruit is used for preparing iced beverages. 

Passiflora porphyretica Masters in Donn.-Sm. Bot. Gaz. 20: 
538, t. 36. 1895. 

Known in Guatemala only from the type (Jutiapa, 420 meters, 
Heyde & Lux 6334). Southern Mexico. 

Stems terete, densely pubescent or hirsute; stipules cordate, 8-10 mm. long, 
reddish or violet-purple, densely ciliate; petioles 1-2.5 cm. long, densely hirsute, 
eglandular; leaf blades orbicular in outline, 5-10 cm. long and broad or smaller, 
broad at the apex and shallowly 2-3-lobate, rounded or subcordate at the base, 
3-nerved, reticulate- veined, strigillose and somewhat scabrous above, densely pale- 
tomentose beneath, the lobes short and broad, rounded to subacute at the apex, 
mucronate; peduncles geminate on the main stems or usually on axillary leafy 
branches, the leaves often absent and the inflorescence thus appearing racemose; 
bracts similar to the stipules, broadly cordate, 10 mm. long, 8 mm. broad, obtuse, 
reddish or violet-purple, glabrate outside, pilosulous within; flowers 3 cm. broad; 
sepals linear-oblong, 1.5 cm. long, 3 mm. broad, very densely pilose outside, gla- 



140 FIELDIANA: BOTANY, VOLUME 24 

brate within; petals linear-oblong, 8 mm. long, obtuse, pink; corona filaments fili- 
form, spotted with purple, biseriate, the outer ones 6-8 mm. long, the inner 2 mm. 
long; ovary globose, densely white-pilose; fruit globose, 1 cm. in diameter; seeds 
transversely sulcate, with 6-8 rugulose ridges. 

Passiflora prolata Masters, Bot. Jahrb. 8: 219. 1887. Granadilla. 

Moist or wet thickets or forest, 200-2,000 meters; endemic; Pe- 
te'n; Chiquimula; Jalapa; Jutiapa; Santa Rosa; Suchitepe"quez; Solola 
(type from Sultan, F. C. Lehmann 1630); Quezaltenango; San Marcos. 

A large or small, herbaceous vine, the stems obtusely angulate, glabrous or 
puberulent; stipules linear-setaceous, 4-15 mm. long, decidous; petioles 1-2 cm. 
long, 2-4-glandular near the middle, the glands subsessile; leaf blades trilobate to 
the middle, 5.5-12 cm. long, 6-13 cm. broad, rounded or subcordate at the base, 
serrulate, membranaceous, glabrous or finely puberulent beneath, the lobes lan- 
ceolate or oblong-lanceolate, caudate-acuminate; peduncles 3-4 cm. long; bracts 
free to the base, ovate-lanceolate, 4.5-5 cm. long, 1.5-2 cm. broad, entire, mem- 
branaceous, green, glabrous or puberulent; flowers 5-7 cm. broad; calyx tube 
campanulate; sepals oblong, 2-2.5 cm. long, 8-10 mm. broad, long-aristate dor- 
sally just below the apex, white within; petals oblong-spatulate, slightly shorter 
than the sepals, obtuse, white; corona filaments several-seriate, filiform, purple, 
those of the 2 outer series 2 cm. long, the innermost half as long; ovary ellipsoid, 
glabrous. 

Passiflora pulchella HBK. Nov. Gen. & Sp. 2: 134. 1817. 
P. pulchella var. bifidata Masters, Bot. Jahrb. 8: 220. 1887 (type from 
Chiquimula, Chiquimula, F. C. Lehmann 1708). 

Moist or dry, often rocky thickets, 900 meters or less; Chiqui- 
mula; Jutiapa; Santa Rosa; Escuintla; Retalhuleu. Southern Mex- 
ico; El Salvador to Panama; Colombia and Venezuela. 

Usually a small vine, herbaceous or often woody below, glabrous throughout 
or nearly so; stipules linear, subfalcate, 5-7 mm. long; petioles 1.2 cm. long, egland- 
ular; leaf blades 2-6 cm. long along the costa, 5-10 cm. broad between the tips of 
the lobes, bilobate one-fourth to one-half their length or sometimes truncate, trun- 
cate to subcuneate at the base, 3-nerved, chartaceous, prominently reticulate- 
veined, ocellate beneath, the lobes divaricate to suberect, truncate or rounded at 
the apex; peduncles solitary, 5-8 cm. long; bracts suborbicular or ovate, 1-1.5 cm. 
long, 8-10 mm. broad, subacute or obtuse, abruptly narrowed at the base, mem- 
branaceous, inserted 2 mm. below the base of the flower, purplish red; flowers 
4.5-5.5 cm. broad, deep blue; sepals oblong, obtuse, 2 cm. long, 7-9 mm. broad; 
petals ovate-lanceolate or oblong-lanceolate, 13-15 mm. long, 4 mm. broad; corona 
filaments several-seriate, filiform, the outermost equaling the petals, those of the 
inner series 4-5 mm. long; ovary subglobose, glabrous; fruit globose, 1-1.5 cm. in 
diameter; seeds obovate, 2-3 mm. long, 1 mm. broad, transversely sulcate, with 
about 6 rugulose ridges. 

Known in El Salvador by the names "calzoncillo" and "cama- 
carlata." 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 141 

Passiflora quadrangularis L. Syst. ed. 10. 1248. 1759. P. macro- 
carpa Masters, Gard. Chron. 1869: 1012. 1869. Granadilla real. 

Planted occasionally in Guatemala, from the capital down to the 
coasts, but infrequent; native of tropical America, but the native 
habitat uncertain ; not found wild in Guatemala or elsewhere in Cen- 
tral America, unless as a rare escape from cultivation. Figure 16. 

A large, coarse, glabrous, herbaceous vine, the stems stout, 4-angulate and 
conspicuously winged along the angles; stipules ovate or ovate-lanceolate, 2-3.5 cm. 
long, acute, narrowed at the base, entire or slightly serrulate; petioles stout, 2- 
5 cm. long, 6-glandular, the glands geminate, almost sessile; leaves broadly ovate 
or oblong-ovate, 10-20 cm. long, 8-15 cm. broad, entire, abruptly acuminate, 
rounded to shallowly cordate at the base, penninerved; peduncles 1.5-3 cm. long, 
triangulate; bracts cordate-ovate, 3-5.5 cm. long, 1.5-4 cm. broad, acute or sub- 
acute, entire or serrulate toward the base, thin-membranaceous; flowers as much 
as 12 cm. broad; calyx tube campanulate; sepals ovate or oblong-ovate, 3-4 cm. 
long, cucullate at the apex, corniculate, greenish or reddish green outside, white, 
violet, or pinkish within; petals oblong-ovate or oblong-lanceolate, 3-4.5 cm. long, 
1-2 cm. broad, obtuse, white tinged with pink; corolla 5-seriate, the outer fila- 
ments as much as 6 cm. long, banded with reddish purple and white at the base, 
blue at the middle, mottled with pinkish blue in the upper half; fruit oblong-ovoid, 
20-30 cm. long and 12-15 cm. broad or even larger, terete or longitudinally some- 
what trisulcate; seeds broadly obcordate or suborbicular, 7-10 mm. long, strongly 
compressed, reticulate at the center on each side, radiately striate at the margins. 

This species is easily recognized by the sharply angled and broadly 
winged stems, and by the very large, green fruits, which are surpris- 
ingly light in weight considering their great size. The pulp is acid 
and in some regions is much used for flavoring beverages and ices. 
It is not often, if ever, eaten alone. In some places in the Central 
American lowlands this vine is cultivated commonly for its fruits but 
it is scarce in Guatemala, and we have not observed the fruits in any 
of the local markets. 



Passiflora reflexiflora Cav., a synonym of which is Tacsonia laevis 
Benth. (PL Hartweg. 118. 1843) has been reported from Guatemala 
on the basis of Hartweg 662, which was reported to have come from 
Guatemala. It is now believed that there was an error in the locality 
data of this number of Hartweg's collections, and that the material 
really came from Ecuador. 

Passiflora Rovirosae Killip, Journ. Wash. Acad. Sci. 12: 259. 
1922. Xicozotz (Maya) ; ala de murcielago. 

At 200 meters or lower; Pete"n (Uaxactun, H. H. Bartlett 12691). 
Southern Mexico; British Honduras. 




FIG. 16. Passiflora quadrangularis. Habit, flower and fruit; reduced. 



142 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 143 






" Sp ' PI - 965 ' "53- Caranilicho 

Moist or wet thickets or forest, 500 meters or less; Peten; Alta 

Honduras a 101 * the Atlantic coast 



; stipules linear-sub- 

SSSS 

or subcordate at the 

5SS 

8-10 mm. broad, acute, green, finely pubescent; flowers 4-6 cm. broad the calvx 









Called "granadilla" in Honduras and doubtless also in Guate- 
mala; "junto amarillo" (Campeche). 

Passiflora sexflora Juss. Ann. Mus. Hist. Nat. 6: 110, t. 37 f 2 
1805. Ala de mmciilago. 



* tcketS ' S metimes in oak OT 



foresoo n v ' es n oa OT P ne 

forest, 300-2,500 meters; Alta Verapaz; Baja Verapaz; El Progreso- 

' 



S, : f ,; SC 7 a; UatemaU; S -atuez; Chma 
o, Solola; Suchite^uez; Quiche; Huehuetenango; Quezalte- 



144 FIELDIANA: BOTANY, VOLUME 24 

nango; San Marcos. Southern Florida; southern Mexico; Honduras; 
Costa Rica; Panama; West Indies; Colombia. 

A large or small, herbaceous or suffrutescent vine, the stems subangulate, 
densely pilose; stipules linear-subulate, about 5 mm. long; petioles 2.5-3 cm. long, 
pilose, eglandular; leaf blades membranaceous, 3-8 cm. long along the costa, 5- 
11 cm. broad, trilobate, rounded or subcordate at the base, entire, hirsutulous and 
often setulose above, softly pilose or tomentose beneath, the middle lobe usually 
shorter than the lateral ones, these acute to rounded at the apex; peduncles gemi- 
nate, 2-10-flowered; bracts and bractlets linear-lanceolate, 3-5 mm. long, irregu- 
larly few-cleft or subentire; flowers 1.5-3 cm. broad; sepals lanceolate, 8-15 mm. 
long, 2-4 mm. broad, acute, densely hirsute outside, glabrous within, white; petals 
linear, 8-10 mm. long, obtuse; corona filaments biseriate, the outer ones as long 
as the petals, white at the apex, purple below, those of the inner surface half as 
long, purple throughout, capitate; ovary globose, densely brown-pubescent; fruit 
globose or depressed-globose, 6-10 mm. in diameter, densely pubescent; seeds 
obovate, slightly compressed, black, transversely 6-7-sulcate, the ridges rugulose. 

This is one of the commonest species of Passiflora in Guatemala. 

Passiflora sicyoides Schlecht. & Cham. Linnaea 5: 88. 1830. 

Moist or wet thickets, sometimes in oak forest, 1,750 meters or 
lower; Santa Rosa (Laguna de los Pinos); Quiche"; Huehuetenango. 
Central and southern Mexico. 

A slender, herbaceous vine, the stems slender, pilose or glabrate; stipules semi- 
ovate, cuspidate, 5-7 mm. long; petals 2-6 cm. long, pilose, bearing at the middle 
2 subopposite stipitate glands 1.5 mm. long; leaf blades thin-membranaceous, 
5-12 cm. long, 4-15 cm. broad, trilobate, entire or nearly so, cordate at the base, 
3-nerved, minutely pilose on both surfaces or almost glabrous, glaucescent beneath, 
the lobes deltoid, acuminate to obtuse; peduncles solitary or geminate, half as long 
as the petioles; bracts setaceous, 3 mm. long; flowers 3-4 cm. broad, the calyx tube 
pilose; sepals oblong-lanceolate, 12-20 mm. long, 6-10 mm. broad, acute, sparsely 
pilose; petals ovate-lanceolate, 5-12 mm. long, greenish white; corona filaments 
uniseriate, 7-8 mm. long, linear, white, with 4-5 transverse purple bands; ovary 
ovoid, stipitate, glabrous; fruit obovoid, tapering to a stipe 2 cm. long; seeds ob- 
cordate, abruptly acute at the base, 5 mm. long, 4 mm. broad, coarsely reticulate. 

Passiflora suberosa L. Sp. PL 958. 1753. Granadilla; chilocayo- 
tillo; granadilla roja. 

In moist or dry, often rocky, mixed or oak forest or in thickets, 
1,000-2,000 meters; Pete"n; Chiquimula; Jalapa; Guatemala; Sacate- 
pe"quez; Chimaltenango; Huehuetenango. Southern Florida; western 
Texas; Mexico; British Honduras to El Salvador and Panama; West 
Indies; widely distributed in South America. 

A slender vine, usually small and more or less woody, glabrous to densely 
pubescent, the larger stems with corky bark; stipules linear-subulate, 6-8 mm. long; 
petioles 0.5-4 cm. long, biglandular, the glands small, stipitate, inserted above the 






STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 145 

middle of the petiole; leaf blades very variable in shape, entire to deeply trilobate, 
rounded or sometimes peltate at the base, membranaceous or sub coriaceous, some- 
times ocellate beneath, the lobes linear to broadly ovate, acute or obtuse, erect or 
divergent; flowers solitary or geminate in the leaf axils or sometimes in leafy axil- 
lary racemes, 1-3 cm. broad; bracts minute, setaceous, soon deciduous; sepals 
ovate-lanceolate, subobtuse, greenish yellow; petals none; corona filaments biseri- 
ate, filiform, the outer ones recurved, white, yellow at the apex, purple below; 
ovary subglobose or ovoid, glabrous; fruit globose or ovoid, dark purple or black, 
glaucous when young, 6-15 mm. in diameter; seeds flattened, abruptly acuminate, 
tapering to the base, 3-4 mm. long, 2 mm. broad, coarsely reticulate. 

Called "calzoncillo" in El Salvador; "zac-cansel-ac," "cansel-ac" 

(Yucatan, Maya). 

Passiflora subpeltata Ortega, Nov. Rar. PI. Hort. Matr. 6: 78. 
1798. P. alba Link & Otto, Icon. PL Rar. 65, t. 33. 1828. 

Moist or dry thickets or hedges, or sometimes in forest, 1,200- 
2,000 meters; Chiquimula; Jalapa; Guatemala; Quiche"; Huehuete- 
nango. Central and southern Mexico; Panama; Greater Antilles; 
Colombia and Venezuela. 

Usually a large vine, herbaceous or suffrutescent below, glabrous throughout 
or nearly so, the stems terete, striate; stipules semioblong, 1-4 cm. long, 0.5-2 cm. 
broad, mucronate, entire, or glandular-crenate at the base; petioles 4-6 cm. long, 
bearing 2-4 minute liguliform glands 1 mm. long or shorter; leaf blades 4-6 cm. 
long, 5-12 cm. broad, trilobate to the middle or deeper, subcordate and often sub- 
peltate at the base, glabrous or nearly so, the lobes oblong or oval, 4 cm. broad or 
narrower, rounded or obtuse at the apex, subentire, glandular-serrulate in the 
sinuses; peduncles 4-6 cm. long; bracts ovate-oblong, 1-1.5 cm. long, 1 cm. broad, 
acute or obtuse, cordate and slightly serrulate at the base, inserted near the base 
of the flower; flowers 4-5 cm. broad; sepals oblong, 5-10 mm. broad, obtuse, green 
outside, white within, carinate, the keel terminating in a green foliaceous horn 
1 cm. long; petals linear-oblong, about equaling the sepals; corona filaments 5- 
seriate, white, the outer ones as much as 2 cm. long, filiform, spreading, the inner 
ones only 2-4 mm. long; ovary ovoid, glaucous; fruit ovoid or subglobose, 2.5-4 cm. 
in diameter; seeds obovate, compressed, 5 mm. long, 3 mm. broad, the beak slightly 
curved, finely reticulate. 

Passiflora trinifolia Masters, Bot. Jahrb. 8: 217. 1887. Hoja 
de murcielago; granadilla; calzoncillo. 

Mostly in moist or wet, mixed, often dense, mountain forest, or 
in thickets, 1,200-2,700 meters; endemic; Baja Verapaz (type from 
Santa Rosa, F. C. Lehmann 1314); Zacapa; Jalapa; Guatemala; 
Quezaltenango; San Marcos. 

Usually a rather small vine, the stems slender, terete, minutely puberulent; 
stipules ovate-lanceolate to orbicular, 5-8 mm. long, 4-6 mm. broad, mucronate; 
petioles mostly 1 cm. long or shorter, bearing 2 flattened glands at or slightly above 



146 FIELDIANA: BOTANY, VOLUME 24 

the middle, or the glands sometimes absent; leaf blades bright green, rather thick 
and firm, 1.5-11 cm. long and 2.5-15 cm. broad, trilobate to about the middle, 
more or less cordate and 5-nerved at the base, reticulate-veined, ocellate, glabrous 
or minutely puberulent, the lobes triangular-ovate, subequal or the lateral ones 
somewhat larger, usually acute or acuminate, the lateral lobes general y widely 
spreading; peduncles geminate, equaling or slightly shorter than the petioles, artac- 
Sate above the middle; flowers 1-2 cm. broad, yellowish green; sepals deltoid- 
lanceolate, 7 mm. long, 6 mm. broad, subcoriaceous; petals none; corona filaments 
uniseriate, filiform, dilated toward the apex, 4 mm. long, erect; ovary subglobose, 
glabrous or minutely puberulent; fruit black, lustrous, globose, 1.5 cm. in diam- 
eter; seeds obovoid, 4 mm. long, very coarsely reticulate. 

This plant is very common in the forests of the Occidente. It is 
particularly attractive because of the neat leaves of a handsome shade 
of green, in their form somewhat suggestive of English ivy (Hedera). 

Passiflora Urbaniana Killip, Journ. Wash. Acad. Sci. 17: 426. 

1927. 

British Honduras (Churchyard Pine Ridge, Sibun River; Honey 
Camp). Described from plants grown at Santiago de las Vegas, 
Cuba, the seeds said to have come from British Honduras. 

Stems terete, ferruginous-villosulous or tomentose; stipules minute semiannu- 
lar, barely 0.5 mm. long; petioles 5-10 mm. long, eglandular, densely ferruginous- 
tomentose; leaf blades oblong or lance-oblong, 4.5-9 cm. long, 2-4 cm broad, 
obtuse or rounded at the apex, subcordate at the base, entire or remotely undulate- 
crenulate, coriaceous or subcoriaceous, minutely hispidulous above, densely and 
softly ferruginous-tomentose beneath; peduncles 5-8 cm. long pilosulous; bracts 
2-3 5 cm long, 1-1.5 cm. broad, viscous, deeply 2-pinnatisect; flowers 5 cm. broad, 
the calyx tube short-campanulate; sepals oblong, 2 cm. long, 6 mm. broad, obtuse, 
greenish and hirsutulous outside, 3-nerved, the middle nerve carinate toward 
apex and terminating in a short horn; petals linear, slightly shorter than the sepals, 
lavender; corona filaments 5-seriate, those of the two outer series filiform, 13 
long, violet at the base, radiate, those of the inner series almost capillary 2 mm. 
long, erect; ovary subglobose, densely villous-tomentose; fruit globose, 3.5 cm ir 
diameter, villous; seeds oblong-cuneate, 4 mm. long, 3 mm. broad, shallowly t 
dentate, finely reticulate. 

CARICACEAE. Papaya Family 

Trees, shrubs, or large coarse herbs, with milky sap; leaves usually very large, 
long-petiolate, palmately lobate or palmately 3-few-foliolate; flowers small or 
rather large, usually white or cream-colored, unisexual or rarely perfect; calyx 
small and short; staminate corolla salverform, the tube slender, the 5 short lobes 
valvate or convolute; stamens 10 (or sometimes only 5), inserted in the throat of 
the corolla, the filaments short; anthers adnate to the filaments, 2-celled ; pistillate 
flowers with 5 distinct petals and no staminodia; ovary compound, 1-celled c 
perfectly 5-celled, free, sessile, the stigmas 5, sessile; ovules numerous, in 2 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 147 

series on the 5 placentae; fruit a large, fleshy berry; seeds numerous, compressed 
or subglobose, the testa roughened; endosperm carnose, the embryo axile. 

There are four, perhaps five, rather poorly differentiated genera 
in the family. One genus is tropical African, the others are all trop- 
ical American. There is one other genus in Mexico and another in 
southern Central America. Jacaratia is in cultivation in Guate- 
mala. 

GARICA L. Papaya 

Trees, shrubs, or large herbs, the trunk usually simple, sometimes branched, 
unarmed; leaves long-petiolate, simple and deeply lobate; inflorescences axillary, 
cymose-racemose, the bracts usually very small, the flowers dioecious; staminate 
corolla with an elongate slender tube, the lobes in bud contorted or valvate; sta- 
mens inserted in 2 series in the throat of the corolla, free, the outer ones with 
elongate filaments, the inner ones short, the connective often produced into a 
ligule beyond the anther cells; style short, the 5 stigmas linear or variously cleft; 
fruit small or often very large, the pulp filling it, or a large cavity often present; 
seeds numerous, covered with a succulent membrane, rugose-tuberculate. 

Species perhaps 30 or more, in tropical America, mostly in South 
America. Two others are described from Panama. 

Leaves usually digitately compound, with 3-7 leaflets; sometimes branched trees. 

C. mexicana. 
Leaves simple, lobate; plants usually with simple stems, herbaceous or somewhat 

woody. 
Stigmas palmately cleft or divided; cultivated or wild plants, the fruits mostly 

large and edible, sometimes small and scarcely edible C. Papaya. 

Stigmas pinnatifid; wild plants, the fruits mostly 3.5-5 cm. long, not edible. 

C. pennata. 

Carica mexicana (A. DC.) L. Wms. Fieldiana, Bot. 29: 368. 
1961. Jacaratia mexicana A. DC. Prodr. 15, pt. 1: 420. 1864. Carica 
dolichaula Donn.-Sm. Bot. Gaz. 23: 247. 1897. Leucopremna mexi- 
cana Standl. Contr. U. S. Nat. Herb. 23 : 850. 1924. Pileus mexicanus 
Johnston, Contr. Gray Herb. n.s. 70: 79. 1924. Jacaratia dolichaula 
Woodson, Ann. Mo. Bot. Gard. 37: 404. 1950; 45: 23, /. 1958. Chaya 
silvestre (perhaps an erroneous name). 

Moist or dry lowland forest or scrub, 800 meters or less; Pete"n; 
Alta Verapaz; Huehuetenango; Izabal. Mexico; British Honduras 
through Central America to Panama. 

A tree 5-12 m. tall or more, sometimes blooming when a shrub 3 m. tall, the 
trunk slender or thick, the bark pale brown, almost smooth, glabrous throughout; 
leaves long-petiolate, the leaflets 3-7, oblong or obovate-oblong, sometimes obo- 
vate-elliptic, mostly 9-18 cm. long, cuspidate-acuminate to almost obtuse, acute 



148 FIELDIANA: BOTANY, VOLUME 24 

at the base, petiolulate or sessile, entire or lobate, glaucescent beneath; peduncles 
axillary, few-many-flowered, the flowers fragrant, greenish to cream-colored; ca yx 
1-2 mm long, the sepals oblong-ovate or subtriangular, obtuse; corolla of the 
Distillate flowers 3-7 cm. long, the segments linear or linear-lanceolate; corolla of 
^esfminate flowers 5-6 cm. long, tubular below, the free lobes 1-2 cm. long and 
oblong-elliptic; fruit obovoid, 4-15 cm. long or perhaps even larger, sometimes 
5-angled and produced at the base into cone-like protuberances or wmgs; seeds 
many and large, often filling the single-celled or falsely 5-celled cavity. 

The Maya name for this plant in Yucatan is reported to be 
"cunche" or "cumche." In other regions of Mexico it is called bo- 
nete orejona" or "papaya orejona." In Mexico it is said that the 
fruits are sometimes eaten raw, especially in salads, or that they are 
made into sweetmeats of various kinds. When maize is scarce the 
Mayas of Yucatan are said to pulverize the pith of the plant and 
mix it with maize in making tortillas, although this seems unlikely. 

Carica Papaya L. Sp. PI. 1036. 1753. Papaya (fruit); papayo 

(plant) . 

Cultivated everywhere at 1,200 meters or lower, and sometimes 
at even higher elevations; often found wild in waste ground, second 
growth, and generally in moist or wet thickets, especially along the 
lowlands of the Atlantic watershed. Native of tropical America, 
but the original home unknown; now grown generally in almost all 
tropical regions. Figure 17. 

Plants with a rather thick, usually simple, columnar trunk at maturity, the 
trunk bearing conspicuous petiolar scars, commonly 3-6 meters high, leafy at the 
ton leave" very large, on long petioles, mostly 30-60 cm. wide but sometimes 
arg ; ur a ny palmateiy 7-lobate, more or less glaucous beneath, the lobes pinnate- 
bbate obtuse or acute; staminate flowers creamy white, in usually lax, peduncu- 
te panicles 10-30 cm. long; staminate calyx 1-1.5 mm. long, the corolla 2.5-3 cm. 
ong the Yobes lanceolate or elliptic-lanceolate, equaling or somewhat porter than 
the tube pistillate sepals 5-10 mm. long, the corolla longer than that of the stami- 
nate flowers the petals distinct, lanceolate, contorted; fruit very variable in shape 
and ste t h ;t of wild plants small and globose-obovoid, that of cultivated plants 
often veiy large and obovoid to oblong, yellow or orange at maturity, more or less 
leaked ^ith green, with a somewhat milky juice, the flesh yellow or orange, very 
juicy and sweet. 

The Maya names of Yucatan are "put" and "chichput" (the wild 
form) The term "papaya" is of Antillean origin, and is used for 
the fruit in most regions, except in Cuba, where it h ^ f com % to jf v ,! 
an obscene meaning, and the papaya there is called "fruta bomba^ 
The Carib name is said to have been originally ababaya LI 
papaya is one of the best of tropical fruits, and many people believe 
that it has few superiors anywhere. It is one for which it is not 




FIG. 17. Carica Papaya. Habit of plant, fruit; both much reduced. 



149 



150 FIELDIANA: BOTANY, VOLUME 24 

necessary to cultivate a taste; most northern people like it upon first 
trial and become very fond of it if they live long in the tropics. 
Almost all papayas are good when thoroughly ripe, and when stewed 
they also are excellent. The fruits vary greatly in shape and size, 
some being extremely large, especially those of Panama. There are 
numerous varieties in Guatemala, but in selecting .them for the table 
little attention seems to be paid to the variations. While found 
everywhere in Guatemala at 1,200 meters or lower, and little planted 
above that limit, the fruits may be found on sale in all markets, for 
they are carried up from the lowlands in trucks or on the backs of 
cargadores, to the highland markets. Many of the fruits seen in the 
markets are scored with longitudinal knife cuts. It is claimed that 
if this is not done, the fruit is bitter; but quantities of unscored fruits 
sold in the markets and bought readily by the people seem just as 
sweet as those thus treated ! The plants are essentially herbaceous. 
They are grown from seeds, and begin to bear when only about 2 
meters high, continuing until they reach a height of about 6 meters. 
When so high, the heavy fruits are likely to cause the plants to fall 
to the ground, especially in case of a strong wind. In protected 
places in the Pacific lowlands extremely tall stems sometimes are 
seen, perhaps 9 meters high, but these plants seldom have fruits, 
and their terminal tuft of leaves is small. The trunks for the most 
part are simple, but it is not unusual to see a plant with a few thick 
branches at the top. 

The larger papayas are sometimes as much as 50 cm. long, with 
a weight of 20 pounds, but such a size is unusual. In appearance 
and flavor the flesh is much like that of a cantaloupe. The imma- 
ture green fruits are cooked and eaten in some regions, but this is 
unusual in Central America, even though they "make better apple pie 
than apples do." A sirup made from the fruit is used in Central 
America as a remedy for coughs. The dried and powdered seeds as 
well as the milky sap are administered internally to kill intestinal 
parasites, and the milk is said to be applied to kill niguas (tropical 
chiggers). 

It has long been known that the milk that exudes from all parts 
of the plant contains a substance called papain, similar in properties 
to animal pepsin. Tough meat wrapped in papaya leaves and left 
overnight is said to become tender, or some leaves may be boiled 
with the meat. This fact is widely published and reported, but it 
seems to be better known abroad than in Central America. If such 
use is made there of the plant, it is not common, and it is not be- 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 151 

cause the average meat of the region does not need such treatment! 
In recent years the papaya has become known to some extent in the 
United States. In favorable seasons it grows and produces well in 
southern Florida, and the fruits reach northern markets in small 
quantities. 

Carica pennata Heilborn, Svensk. Bot. Tidskr. 30: 222, /. 4. 
1936. Papayo de monte; papayo de montana. 

Moist or dry, rather open forest, thickets, and open clearings, 
often in sandy soil near streams on the plains, Pacific plains and 
bocacosta, 1,300 meters or less; Santa Rosa; Escuintla; Sacatepe"quez; 
Suchitepe'quez (described from plants grown in Sweden, the seeds 
collected at Tiquisate by N. Johansson in 1929); Retalhuleu; Que- 
zaltenango; San Marcos. Probably also in El Salvador. 

Plants herbaceous and usually 2 meters high or less, apparently annual, the 
stems slender; leaves scattered along the stems or at maturity mostly near the top 
of the stem, very large or small, long-petiolate, mostly 5-lobate, deeply cordate 
at the base, the middle lobe pinnatifid with acute to long-attenuate segments, the 
other lobes 3-fid or 2-fid, glaucous beneath; staminate inflorescences short or elon- 
gate and many-flowered, the calyx 1.5 mm. long; corolla white, fading to yellow- 
ish, 3-3.5 cm. long, the lobes usually much shorter than the tube; pistillate inflores- 
cences few-flowered, the flowers 2.5 cm. long, the sepals minute; petals lanceolate 
or linear, 5 mm. broad, acuminate; style 1 mm. long, the 5 stigmas 8 mm. long, 
pinnatifid with 5-7 lobes; fruit globose-obovoid, usually 3-5 cm. long or often 
somewhat larger, yellow at maturity; seeds 3 mm. long or slightly larger, very 
numerous. 

It is assumed that this species is the common wild papaya of the 
Pacific plains and foothills. It grows there abundantly, forming ex- 
tensive stands in open places, and often growing profusely in the 
more open forest. Most of the plants appear to last only a single 
season, dying if the soil becomes very dry. The fruits are small, 
orange-yellow within, and of poor flavor. They are not eaten. 
There is not available satisfactory material for testing the value of 
the stigma character on which Heilborn separated this species from 
C. Papaya. Tropical plants grown in northern hothouses often be- 
have queerly, and the stigma characters should be investigated in 
the field before too much importance is attached to them, in spite of 
the fact that the author of the species established a separate sec- 
tion for it. 



Jacaratia dodecaphylla (Veil.) A. DC., a tree native in Brazil, is 
in cultivation at Finca Panama, Suchitepe'quez. It is a good-sized 



152 FIELDIANA: BOTANY, VOLUME 24 

tree with very prickly trunk and branches, the long-petiolate leaves 
palmately compound, with 7 or more, entire, narrow leaflets, glau- 
cous beneath. 

LOASACEAE 

Reference: I. Urban & E. Gilg, Monographia Loasacearum, Nova 
Act. Leop. Carol. Acad. 86: 1-380. 1900. 

Herbs or rarely shrubs, erect or scandent, dichotomously branched, often cov- 
ered with stinging hairs; hairs of the leaves usually rough and often glochidiate or 
anchor-shaped; leaves opposite or alternate, entire, lobate, pinna tifid, or 2-3 times 
pinnatifid; stipules none; flowers small or large, regular, perfect; inflorescence of 
solitary flowers, racemose or cymose, the peduncles often opposite the leaves, the 
pedicels bibracteolate; corolla usually white or yellow; calyx tube adnate to the 
ovary, the calyx lobes 4-5, imbricate or contorted, herbaceous or chartaceous, 
persistent; petals 4-5, inserted in the throat of the calyx, sessile or unguiculate, 
flat or cucullate, induplicate-valvate; petaloid scales alternate with the petals, 
cucullate, sometimes with dorsal appendages; stamens 4-12 or numerous, inserted 
with the petals, the filaments filiform, often collected in fascicles opposite the 
petals; anthers oblong or didymous, introrsely and longitudinally dehiscent, the 
filaments filiform; staminodia present between the stamens and petals or absent, 
sometimes petaloid; ovary inferior, rarely half superior, 1-celled or rarely 2-3- 
celled, confluent with the calyx tube, often hispid at the free apex; style filiform 
or stiff, subulate, entire, or 2-3-fid, the lobes linear; stigma punctiform or obtuse; 
ovules solitary or numerous, affixed at the apex of the cell or on parietal placentae, 
pendulous, anatropous; capsule generally 1-celled, often costate, the costae straight 
or spirally twisted, 3-5-valvate at the apex or longitudinally 3-10-valvate, the 
valves alternate with the placentae, the seeds 1-many, usually small, oblong or 
angulate; testa membranaceous or cellulose, often laxly reticulate; endosperm 
none or carnose, rarely corneous; embryo straight, linear, oblong, or cylindric; 
cotyledons plano-convex and foliaceous or hemispheric, the radicle short or 
elongate. 

About 14 genera, all except one confined to America. One other 
genus, Klaprothia, is known from Costa Rica and Panama, two 
others from Mexico. 

Ovary 1-ovulate; plants scandent; leaves palmate-nerved, deeply cordate at the 

base; plants covered with stinging hairs. Gronovia. 

Ovary with few-many ovules; plants not scandent. 

Plants covered with stinging hairs; leaves pinnately parted Loasa. 

Plants without stinging hairs; leaves merely dentate or shallowly lobate. 

Leaves all opposite; flowers minute, the petals 2 mm. long or less: capsule 

spirally twisted Sclerothrix. 

Leaves all or mostly alternate; flowers much larger; capsule not spirally 
twisted. 

Capsule 3-carpellate Mentzelia. 

Capsule 5-carpellate Eucnide. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 153 

EUGNIDE Zuccarini 

Annuals or sometimes more enduring, with brittle stems, the pubescence 
mostly of long and soft hairs, but these often barbellate at the apex and more or 
less irritant; lower leaves opposite, the upper ones alternate, usually long-petiolate; 
flowers 5-parted, solitary or in cymes, often very large, yellow or white; calyx tube 
subglobose to campanulate, the sepals imbricate; petals imbricate, united at the 
base; stamens numerous, in 2-5 series, united at the base, united with the petals 
and falling with them; staminodia none; style filiform; ovary inferior, with 5 or 
rarely 4 placentae, the ovules very numerous, horizontal; capsule obovoid or ellip- 
soid, dehiscent by 5 apical valves; stamens numerous, small, cylindric or elongate; 
endosperm present, the embryo straight. 

About 10 species, ranging from southwestern United States to 
Guatemala. 

Flowers about 2.5 cm. long E. bartonioides. 

Flowers about 7.5 cm. long E. guatemalensis. 

Eucnide bartonioides Zucc. Linnaea 18: 509. 1844. 

Known in Guatemala only from Guastatoya, El Progreso, 300 
meters, W. A. Kellerman 8062. Southern Mexico. 

Plants probably annual, simple or branched, 40 cm. high or less, densely pilose 
throughout with slender spreading whitish hairs, many of these barbate at the apex, 
the stems pale; leaves mostly on long slender petioles, rounded-cordate, thin and 
soft, mostly 2.5-6.5 cm. long and almost or quite as broad, rounded or obtuse at 
the apex, deeply cordate at the base, shallowly lobate and sinuate-dentate, pal- 
mately nerved; peduncles axillary, 1-flowered, short at first but in age often much 
elongate, sometimes longer than the leaves; calyx densely long-pilose with spread- 
ing hairs, about 12 mm. long, the lobes linear-lanceolate, longer than the tube; 
corolla yellow, about 2.5 cm. long, the petals rounded at the apex, erect, glabrous or 
nearly so; stamens slightly or much exceeding the petals; capsule about 1 cm. long. 

The hairs, especially on the calyx, are often much thickened at 
the base. 

Eucnide guatemalensis Standl. & Steyerm. Field Mus. Bot. 23: 
178. 1944. 

Known only from the type, Huehuetenango, Cerro Chiquihui, 
above Carrizal, about 2,000 meters, Steyermark 50818. 

A coarse branched herb, the branches thick, densely viscid-hispid; leaves on 
stout petioles 2.5-5.5 cm. long, rounded in outline or ovate-rounded, 5-9 cm. long, 
4.5-8.5 cm. broad, deeply and narrowly cordate at the base, very shallowly 5-9- 
lobate, the lobes low and rounded, undulate or undulate-dentate, densely hispid 
on both surfaces; flowers numerous, the peduncles stout, 3-6 cm. long, densely 
viscid-hispid; hypanthium obconic or broadly turbinate, 9 mm. long, 10-12 mm. 
broad at the apex, densely hispid; sepals oblong-linear or narrowly lanceolate, 
3-3.5 cm. long, 5-7 mm. broad, long-acuminate, densely viscid-hispid; petals about 



154 FIELDIANA: BOTANY, VOLUME 24 

5.5 cm. long, and almost 2 cm. broad, rounded at the apex, whitish green, very 
sparsely pilose outside with long, soft, white hairs or almost wholly glabrous; 
stamens very numerous, longer than the petals, about 7 cm. long, the anthers 
1.5 mm. long; capsule 1.5 cm. long. 



GRONOVIA L. 

Scandent herbs, pilose and provided with stinging hairs, the hairs of the stems 
uncinate at the apex; leaves alternate, long-petiolate, broadly cordate, 5-lobate; 
flowers small, pedicellate, subcorymbose, the inflorescences opposite the leaves, 
few-many-flowered, the pedicels bearing small linear bractlets; calyx tube short, 
oblong, subangulate; sepals 5, united below; petals 5, yellowish green, inserted in 
the throat of the calyx, shorter than the sepals and narrower, linear-spatulate; 
stamens 5, inserted with the petals, the filaments filiform, puberulent, the anthers 
oblong; staminodia none; ovary inferior, 1-celled, the style inserted on an epigy- 
nous disc, filiform, the stigma capitellate; ovule solitary, pendulous from the apex 
of the cell; fruit small, crowned by the persistent calyx, subglobose, coriaceous, 
costate, indehiscent; seed with membranaceous-striate testa; endosperm none; 
cotyledons carnose, the margins lobulate, induplicate. 

The genus is wholly American. One or two other species are 
known in Mexico. 

Gronovia scandens L. Sp. PL 202. 1753. Guisquilillo (fide 
Aguilar) . 

Moist thickets or waste ground, 1,500 meters or less; Zacapa; 
Santa Rosa; Guatemala; Suchitepe*quez; Huehuetenango. South- 
ern Mexico; El Salvador and Honduras to Panama; southward to 
Ecuador. 

Usually a small herbaceous vine with rather succulent stems, the stems bear- 
ing numerous long uncinate hairs mixed with shorter ones, sparsely branched; 
leaves thin, on petioles 3-6 cm. long, the blades 3-8 cm. long, deeply 5-lobate, 
deeply cordate at the base, the terminal lobe subulate-acuminate, the lobes all 
sublobate or coarsely dentate, sparsely or densely pilose on both surfaces, part of 
the hairs simple, part of them glochidiate at the apex; inflorescences pedunculate, 
10-18-flowered; calyx 4-7 mm. long, yellowish green, the adnate basal portion 
3-4 mm. long, the lobes oblong, acute; stamens 2-3 mm. long, the petals scarcely 
longer; fruit obovoid or short-cylindric, 2.5-4 mm. long. 

Known in El Salvador as "pan caliente," "picapica," "chichicaste" 
and "Juan caliente"', "lalmuch" (Yucatan, Maya); "pan caliente" 
and "no me tientes" in Honduras; "pringamoza" or "pringamosca" 
in Nicaragua. The hairs of the plant sting severely and painfully. 
As remarked by Bentham and Hooker, the general appearance of 
this weedy plant is suggestive of many Cucurbitaceae, which the 
Guatemalan common name also suggests. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 155 

LOASA Jussieu 

Herbs, usually erect, hispid and setose, the hairs often stinging; leaves alter- 
nate or opposite, entire, lobate, or decompound; flowers large or small, axillary, 
solitary, racemose, or subpaniculate, mostly white or yellow; calyx tube turbinate 
to ovoid, clavate, or cylindric, the 5 lobes equal; petals 5, cucullate, spreading or 
suberect, often connivent, alternating with 5 scales, these 2-3-setose dorsally and 
often appendaged at the base; stamens very numerous, the filaments filiform, col- 
lected in fascicles opposite the petals, the anthers didymous or oblong; staminodia 
10, filiform; ovary 1-celled, the style subulate, the stigma obtuse, acute, or shortly 
3-fid; ovules numerous, affixed to 3-5 parietal placentae; capsule terete or costate, 
crowned by the persistent calyx, 3-valvate at the apex, many-seeded; seeds small 
or large, oblong or angulate, the testa usually cellulose and spongious; endosperm 
carnose, the embryo terete or compressed, the cotyledons plano-convex, rarely 
foliaceous. 

Probably 100 species, all in tropical America and mostly South 
American. Two other species, with large, showy, yellow flowers, are 
found in Costa Rica and Panama. 

Loasa triphylla Juss. Ann. Mus. Paris 5: 27, t. 5, f. 2. 1804. 

A plant of wide distribution in western South America, repre- 
sented in southern Mexico and Central America by the following 
variety: 

Loasa triphylla var. rudis (Benth.) Urban & Gilg, Nova Act. 
Leop. Carol. Acad. 86: 239. 1900. L. rudis Benth. PI. Hartweg. 75. 
1839 (type from Santa Maria, probably in Quezaltenango, Hartweg} . 
Chichicaste. 

Moist or wet thickets or forest, often along streams, sometimes 
on white sand slopes, 300-2,800 meters; Alta Verapaz; Retalhuleu; 
Quezaltenango; San Marcos. Southern Mexico; Costa Rica; Panama. 

A rather weak, usually erect or sprawling herb, a meter high or less, pubescent 
everywhere with long hairs glochidiate at the apex and with long, fulvous, simple 
setae; leaves long-petiolate, the blades often 15 cm. long or more, the lower ones 
with 2-4 pairs of petiolulate leaflets, these oval or oval-oblong, serrate, the upper 
leaves simple, short-petiolate; flowers rather few, large, in secund racemes, long- 
pedicellate, about 2 cm. broad; petals white, 1 cm. long, caducous; capsule erect, 
13-16 mm. long, 1 cm. broad above, turbinate-clavate, echinate with long, fulvous 
setae; seeds blackish, obovate, 1 mm. long. 

The plant is all too plentiful in many parts of the mountains or 
plains of western Guatemala, growing in abundance where it is 
kept permanently wet by the spray of water. The plant stings badly, 
leaving a persistent itching sensation. It is able to sting, like a nettle, 
through the cloth of ordinary trousers. 



156 FIELDIANA: BOTANY, VOLUME 24 

MENTZELIA L. 

Reference: Josephine Darlington, A monograph of the genus Ment- 
zelia, Ann. Mo. Bot. Card. 21: 103-226. 1934. 

Annual or perennial herbs, rarely shrubs or small trees, usually erect, covered 
with various kinds of rigid, barbate, but not stinging hairs, the stems usually 
whitish or yellowish in age; leaves mostly alternate, lobate, dentate, or pinnatifid; 
inflorescence terminal, cymose, the flowers sometimes solitary, small or large, white 
or yellow; calyx tube cylindric to ovoid or turbinate, the limb 5-lobate, the lobes 
imbricate, deciduous or persistent; petals 5-10, imbricate, free or united and nar- 
rowed or contracted at the base, apiculate; stamens 10-many, inserted with the 
petals but free from them; filaments distinct or in clusters opposite the petals, 
filiform or linear, or the 10 outer ones broadened, sometimes petaliform, in 1-5 
series; anthers introrse, laterally dehiscent; style elongate, filiform, trifid at the 
apex; ovary 1-celled, 2-many-ovulate, the placentae usually 3; capsule short, cylin- 
dric, oblong or turbinate, more or less distinctly costate at maturity, 3-7-valvate 
at the apex; seeds flat or angulate, sometimes winged; endosperm copious or scant, 
the embryo erect or curved, the radicle conic or subterete. 

Species about 60, all American. Only one is known from Central 
America. 

Mentzelia aspera L. Sp. PI. 516. 1753. Pegapega. 

Moist or dry thickets, often in rocky places, 200-1,500 meters; 
Zacapa; Jutiapa; Baja Verapaz; Guatemala; Quiche"; Huehuetenango. 
Arizona and Mexico; British Honduras; Honduras; Nicaragua; Pan- 
ama; West Indies southward to Brazil and Bolivia. 

Plants annual, often much branched from the base or throughout, erect or 
ascending, a meter high or usually lower, the stems generally yellowish, very brittle, 
covered with short, glochidiate hairs intermixed with antrorse, spine-like ones; 
leaves alternate, lanceolate or ovate-lanceolate, 3-10 cm. long, slender-petiolate, 
acute or short-acuminate, serrate or dentate and often hastately trilobate, rough- 
pubescent on both surfaces with chiefly glochidiate hairs; flowers sessile, axillary; 
calyx tube terete, attenuate to the base, 6-11 mm. long, 2-3.5 mm. broad; calyx 
lobes lanceolate or linear-lanceolate, long-acuminate, 3-4 mm. long or in age longer; 
petals obovate to obovate-orbicular, acute, 4-8 mm. long; stamens 20-30, the outer 
ones dilated and petaloid; capsule cylindric, 1.5-2.5 cm. long, 3-5 mm. thick, in- 
conspicuously striate, 3-valvate at the apex; seeds 6-9, grayish yellow, irregularly 
tuberculate, undulate-striate, not winged. 

Called "pegarropa" in Yucatan, and the Maya names are reported 
as "tzayuntzay" and "tsootscab." An unattractive plant and unpleas- 
ant to handle, although it does not sting. The rough leaves adhere 
tenaciously to clothing. 

SCLEROTHRIX Presl 

Annual or longer-lived, slender, weak herbs. Leaves opposite, petiolate, usu- 
ally membranaceous, simple; inflorescence racemose, few-flowered, arising in the 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 157 

axils of leaves; calyx tube narrow, costate, the costae spirally twisted; calyx lobes 
4, triangular; petals 4, sessile, obovate, apiculate; stamens 4-12, 1-3 of them oppo- 
site the petals; anthers small, oblong; ovary inferior, 1-celled; style subulate; cap- 
sule clavate, hispid-hirsute, few-many-seeded. 

A small genus of one or perhaps two species ranging from Mexico 
through Central America well into South America. 

Sclerothrix fasciculata Presl, Symb. Bot. 2: 3, t. 53. 1833. 
Ancyrostemma micranthum Poepp. & Endl. Nov. Gen. & Sp. 3: 65, 
t. 272. 1845. Pegapega. 

In old fields or more often in sand or gravel along streams, occa- 
sionally in the forest, from 300-1,600 meters; San Marcos; Re- 
talhuleu; Quezaltenango; Chimaltenango; Escuintla; Suchitepe"quez. 
Mexico through Central America south to Peru and Bolivia. 

Small, herbaceous plants with succulent stems to 50 cm. tall; the stem sparsely 
to densely pubescent with retrorse hairs, somewhat branched or nearly simple; 
leaves lanceolate to broadly elliptic or ovate, acute or acuminate, serrate, ap- 
pressed hirsute pubescent on both surfaces, 2-10 cm. long and 0.7-4 cm. broad; 
petioles slender, mostly about 1 cm. long; inflorescences few-flowered, axillary 
racemes, pubescent; calyx clavate, costate and spirally twisted, densely short, stiff 
hirsute pubescent; calyx lobes short, triangular or nearly so, acute, about 1 mm. 
long; petals about 1 mm. long, sessile. 

BEGONIACEAE. Begonia Family 
By LYMAN B. SMITH and BERNICE G. SCHUBERT 

References: A. DC. in DC. Prodr. 15, pt. 1: 266. 1864. Irmscher 
in Engler & Prantl, Pflanzenfam. ed. 2. 21: 548-588. 1925. C. Che- 
valier, Les Begonias, 1938. 

Annual or perennial herbs, shrubs or rarely small trees; stem elongate and 
succulent or woody or sometimes reduced to a tuberous rhizome, sometimes 
climbing; leaves usually alternate or rarely subverticillate, entire, serrate, lobed 
or digitately parted, usually asymmetric, sometimes peltate; stipules 2, free, 
persistent or deciduous; peduncles axillary; inflorescence mostly cymose, some- 
times 1-flowered or racemose, cymes regular or strongly one-sided; flowers unisexual; 
tepals free or connate; stamens indefinite, inserted on the receptacle, filaments 
free or united; styles usually 3, free or connate, usually bifid; ovary inferior in 
the American species, usually 3-celled and 3- winged or 3-horned; placentae usually 
axile, simple or divided; fruit usually capsular. 



BEGONIA L. 

General characters of the family, of which it contains more than 90 per cent 
of the species; tepals free, usually 4 in two pairs, in staminate flowers, and 5 in 



158 FIELDIANA: BOTANY, VOLUME 24 

pistillate flowers; placentae simple or divided; fruit usually bearing 3 unequal 
wings. 

About 800 species. Nearly pantropical. We take the following 
from Standley's notes: "The species or varieties of Begonia grown in 
Guatemalan gardens or as pot plants are numerous. They are largely 
those seen commonly in the United States and are probably of South 
American origin. It is noteworthy that of all native Central Amer- 
ican species only B. heracleifolia has much if any importance as an 
ornamental plant. Most of the others are decidedly inferior in orna- 
mental value, and many of them decidedly unattractive. Rex bego- 
nias are noticed frequently as pot plants, especially in the mountains, 
but they must be protected from wind and rain. The cultivated 
plant known in the United States as Begonia rubra [a form or hybrid 
of B. maculata Raddi] is a common garden plant of Guatemala. In 
the gardens of Coban the plants of this species often rise well above 
the garden walls." 

Plants acaulescent or subacaulescent with the leaves clustered near the apex of 

the stem, or with slender creeping rhizomes if the internodes elongate. 
Leaves peltate. 

Petiole with a ring of large lacerate scales at its summit and usually several 
more rings below; inflorescence of many small flowers; capsule- wings 

lunate B. manicaia. 

Petiole glabrous or if pubescent the indument not in rings. 
Inflorescence regularly cymose, about as broad as long. 

Leaves 17-40 cm. long; inflorescence many-flowered. . .B. nelumbiifolia. 
Leaves 5-10 cm. long; inflorescence few-flowered. 

Staminate and pistillate tepals 2, suborbicular; the only capsule-wing 

narrow, falcate-ascending B. Calderonii. 

Staminate tepals 4, narrowly elliptic; pistillate tepals 5; largest cap- 
sule-wing ovate B. sciadiophora. 

Inflorescence strongly unilateral, elongate; staminate tepals 4, pistillate 

tepals 5 B. militaris. 

Leaves basifixed. 

Leaves palmately compound B. caroliniifoUa. 

Leaves simple. 

Leaves deeply palmate-lobed. 
Inflorescences strongly unilateral. 

Flowers appearing with the leaves; bracts persistent. .B. heracleifolia. 

Flowers appearing before the leaves; bracts deciduous . .B, crassicaulis. 

Inflorescences scarcely or not at all unilateral; flowers appearing with the 

leaves; bracts deciduous. 
Stipules deciduous; staminate tepals 4; pistillate tepals 3; fruit deflexed 

with largest wing basal B. acutiloba. 

Stipules persistent; staminate tepals 2 or rarely 4; pistillate tepals 2 or 

rarely 3; fruit erect with largest wing central B. Lindleyana. 

Leaves not lobed or with the lobes few and superficial. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 159 

Pistillate tepals 4, subequal; capsule- wings rounded, unequal to subequal; 

petiole tomentose B. pinetorum. 

Pistillate tepals 2, or if 3 with the inner one much reduced. 

Capsules erect or suberect (imperfectly known in B. confusa); leaves 

never bullate. 

Petiole with a ring of large lacerate scales at its summit and usually 
several more rings below; inflorescence of many small flowers; 

capsule-wings lunate, subequal B. manicata. 

Petiole glabrous or the indument not in rings. 

Indument of the petioles consisting of fimbriate scales. 

Largest capsule- wing narrower than high; anthers longer than 

the filaments; leaves entire or slightly serrate . . B. strigillosa. 

Largest capsule- wing wider than high; anthers much shorter 

than the filaments; leaves ciliate-denticulate . .B. stigmosa. 

Indument of the petioles, if any, consisting of simple linear tri- 

chomes. 
Tepals fleshy, at least the outer ones bearing some indument. 

Leaves oblique, up to 6 dm. long, petioles 2-4 dm. long, 
densely retrorse-pilosulous; largest capsule- wing dolabri- 
form or subovate, 25-30 mm. wide B. fusca. 

Leaves straight, not over 18 cm. long, petioles 16-23 cm. long, 
sparsely pilose; internodes 1-9 cm. long, not over 5 mm. 

thick B. confusa. 

Tepals thin, wholly petaloid, glabrous or (in B. Lindleyana) with 

an evanescent pubescence. 

Stipules 1-nerved, thin; capsule- wings subequal, all narrowly 
lunate, rounded; tepals bright red B. tincioria. 

Stipules with many thick nerves; capsule- wings usually very 

unequal, at least the largest somewhat angled. 
Cyme nearly or quite regular; petioles more or less fuscous- 
lanate B. Lindleyana. 

Cyme strongly unilateral; petioles usually soon glabrous. 

B. plebeja. 

Capsules reflexed (imperfectly known in B. confusa), with one wing 
very large; stem usually slender and repent with distinct inter- 
nodes. 

Leaves bullate, broadly ovate, oblique B. pustulata. 

Leaves not bullate. 

Leaves symmetric, straight, cordate-ovate, undulate but not lobed; 
stipules lacerate B. trichosepala. 

Leaves asymmetric at least at base, mostly oblique, sometimes 
lobed. 

Petioles densely pilosulous or hirsute; leaves ample, 1-6 dm. 
long, deeply cordate at base, shallowly if at all lobed. 

Cymes much-branched; leaves shallowly lobed B. fusca. 

Cymes few-branched; leaves not lobed B. Popenoei. 

Petioles sparsely pilose to glabrous; leaves mostly lobed. 

Tepals glabrous; stipules mostly persistent B. ludicra. 

Tepals pubescent; stipules deciduous. 



160 FIELDIANA: BOTANY, VOLUME 24 

Leaves straight, broadly ovate, shallowly and bluntly few- 
lobed; outer tepals covered with dark red septate tri- 

chomes B. confusa. 

Leaves obliquely elliptic, deeply and sharply lobed; tepals 

puberulent B. acutiloba. 

Plants caulescent with erect or ascending stems and distinct internodes. 

Leaves peltate, densely pale-lanate at least beneath B. peltata. 

Leaves basifixed. 

Ovary bearing 3 equal, narrowly triangular acuminate horns; leaves oblique, 

cordate-ovate B. Heydei. 

Ovary bearing broad or marginiform, usually unequal wings. 

Leaves symmetric, ovate, very sparsely undulate-serrate to entire; stami- 

nate tepals 4, pistillate 5 B. glabra. 

Leaves asymmetric, usually oblique. 

Anthers spherical or obovoid; staminate tepals 4; pistillate tepals usually 
5, rarely 3 (not known in B. cebadillensis). 

Pistillate tepals 3; capsule-wings subequal, narrow B. oaxacana. 

Pistillate tepals 5 (unknown in B. cebadillensis) . 

Tepals glabrous; inflorescence 1- (rarely 2-) flowered. 
Outer staminate tepals broadly reniform, entire; stipules and 

bracts minute, scarious, deciduous B. cebadillensis. 

Outer staminate tepals broadly ovate or elliptic, often serrate; 
stipules and bracts large, foliaceous, colored, persistent. 

B. gradlis. 

Tepals pubescent, the outer serrate; inflorescence usually several- 
flowered; bracts deciduous. 

Stipules deciduous B. biserrata. 

Stipules persistent, foliaceous B. ignea. 

Anthers much longer than broad; staminate tepals 2-4; pistillate tepals 2, 

3 or 5. 
Bracts persistent, incised; pistillate tepals 5. 

Capsule- wings subequal; seeds obtuse; stamens on a column. 

B. Franconis. 

Capsule-wings very unequal, the largest being broadest well above 

the middle; seeds acuminate; stamens free B. Fischeri. 

Bracts deciduous; pistillate tepals 2 or 3. 

Cymes few-flowered (not truly erect-caulescent types but some- 
times may be confused with them). 

Tepals glabrous; stipules mostly persistent B. ludicra. 

Tepals pubescent; stipules deciduous B. acutiloba. 

Cymes many-flowered. 

Leaves pinnate-nerved, obliquely elliptic B. convallariodora. 

Leaves palmate-nerved. 

Lobes of the leaf narrowly triangular, acuminate, denticulate; 
ultimate branches of the inflorescence nearly aborted, mak- 
ing the flowers appear umbellate; largest capsule- wing 

spreading, elliptic or falcate B. involucrata. 

Lobes of the leaf short and broad, entire; branches of the in- 
florescence well developed; largest capsule- wing ovate. 

B. Sartorii. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 161 

Begonia acutiloba Liebm. Vid. Medd. Kjoebenhavn 1852: 14. 
1853; emend. Smith & Schubert, Contr. Gray Herb. 161: 28, t. 3. 
1946. 

Forested slopes, 1,800-2,000 meters; Huehuetenango (Cerro Chi- 
blac, between Ixcan and Finca San Rafael, Sierra de los Cuchuma- 
tanes, Steyermark 49379). Southern Mexico. 

Herbaceous; rhizome erect or repent with very short to distinct internodes, 
glabrous, purplish red; stipules deciduous, not known; petioles 15-24 cm. long, 
moderately hirsute, becoming less so; leaf -blades asymmetric, obliquely elliptic, 
sinuate-lobate with about 5 acuminate lobes, cordate at base, 20-30 cm. long, 
irregularly denticulate, rufous-hirsute on the nerves beneath; peduncles 25-30 cm. 
long, puberulent; cymes regular, few-flowered; bracts quickly deciduous, ovate- 
oblong, obtuse, entire, thin, glabrous; pedicels 1-2 cm. long, rufous-puberulent; 
staminate tepals 4, entire, white, the outer ovate, obtuse, 10-14 mm. long, pubescent 
with light brown trichomes, the inner narrowly obovate, 6 mm. long; anthers 
oblong, obtuse; pistillate tepals 3 (only very immature ones known), entire, 
the two outer ovate, densely tomentose, the inner one much smaller; styles 3, 
bifid; capsule deflexed, ellipsoid, unequally 3-winged, the largest wing subbasal, 
oblong, obtuse, half the height of the capsule. 

Begonia biserrata Lindl. Journ. Hort. Soc. 2: 313. 1847; Hook. 
Bot. Mag. 79: t. 4746. 1853. Knesebeckia biserrata Klotzsch, Begon. 
45. 1855. Begonia palmaris A. DC. Ann. Sci. Nat. se>. 4, 11: 126. 
1859. B. palmata Sesse" & Mocino, PI. Nov. Hisp. 163. 1890, not 
Don. 

Moist thickets and forests, 1,400-2,000 meters; Chiquimula; San- 
ta Rosa; Guatemala; Quiche"; Huehuetenango. Southern Mexico. 

Herbaceous, 6-9 dm. high, apparently dioecious; stem erect, usually simple, 
pubescent, internodes elongate; stipules soon deciduous; petioles 1-13 cm. long; 
leaf-blades obliquely palmate with 4 major nerves and 4-6 acute to acuminate 
lobes, cordate with a cuneate to truncate center, to 19 cm. broad and nearly as 
long, sharply and doubly serrate, nearly concolorous, pubescent on both sides; 
inflorescence unilateral, peduncles in the axils of reduced leaves, 15-65 mm. long, 
2-6-flowered; bracts rapidly deciduous, ovate or lanceolate, acute, 6 mm. long, 
serrulate; pedicels hirsute, to 25 mm. long; flowers more or less nodding; staminate 
tepals 4, serrate, the outer broadly ovate, 10-25 mm. long, pubescent, the inner 
lanceolate, much smaller; stamens numerous in a spherical cluster, filaments 
much longer than the subspherical anthers, connective not produced; pistillate 
flowers bracteolate, tepals 5 with 3 large outer ones, like the staminate; ovary 
3-celled, placentae bifid, ovuliferous on all sides, styles connate at base, bifid with 
the stigmatic surface linear, spiral and continuous; capsule ellipsoid, 15-17 mm. 
long, pubescent, wings oblong, obtuse, unequal, the largest 17 mm. wide; seeds 
ellipsoid, blunt. 

Begonia Calderonii Standl. Field Mus. Bot. 8: 27. 1930 (type 
from El Salvador, Calderon 2416). B. falcata Smith & Schubert, 



162 



FIELDIANA: BOTANY, VOLUME 24 




FIG. 18. Begonia Calderonii. A, Habit; X Y 2 . B, Staminate flower; X 1. 
C, Stamen; X 5. D, Pistillate flower; X 1. E, Style; X 5. F, Fruit; > 

Contr. Gray Herb. 127: 28, t. 2, figs. 19-21. 1939 (type from Suchi- 
tepe"quez, Skutch 1485). 

In forest usually on rocks, 1,250-2,000 m.; Zacapa; Sacatepe"- 
quez; Chimaltenango; Suchitepequez; Quezaltenango; San Marcos. 
El Salvador. Figure 18. 

Herbaceous; rhizome branching, 3-5 mm. thick, the younger parts rufous- 
hirsute, internodes evident, up to 3 cm. long; stipules lanceolate acuminate 
with a long soft bristle at apex, 7-12 mm. long, membranaceous, ferrugineous, 
subglabrous; petioles very slender, erect, 7-18 cm. long, sparsely rufous-hirsute 
with slender septate trichomes 2 mm. long; leaf-blades peltate, ca 7-nerved, 
broadly ovate or elliptic, rather abruptly acuminate, broadly rounded at base, 
5 5-10 cm long, 3-6 cm. wide, sometimes very slightly lobed, entire or obscurely 
denticulate, green and glabrous above, paler beneath and rufous-hirsute especially 
on the nerves; peduncles nearly 2 dm. long, equaling or exceeding the leaves; 
inflorescence cymose, few-flowered, bisexual, bracts deciduous, unknown pedicels 
15-20 mm. long, glabrous or hirsute; staminate tepals 2, 6-10 mm. long, 7-1 d mm. 
wide, suborbicular, cordate at base, entire, subfleshy, white, obscurely nerved; 
stamens numerous, anthers oblong, filaments short; pistillate flowers ebracteolate, 
tepals 2, suborbicular, 9-11.5 mm. long, 11-12 mm. wide; ovary slightly hirsute, 
styles 3, 2-parted, placentae bilamellate, capsule decurved, subelhpsoid, t 
single wing falcate, obtuse, 15-20 mm. long, 5-6 mm. wide. 

Begonia caroliniifolia Regel, Gartenfl. 1: 259, t. 25. 1852. 
B. rotata Liebm. Vid. Medd. Kjoebenhavn 1852: 11. 1853. Gireoudia 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 163 

caroliniaefolia Klotzsch, Monatsber. Akad. Berlin 125. 1854 G ro- 
tate Klotzsch, Begon. 91. 1855. 

Epiphytic and on boulders, 100-1,200 meters; Izabal; Huehuete- 
nango. Southern Mexico. 

Herbaceous; rhizome erect or prostrate, 3-6 dm. long, fleshy, 2 cm. or more 
thick, knobby, white-tuberculate, setose when young but later glabrous, internodes 
very short; stipules deciduous, ovate, acuminate, entire, ferruginous- villous; petioles 
erect, stout, 15-60 cm. long, rufous-pilose when young but soon glabrous; leaf- 
blade palmately compound, leaflets petiolulate for 1-5 cm., asymmetric, simple 
or with one lateral lobe toward apex, linear-lanceolate, cuneate, acuminate, 1-3 
dm. long, 1-7 cm. wide, thin, rufous-villous beneath when young, margin undulate 
setose; peduncle exceeding the leaves, to 85 cm. long, stout, soon glabrous; in- 
florescence irregularly cymose with one primary branch tending to elongate and 
make the other appear lateral, many-flowered, to 14 cm. long, lax, soon glabrous 
bisexual; bracts deciduous, elliptic, obtuse, 8-12 mm. long, pilose, entire; pedicels 
slender, 5-18 mm. long; staminate tepals 2, suborbicular, 6-8 mm. long, pink or 
white; stamens rather numerous, filaments shorter than the oblong anthers 
connective produced, short, blunt; pistillate flowers ebracteolate, tepals like the 
staminate; ovary 3-celled, placentae bifid, ovuliferous on all sides, styles tardily 
deciduous, nearly free, the stigmatic surface lunate-capitate at their apices- 
capsule more or less nodding but not sharply reflexed, suborbicular, 8-10 mm long' 
wings very unequal, the largest 20 mm. wide, 15 mm. high, nearly truncate above' 
rounded below; seeds ellipsoid, blunt. 

Begonia cebadillensis Houghton ex Smith & Schubert, Contr 
Gray Herb. 161: 26, t. 3. 1946. 

Wet, shady quebrada, 900-950 meters; Santa Rosa (Standley 
78204). El Salvador. 

Erect caulescent herb; stipules deciduous, lanceolate, acuminate ciliate- 
lacerate, 2 mm. long, petioles slender, to 7 cm. long, leaves strongly oblique, 
subelhptic, acuminate, sublobate, shallowly cordate at base, to 4 cm. long and 9 cm 
wide, dentate, sparsely puberulent on both sides; peduncles axillary, much shorter 
than the leaves; inflorescence 1-few-flowered; bracts deciduous, like the stipules- 
pedicels 6-23 mm. long, sparsely glandular; staminate tepals 4, entire rose the 
outer broadly reniform, to 10 mm. long, about twice as long as the stamens' the 
inner narrowly elliptic, about equaling the stamens; stamens many on a short 
column, anthers obovoid, much shorter than the filaments; pistillate tepals un- 
known; styles persistent, articulate, short-bilobate, placentae bilamellate ovulif- 
erous throughout; capsule oblong-ellipsoid, unequally 3-winged, the wings tri- 
angular. 

Begonia confusa Smith & Schubert, Contr. Gray Herb 161- 
27, t. 3. 1946. 

Creeping on rocks, 1,200-2,000 meters; Huehuetenango (type 
from Sierra de los Cuchumatanes, Sieyermark 49185). 



164 FIELDIANA: BOTANY, VOLUME 24 

Herbaceous; rhizome slender, prostrate, red, glabrous, internodes 1-9 cm. 
long; stipules deciduous, lance-oblong, entire, 23 mm. long, pilose, petioles erect, 
16-23 cm. long, sparsely pilose, blades straight, asymmetric, broadly ovate, 
acuminate, 1-3-lobed, cordate at base, to 18 cm. long, entire or sparsely denticulate, 
sparsely puberulent on both sides, more densely on the nerves beneath; young 
peduncles much shorter than the leaves; cymes few-flowered, regular, dense when 
young; bracts deciduous, elliptic, entire, membranaceous, the lowest 12 mm. long; 
pedicels about 10 mm. long; staminate tepals 2, broadly ovate, obtuse, entire, 
red-puberulent, 12 mm. long; stamens numerous, anthers oblong, longer than 
the filaments, connective produced, obtuse; pistillate flowers probably ebracteolate, 
only very young ones known; tepals 3, the outer ones broadly ovate, fleshy, red- 
puberulent, the inner one much smaller, elliptic, glabrous; styles 3, bifid; ovary 
unequally 3-winged, pubescent, the wings ascending. 

Begonia convallariodora C. DC. Bot. Gaz. 20: 538. 1895; 
Smith & Schubert, Ann. Mo. Bot. Card. 45: 67, fig. 20. 1958. 

Thickets or moist forest, sometimes epiphytic (Panama), 1,250- 
1,350 meters; Alta Verapaz (type from Pansamala, von Tuerckheim 
886); Huehuetenango. Southern Mexico to Panama. Figure 19. 

Scandent to erect herb 1-4 m. high with little-branched slender stem, glabrous; 
stem ribbed, internodes of main axis 5.5-14 cm. long; stipules somewhat persistent, 
obliquely elliptic, membranaceous, base truncate and apex acute and apiculate, 
entire, 25 mm. long, 7 mm. wide; petioles 12-40 mm. long; leaf-blades asymmetric, 
obliquely elliptic, pinnately veined, abruptly acuminate, base usually with broader 
half produced into a conspicuous lobe, 8.5-15 cm. long, 4-7.5 cm. wide, more or 
less denticulate; peduncles 3-8 cm. long; cymes many-flowered, regular; bracts 
very early deciduous; pedicels 6-16 mm. long; staminate tepals entire, white or 
pinkish, 2 or less often 3 or 4, the outer orbicular, cordate at base, 4-8 mm. long, 
the inner elliptic, 1-3.5 mm. long; stamens many, anthers oblong-elliptic, longer 
than the filaments, connective produced, obtuse; pistillate bracteoles early decidu- 
ous, elliptical, entire, acute; tepals entire, 2 or less often 3, the outer orbicular to 
rounded ovate, 6-9 mm. long, the third much smaller; styles 3, bifid; placentae 
bilamellate, ovuliferous throughout; capsule ellipsoid, unequally 3-winged, the 
largest wing obtuse, 7-12 mm. wide. 

The flowers are said to smell like Convallaria, hence the name. 

Begonia crassicaulis Lindl. Bot. Reg. 28: misc. 22, t. 44- 1842. 

Thickets or forest, sometimes epiphytic, 1,300-1,400 m. Endemic. 
Figure 20. 

Herbaceous; rhizome short, simple or with very short bulbous branches, 3-4 
cm. thick, pilose, soon glabrous, internodes very short; leaves developing after 
the flowers, stipules triangular, acuminate, 15-25 mm. long, entire, brown, thin, 
petioles erect, elongate, fuscous-lanate at least when young, leaf-blades palmately 
5-7-lobed about half-way to base, 12-22 cm. wide, sinuate-dentate with acute 
or acuminate lobes, fuscous-lanate beneath at first, peduncles erect, 7-30 cm. long, 
fuscous-lanate; inflorescence unilaterally cymose so that the larger side appears 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 



165 




FIG. 19. Begonia convallariodora. A, Branch; X %. B, Staminate flower; 
X 1. C, Stamen; X 5. D, Pistillate flower; X 1. E, Style; X 5. F, Fruit; X 1. 



to be a continuation of the peduncle; bracts quickly deciduous, broadly elliptic, 
obtuse, 10 mm. long, serrulate; pedicels slender, 6-11 mm. long; staminate tepals 2, 
suborbicular, 6-10 mm. long, white or rose; stamens numerous, anthers oblong; 
pistillate flowers with 2 large ample deciduous bracts at base; tepals 2, like the 
staminate; ovary 3-celled, placentae bifid, ovuliferous on all sides; styles connate 
at base, stigmatic papillae lunate-capitate at their apices; capsule erect or some- 
what nutant, ellipsoid, 12-16 mm. long, wings slightly unequal, the largest ovate, 
subacute, 8 mm. wide, seeds ellipsoid, blunt. 

Begonia Fischer! Schrank var. tovarensis (Klotzsch) Irmscher, 
Bot. Jahrb. 76: 23. 1953; Smith & Schubert, Ann. Mo. Bot. Card. 
45: 57, /. 17. 1958. B. tovarensis Klotzsch, Begon. 31. 1855. B. Mo- 
ritziana Klotzsch, I.e., not Kunth & Bouch 1848. 

Swamps, 50-1,700 meters; Pete"n; Alta Verapaz; Baja Verapaz; 
Izabal; Santa Rosa; Sacatepe*quez ; Chimaltenango. Southern Mex- 
ico and Cuba to Venezuela, Bolivia and Peru. 



166 



FIELDIANA: BOTANY, VOLUME 24 





H 




FIG. 20. Begonia crassicaulis. A, Habit; X %. B, Leaf; X Y^,. C, Stami- 
nate flower and bracts; XI. D, Staminate flower opened; XI. E, Stamen; X 5. 
F, Pistillate flower and bracts; X 1. G, Style; X 5. H, Fruit; X 1. 

Herb, 2-12 dm. high, sparsely brown-pilose to glabrous; stem erect, red; 
stipules deciduous, ovate-oblong, to 10 mm. long, ciliate-serrulate, petioles 5-60 
mm. long, leaf-blades more or less asymmetric, broadly ovate or suborbicular, 
acute or rounded, cordate at base, 2-9 cm. long, palminerved, crenate-serrate, 
ciliate, nearly or quite glabrous above, sparsely hirsute beneath; peduncles axillary, 
15-45 mm. long; cymes few-flowered; bracts persistent, ovate, 2-4 mm. long, 
fimbriate; pedicels 5-17 mm. long; staminate tepals 4, the outer orbicular, 8 mm. 
long, the inner smaller, narrowly obovate; stamens free, numerous, filaments short, 
anthers oblong; pistillate flowers bracteolate, tepals 5, obovate, 3-6 mm. long; 
styles 3, 2-parted, the stigmatic tissue linear, spiral, continuous, placentae bila- 
mellate, ovuliferous on all sides; capsule ellipsoid, 10-15 mm. long, its wings 
decurrent, very unequal, the largest usually ascending, tapering and often hooked, 
to 23 mm. wide, seeds curved with attenuate apices. 

Begonia Franconis Liebm. Vid. Medd. Kjoebenhavn 1852: 21. 
1853. B. modesta Liebm. I.e. 20. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 167 





FIG. 21. Begonia Franconis. A, Partial habit; X K- B, Staminate flower; 
X 1. C, Stamen; X 5. D, Pistillate flower; X 1. E, Style; X 5. F, Fruit; X 1. 



On rocks, 1,100 meters; Alta Verapaz. Southern Mexico. Fig- 
ure 21. 

Herbaceous annual, caulescent, branching, 5-50 cm. high, stem with mixed 
coarse and fine pubescence, erect or decumbent at base; stipules persistent, lance- 
olate, acute, 2-3 mm. long, ciliate, petioles slender, 5-70 mm. long, leaf-blades 
obliquely ovate, acute, cordate to subtruncate at base, 3.5-8 cm. long, crenate- 
dentate, ciliate, thin; peduncles axillary, 1-few-flowered, to 3 cm. long; bracts per- 
sistent, ovate, acute, 1.5-2 mm. long, lacerate; pedicels 4-10 mm. long, filiform; 
staminate tepals 4, entire, rose or white, the outer broadly ovate or orbicular, 
3.5-7 mm. long, the inner elliptic, shorter; stamens few on a column, anthers ellip- 
soid, shorter than the filaments; pistillate bracteoles subulate, minute; pistillate 
tepals 5, subequal, elliptic; styles 3, bilobed, placentae simple; capsule ovoid or 
ellipsoid, with the lunate, subequal wings broadest at the top, middle or base and 
much extended above the locules; seeds obtuse. 

Begonia fusca Liebm. Vid. Medd. Kjoebenhavn 1852: 7. 1853. 
B. maxima Klotzsch, Begon. 39. 1855, nomen, in synon. Magnusia 
fusca (Liebm.) Klotzsch, Begon. 102. 1855, and M . fusca Klotzsch, 
I.e. 103, t. 9, C, figs. a-m. M. maxima Klotzsch, I.e. Begonia John- 
stonii Standl. ex J. R. Johnston, Cat. Plant. Guatemala 12. 1938, 
nomen (type from Alta Verapaz, Johnston 1152). Flor de mano de 
leon de penasco (fide Steyermark). 

Epiphytic or terrestrial on shady slopes, 1,300-2,500 meters; 
Alta Verapaz; Baja Verapaz; Zacapa; Huehuetenango; Quezalte- 
nango; San Marcos. Southern Mexico. 

Herbaceous, to 15 dm. high (Steyermark!); rhizome procumbent or oblique, 
15-40 mm. thick, setose (Klotzsch!), with very short internodes or sometimes 



168 FIELDIANA: BOTANY, VOLUME 24 

only about 5 mm. thick and with distinct internodes; stipules deciduous, broadly 
ovate, subacuminate, pilose at first, serrate-setose (Klotzsch!) or entire; petioles 
erect, 2-4 dm. long, densely retrorse-pilosulous; blades oblique, very broadly 
ovate or suborbicular, abruptly acuminate, often with several superficial acuminate 
lobes beside the apex, cordate, up to 6 dm. long (Standley!), palmately 7-9-nerved, 
angulate-dentate, ciliate-serrate, thin, finely pubescent on both sides; peduncle 
exceeding the leaves, to 6 dm. long, 5-10 mm. thick, pilosulous at first; cyme 
regular, much branched, 2-5 dm. long, pubescent; bracts deciduous, suborbicular, 
pubescent, the upper ones entire, thin, the lower ones over 4 cm. long, sometimes 
serrulate; pedicels slender, 1-4 cm. long, pubescent; staminate tepals 2, broadly 
ovate or suborbicular, subacute or obtuse, 6-11 mm. long, fleshy, sparsely pilose, 
pink or white; stamens numerous, filaments shorter than the oblong anthers, 
connective produced into a broad flat cap above the anther; pistillate tepals 2 or 
sometimes a third and much smaller one inside the others, like the staminate but 
much smaller; ovary 3-celled, placentae bifid, ovuliferous throughout, styles decidu- 
ous, slightly connate at base, bifid, the stigmatic surface linear, spiral, continuous; 
capsule more or less nutant, broadly ovoid, pubescent at first, 8-12 mm. long, 
wings very unequal, the largest dolabriform or subovate, 25-30 mm. broad. 

Begonia glabra Aubl. PI. Guian. 2: 916, t. 349. 1775; Smith & 
Schubert, Ann. Mo. Bot. Card. 45: 54, /. 17. 1958. B. scandens Sw. 
Prodr. 86. 1788. B. elliptica HBK. Nov. Gen. & Sp. 7: 180, t. 6^1. 
1825. B. lucida Otto & Dietr. Allg. Gartenz. 16: 162. 1848. B. Mo- 
ritziana Kunth & Bouche", Ind. Sem. Hort. Berol. 16. 1848. B. phy- 
salifolia Liebm. Vid. Medd. Kjoebenhavn 1852: 19. 1853. Wageneria 
glabra Klotzsch, W. Moritziana Klotzsch and W. lucida Klotzsch, 
Monatsber. Akad. Berlin 126. 1854. W. deflexa Klotzsch, Begon. 
113. 1855. W. montana Klotzsch, I.e. 115. Begonia locellata A. DC. 
Ann. Sci. Nat. se>. 4. 11: 137. 1859. 

Epiphytic and terrestrial in forest, 75-1,300 meters; Pete"n; Alta 
Verapaz; Izabal; Suchitepe"quez; Huehuetenango; Quezaltenango; 
San Marcos. Southern Mexico and the West Indies to Peru and 
Bolivia. 

Stem scandent, rooting at the nodes, glabrous; stipules persistent, ovate- 
oblong, setiferous at apex, entire, 10-22 mm. long, petioles 1-8 cm. long, leaf- 
blades variable, straight, nearly symmetrical, broadly ovate, 4-12 cm. long, 
short-acuminate, rounded or barely cordate at base, sparsely serrate and ciliate 
to entire, glabrous; peduncles axillary, 6-12 cm. long; cymes unisexual, many- 
flowered, diffuse; bracts persistent, minute; pedicels 6-17 mm. long, fine; staminate 
tepals 4, entire, white, the outer broadly obovate, 3-6 mm. long, the inner narrower; 
stamens free, few, anthers oblong; pistillate flowers minutely bracteolate; pistillate 
tepals 5, elliptic, 4-6 mm. long; styles 3, 2-parted, completely covered by stigmatic 
papillae, placentae simple; capsule erect, ellipsoid, 6-12 mm. long, the largest 
wing oblong to deltoid, spreading or barely ascending, 10-23 mm. wide, the other 
two marginiform, very narrow. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 



169 




FIG. 22. Begonia gradlis. A, Partial habit; X H. B, Staminate flower; X 1. 
C, Stamen; X 5. D, Pistillate flower and bracts; X 1. E, Fruit; X 1. 



Begonia gracilis HBK. Nov. Gen. & Sp. 7: 184. 1825. 

Solola (slopes above San Lucas Tollman, Volcan Tollman, Steyer- 
mark 47605). Mexico, where It grows in moist forest or sometimes 
on exposed rocks, at 1,500-2,500 meters. Figure 22. 

Herbaceous, erect-caulescent from a tuberous base, very variable; stem 
pubescent or glabrous, often bearing bulblets in the leaf-axils; stipules persistent, 
green, oblong, acute, serrulate-ciliate; petioles slender, 1-10 cm. long but the 
upper ones short; leaves oblique, ovate or elliptic and acute or acuminate, the 
lowest sometimes reniform, cordate at base, more or less lobed, doubly serrate, 
3-12 cm. long, pubescent or glabrous; peduncles 1- (rarely 2-) flowered; bracts 
persistent, broadly elliptic, serrate; pedicels to 3 cm. long; staminate tepals 4, 
pink, glabrous, the outer broadly ovate or elliptic, more or less serrate or entire, 
10-22 mm. long, the inner narrowly obovate, shorter, entire; stamens many, 
on a short column, anthers obovoid, much shorter than the filaments; pistillate 
flowers ebracteolate; pistillate tepals 5, like the staminate; styles bifid, placentae 
bilamellate; capsule ellipsoid, unequally 3- winged, the largest wing triangular, 
the others ovate to lunate. 



170 



FIELDIANA: BOTANY, VOLUME 24 






FIG. 23. Begonia heracleifolia. A, Habit; X ^. B, Staminate flower and 
bracts; X 1. C, Stamen; X 5. D, Pistillate flower and bracts; X 1. E, Style; 
X 5. F, Fruit; X 1. 

Begonia heracleifolia Schlecht. & Cham. Linnaea 5: 603. 1830. 
B. radiata R. Graham, in Edinb. Phil. Journ. 182. July 1833. B. punc- 
tata Link & Otto, Ic. PL Rar. 1: 16, t. 7. 1840-44. Gireoudia hera- 
cleifolia Klotzsch and G. punctata Klotzsch, Monatsber. Akad. Berlin 
125. 1854. G. heracleifolia var. a viridis Klotzsch and var. ft punctata 
Klotzsch, Begon. 95. 1855. Begonia nigricans Klotzsch, Begon. I.e., 
nomen, in synon. B. heracleifolia var. nigricans Hook. Bot. Mag. 83: 
t. 4983. 1857. B. longipila Lem. 111. Hortic. 8: t. 307. 1861. B. hera- 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 171 

cleifolia var. /3 nigricans A. DC., var. y longipila A. DC. and var. 5 
punctata A. DC. Prodr. 15, pt. 1: 335. 1864. B. heracleifolia var. 
Sunderbrucki Hort. ex C. Chevalier, Begon. 268. 1938. Tonichut 
(Quecchi). 

Terrestrial and saxicolous in forests, 50-1,500 meters; Pete"n; Alta 
Verapaz; Zacapa; Santa Rosa; Guatemala. Southern Mexico, British 
Honduras. Figure 23. 

Herbaceous, very variable in indument, coloration of leaves, size of inflo- 
rescence and form of capsule- wings; rhizome repent, 7-15 cm. long, 2 cm. thick, 
simple, setose, internodes very short; leaves developing with the flowers, stipules 
triangular, setose-acuminate, entire, 12-20 mm. long, petioles erect, stout, 3-40 
cm. long, from densely hirsute with slender sometimes divided trichomes to nearly 
glabrous; trichomes especially large and dense at the apex of the petiole, leaf- 
blade palmately 7-lobed ^ to % toward base, subcordate, 8-26 cm. wide, irregu- 
larly sinuate-dentate with acuminate lobes, nearly or quite glabrous above, more 
or less hirsute beneath on the nerves and margin, thin, green throughout or with 
broad, nearly black margins; peduncles mostly exceeding the leaves, 2-7 dm. 
long, more or less hirsute; inflorescence laxly cymose but tending to develop one 
side much more than the other, bisexual, up to 25 cm. long, many-flowered, some- 
times red-punctate; bracts persistent, broadly ovate or elliptic, entire or serrate, 
green or red, the lowest 2 cm. long; pedicels slender, 8-18 mm. long; staminate 
tepals 2, suborbicular, 10-14 mm. long, white or rose; stamens numerous, subfree, 
filaments short, anthers oblong, the connective produced into a broad rounded 
appendage; pistillate flowers ebracteolate; pistillate tepals 2, like the staminate; 
ovary 3-celled, placentae bifid, ovuliferous on all sides, styles connate at base, 
the stigmatic tissue lunate at their apices; capsule erect or somewhat nutant, 
suborbicular, 8-12 mm. long, wings unequal, the largest ovate with the upper side 
nearly horizontal, usually subacute, 8-12 mm. wide, seeds ellipsoid, blunt. 

Standley says of this species: "A common ornamental plant in 
Guatemala, in pots and gardens. Wild at many places in the coun- 
try. The finest display of it that I have ever seen was in Alta Vera- 
paz on the road above Pancajche, where for long distances it covered 
completely the roadside banks. The display of beautiful pink flowers 
equaled almost the best that could be produced at horticultural ex- 
hibitions in the United States, and gave the effect almost of well- 
furnished greenhouse benches." 

Begonia Heydei C. DC. Bot. Gaz. 20: 540. 1895 (type from 
Quiche", Heyde & Lux 3094); Smith & Schubert, Ann. Mo. Bot. Gard. 
45: 62, /. 18. 1958. B. Pittieri C. DC. Bull. Herb. Boiss. II. 8: 316. 
1908. B. triloba C. DC. I.e. 322 (type from Alta Verapaz, von Tuerck- 
heim II1954). 

Usually epiphytic in forests, 900-2,100 meters; Alta Verapaz; 
Quiche"; Huehuetenango. Costa Rica; Panama. Figure 24. 



172 



FIELDIANA: BOTANY, VOLUME 24 




FIG. 24. Begonia Heydei. A, Branch; X K- B, Staminate flower; X 1. 
C, Stamens; X 5. D, Pistillate flower and bracts; X 1. E, Style; X 5. F, Fruit; 
X 1. 



Erect, to 6 dm. high, branching, glabrous, internodes elongate; stipules decidu- 
ous, lance-ovate, subulate, 5 mm. long, petioles 10-34 mm. long, leaf-blades 
oblique, cordate-ovate, acuminate, 6-10 cm. long, 2-3.5 cm. wide, remotely denticu- 
late; peduncles axillary, 15-28 mm. long; cymes few-flowered, exceeded by the 
leaves; bracts deciduous; pedicels 8-10 mm. long; staminate tepals 4, entire, 
obtuse, the outer ovate or elliptic, 7 mm. long, setose toward base, the inner 
slightly shorter; stamens many on a short column, anthers obovate, nearly equaling 
the filaments; pistillate bracteoles elliptic, entire, about equaling the ovary; 
pistillate tepals 3, elliptic or obovate, entire, setose toward base; ovary 3-celled, 
placentae simple; styles deciduous, connate at base, bifid; capsule rhomboidal, 
attenuate at base, bearing three equal slender acuminate horns 9-11 mm. long. 

Begonia ignea (Klotzsch) Warsc. ex A. DC. Prodr. 15, pt. 1: 
306. 1864 (type from cultivated material from Guatemala, without 
further locality, Warsczewicz) . Knesebeckia ignea Klotzsch, Begon. 
46. 1855. Costa Rica. 

Caulescent erect herb, 6-9 dm. high, stem glandular-hispid, base subtuberous; 
stipules persistent, suborbicular, cordate, 6-12 mm. long, serrulate, green, petioles 
to 12 cm. long, leaf-blades oblique, broadly ovate, sharply 4-6-lobed, cordate at 
base, 5-14 cm. long, 7-22 cm. wide, serrate, hispid; peduncles axillary, 4-9 cm. 
long; cymes few-flowered, dichotomous, bracts deciduous, broadly ovate, obtuse, 
4-6 mm. long, ciliate, rose; pedicels pilose; staminate tepals 4, rose, the outer ovate, 
acute, 16-20 mm. long, ciliate-serrulate, pilose, the inner shorter, obovate, obtuse, 
entire, glabrous; stamens numerous, on a column, anthers obovoid, much shorter 
than the filaments; pistillate bracteoles like the bracts; pistillate tepals 5, like the 
staminate; styles 3, persistent, irregularly short-branched; capsule ellipsoid, pilose, 
unequally 3- winged (!A. DC.), the largest wing obtuse. 



STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 173 

Begonia involucrata Liebm. Vid. Medd. Kjoebenhavn 1852: 
15. 1853; Smith & Schubert, Ann. Mo. Bot. Card. 45: 60, /. 18. 1958. 
B. broussonetiaefolia A. DC. Ann. Sci. Nat. se>. 4. 11: 133. 1859 (type 
from Guatemala, Friedrichsthal) . B. laciniosa A. DC. Prodr. 15, 
pt. 1:340. 1864. 

Known here only from the type of B. broussonetiaefolia, and with- 
out further locality than "Guatemala." Nicaragua; Costa Rica; 
Panama. 

Erect, 1-2 m. high, fruticose, stem fulvous-tomentose or becoming glabrous; 
stipules deciduous, ovate, setose-apiculate, 14-25 mm. long, entire, brown, mem- 
branaceous, glabrous or sparsely pilose, petiole 12-18 cm. long, fulvous-tomentose 
or becoming glabrous; leaf -blades oblique, cordate at base, 2-5-lobed at apex 
with acuminate points, 15-28 cm. wide, to 20 cm. high, denticulate, evenly pilosul- 
ous above, more densely on the nerves beneath; peduncles axillary, exceeding the 
leaves, fulvous-tomentose, becoming glabrous; cymes many-flowered, dense, their 
primary branches short and their ultimate branches so nearly aborted that the 
inflorescence appears almost umbellate; bracts deciduous, the outer ovate, ample 
and completely enclosing the young inflorescence, pedicels 5-25 mm. long; staminate 
tepals 2, suborbicular, 6-11 mm. long, entire, sparsely pilose to glabrous, white; 
stamens numerous, anthers oblong, about equaling the filaments, connective 
produced, broadly obtuse; pistillate bracteoles lacking; pistillate tepals 2, like 
the staminate; ovary 3-celled, placentae bilamellate; styles connate at base, bifid; 
capsule ellipsoid, very unequally 3-winged, the largest wing elliptic or falcate, 
obtuse, 8-10 mm. wide. 

Begonia Lindleyana Walp. Rep. 2: 209. 1843; Smith & Schu- 
bert, Caldasia 4, no. 16: 11, t. 2. 1946. B. vitifolia Lindl. Bot. Reg. 
28: misc. 21. 1842, not Schott 1827. B. sarchophylla Liebm. Vid. 
Medd. Kjoebenhavn 1852: 12. 1853. B. sericoneura Liebm. and 
B. cardiocarpa Liebm. I.e. 13. Gireoudia cardiocarpa Klotzsch, Mon- 
atsber. Akad. Berlin 125. 1854. G. fibrillosa Klotzsch and G. pilifera 
Klotzsch, Begon. 86. 1855. G. vitifolia Klotzsch, I.e. 87. G. sarcho- 
phylla Klotzsch, I.e. 88. G. sericoneura Klotzsch, I.e. 89. Begonia 
lanuginosa A. DC. Ann. Sci. Nat. se*r. 4. 11: 131. 1859. B. pilifera 
A. DC. Prodr. 15, pt. 1: 337. 1864. B. Biolleyi C. DC. Bull. Soc. 
Bot. Belg. 35, pt. 1: 263. 1896. B. nicaraguensis Standl. Field Mus. 
Bot. 4: 237. 1929. Flor de piedra (fide Steyermark); choo-kek (fide 
Steyermark) . 

Terrestrial and saxicolous, 100-1,150 meters; Pete"n; Alta Vera- 
paz; Izabal; Huehuetenango. Southern Mexico to Colombia. 

Herbaceous, fuscous-villous throughout, at least when young; rhizome usually 
erect, up to 3 dm. long, subligneous at base, 10-25 mm. thick, internodes very 
short; stipules persistent, imbricate, subtriangular, acuminate-setiferous, 15-25 
mm. long, reticulate-veined, entire; petioles erect, 3-22 cm. long, 2-5 mm. thick; 



174 FIELDI ANA: BOTANY, VOLUME 24 

leaves palmately 7-9-nerved, oblique, very broadly ovate, abruptly acuminate, 
deeply cordate at base, from almost evenly rounded and entire to deeply acumi- 
nate-lobed and coarsely dentate, 8-22 cm. long, thin, soon glabrous above; peduncles 
exceeding the leaves, up to 48 cm. long; cymes usually subregular, diffuse and 
broader than high, few- to many-flowered; bracts deciduous, very broad, obtuse, 
the lowest 17-20 mm. long, serrate, ciliate, very thin; staminate tepals 2 or 4, 
the outer ones suborbicular, 6-13 mm. long, entire, white, the inner when present 
smaller and narrowly obovate; stamens numerous, filaments short, anthers oblong, 
obtuse; pistillate flowers bracteolate; pistillate tepals 2 or sometimes a smaller one 
inside the others, suborbicular, smaller than the staminate; ovary 3-celled, placentae 
bifid, ovuliferous throughout, styles short-connate, the stigmatic tissue lunate- 
capitate at their apices; capsule suberect, ellipsoid, 8-12 mm. long, wings unequal, 
the largest ovate, obtuse or acute, subascending, as high as the capsule, 11-15 mm. 
wide. 

Begonia ludicra A. DC. Ann. Sci. Nat. se>. 4. 11: 133. 1859. 
B. reptans Liebm. Vid. Medd. Kjoebenhavn 1852: 5. 1853, not Benth. 
Weilbachia reptans Klotzsch & Oerst. Begon. 120, 1. 11, fig. A. 1855. 
Begonia Liebmanni A. DC. Prodr. 15, pt. 1: 345. 1864. 

Epiphytic or terrestrial in wet forest; 1,410-2,250 meters; Alta 
Verapaz; Quezaltenango; San Marcos. Southern Mexico; Panama. 

Herb, stem repent or erect, to 1 m. long, slender, with elongate internodes, 
soon glabrous; stipules mostly persistent, broadly ovate, 7-15 mm. long, entire, 
petioles erect, 6-30 cm. long, slender, pubescent or glabrous; blades extremely 
variable, oblique, ovate to subpalmate, acuminate, with one or more lobes barely 
developed to elongate and slender, the base from typically deeply cordate with 
overlapping lobes through subtruncate or broadly rounded to cuneate with a 
minute cordate termination, entire or sparsely dentate, 7-17 cm. long, glabrous 
or somewhat pubescent on the nerves beneath; peduncles 6-30 cm. long; cyme 
few-flowered, dense; bracts deciduous, ample, the lowest 12 mm. long; pedicels 
slender, 10-20 mm. long; staminate tepals 2 or 4, the outer suborbicular, fleshy, 
10-18 mm. long, the inner narrowly obovate; stamens numerous, the filaments 
much shorter than the oblong anthers; pistillate flowers ebracteolate; pistillate 
tepals 2, like the staminate; ovary 3-celled, styles much connate; capsule nutant 
to very sharply reflexed, asymmetric, subellipsoid, 13-14 mm. long, wings more or 
less decurrent from the apex of the capsule, very unequal, the largest ovate, 
obtuse, 13 mm. long. 

Begonia manicata Brongn. ex Gels. Journ. Jard. (Ann. Fl. et 
Pom.) 104, 256. 1842; Gels, ex Vis. Orto Padov. 135. 1842; Mart. Fl. 
Bras. 4, pt. 1: t. 101. 1861. B. schizolepis Liebm. and B. lepidota 
Liebm. Vid. Medd. Kjoebenhavn 1852: 17. 1853. Gireoudia mani- 
cata Klotzsch, Monatsber. Akad. Berlin 125. 1854. 

Rocks or rich slopes, 500-1,500 meters; Alta Verapaz; Zacapa; 
Chiquimula; San Marcos. Southern Mexico. 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 175 

Herbaceous; rhizome ascending, 5-15 cm. long, fleshy, internodes short but 
usually distinct; stipules persistent, lanceolate, setose-acuminate, 12-19 mm. long; 
petioles rather stout, up to 28 cm. long, bearing at apex a whorl of large bright red 
fimbriate scales and usually more below at intervals, often scaly clear to the 
base but then the whorls successively more irregular until broken up into tufts 
of scales; leaves broadly oblique-ovate, acute to acuminate, cordate at base, 
angulate-dentate and denticulate, dark-ciliate, up to 25 cm. long, green often 
with nerves red at base, glabrous above, beneath with coarse fimbriate scales on 
the main nerves; peduncles exceeding the leaves, bracts quickly deciduous, 6-17 
mm. long; cyme bisexual, much branched, usually strongly unilateral, pedicels 
very slender, 2-15 mm. long; staminate tepals 2, obovate or suborbicular, to 8 mm. 
long, whitish; stamens few, the oblong anthers mostly longer than the filaments; 
pistillate flowers bearing minute quickly deciduous bracteoles; pistillate tepals 2, 
to 7 mm. long; ovary 3-celled, placentae bifid, ovuliferous on all sides, styles 
connate at base, the stigmatic tissue lunate-capitate; capsule ellipsoid, 8-12 mm. 
long, the wings forming an almost even ring about 5 mm. wide all around it and 
acute or obtuse at base, subequal. 

Begonia manicata Brongn. var. peltata Smith & Schubert, 
var. nov., differt foliis peltatis. 

Alta Verapaz : crevices in upper part of limestone bluff, Montana 
Yxocubvain, 2^ miles west of Cubilgiiitz, alt. 300-500 m., March 12, 
1942, Steyermark 44980 (Gray, type). 

Begonia militaris Smith & Schubert, Contr. Gray Herb. 154: 
24, t. 2. 1945. (Type and only known collection from Chama, Alta 
Verapaz, alt. 270 m., Johnson 178.) 

Herb, 18-21 cm. high; rhizome repent, scarcely more than 2 mm. thick, more 
or less pilose, internodes short but distinct; stipules persistent, to 1 cm. long, 
narrowly triangular, acuminate-setose, brown; petioles 5-10 cm. long, slender, 
pilose; leaf-blades peltate, broadly ovate or elliptic, abruptly acute, 5-6 cm. long, 
3-4 cm. wide, entire or faintly undulate, glabrous above, sparsely pilose beneath; 
peduncles exceeding the leaves; inflorescence very laxly paniculate, subsecund, 
glabrous; bracts deciduous, small, broadly elliptic; pedicels very slender, to 8 
mm. long; staminate tepals 4, entire, the outer elliptic, 6 mm. long, roseate, the 
inner narrowly obovate; stamens on a short column, anthers obovate, longer 
than the filaments, connective produced, obtuse; pistillate bracteoles elliptic, 
about equaling the ovary; pistillate tepals 5, narrowly elliptic, obtuse, entire, to 
7 mm. long, the outer roseate; styles 3, shallowly bifid; ovary ellipsoid, placentae 
bilamellate; capsule 3- winged, the two smaller wings lunate, the third narrowly 
triangular, spreading, 14-17 mm. wide. 

Begonia nelumbiifolia Schlecht. & Cham. Linnaea 5: 604. 
1830; Smith & Schubert, Ann. Mo. Bot. Gard. 45: 44, /. 11. 1958. 
B. derycxiana Lem. Hortic. Univ. 5: misc. 355. 1844. Gireoudia 
nelumbiifolia Klotzsch, Monatsber. Akad. Berlin 125. 1854. B. cau- 
dilimba C. DC. Smithson. Misc. Coll. 69, no. 12: 9. 1919. 



176 FIELDIANA: BOTANY, VOLUME 24 

Damp thickets and forests, alt. 1-1,650 meters; Alta Verapaz; 
Izabal; Santa Rosa. Southern Mexico to Colombia. 

Herbaceous; rhizome repent, short, 15 mm. thick, setose, internodes extremely 
short; stipules persistent, lanceolate, over 2 cm. long, entire, rather firm, pilose; 
petioles 15-45 cm. long, 8 mm. thick, sparsely rufous-hirsute, soon glabrous; leaf- 
blades peltate, obliquely very broadly ovate or subelliptic, 7-9-nerved, evenly 
rounded except for the abruptly acuminate apex or slightly produced at the ends 
of the nerves, remotely denticulate, soon glabrous, the margin ciliate and some- 
times purple, 17-40 cm. long, 10-28 cm. wide, thin; peduncle usually exceeding the 
leaves, to 66 cm. long, 6 mm. thick, soon glabrous; cymes regular, much branched, 
diffuse, 2-5 dm. broad; bracts deciduous, ovate, obtuse; pedicels slender, 9-22 mm. 
long; staminate tepals 2, suborbicular, 6-8 mm. long, white or pale pink; stamens 
few, anthers narrowly obovate, equaling or longer than the filaments; pistillate 
flowers ebracteolate; pistillate tepals 2, like the staminate; ovary 3-celled, placentae 
bifid, ovuliferous on all sides, styles connate at base, distinctly divided, the stig- 
matic surface linear, spiral, continuous; capsule erect or somewhat nutant, broadly 
ovoid, 6-10 mm. long, wings very unequal, the largest ovate or deltoid, obtuse, 
to 15 mm. wide, seeds ellipsoid, blunt. 

Begonia oaxacana A. DC. Prodr. 15, pt. 1: 312. 1864; Smith & 
Schubert, Ann. Mo. Bot. Card. 45: 58, /. 18. 1958. B. oaxacana 
|8. pilosula A. DC. I.e. B. Luxii C. DC. Bot. Gaz. 20: 541. 1895. 
B. serrulatoala C. DC. Bull. Herb. Boiss. II. 8: 321. 1908. Nitro 
(fide Steyermark). 

Wet forests and thickets, common, 1,600-3,000 meters; Alta 
Verapaz; El Progreso; Zacapa; Sacatepe'quez; Chimaltenango; So- 
lola; Suchitepe"quez; Quiche"; Huehuetenango; Quezaltenango; San 
Marcos. Southern Mexico; El Salvador. 

Terrestrial or epiphytic, herbaceous to almost shrubby, suberect or climbing, 
very variable in all its parts, 3-15 dm. high; stems 2-12 mm. thick, glabrous to 
pilose, green or red; stipules deciduous, oblong, 12 mm. long, setose-mucronate, 
entire; petioles slender, 4-15 cm. long, sparsely hirsute to glabrous; leaf-blades 
transverse or oblique or rarely some almost straight, broadly or narrowly ovate, 
acuminate, palmately 5-7-nerved, cordate at base, slightly or usually not at all 
lobed, 6-18 cm. long, 3-11 cm. wide, serrulate, sparsely hirsute on both sides or 
nearly glabrous above, thin; peduncles axillary, 2-5 cm. long; cymes few-flowered, 
bisexual; bracts tardily deciduous, broadly ovate or suborbicular, obtuse, 6-12 mm. 
long, entire; pedicels 10-15 mm. long; staminate tepals 4, the outer suborbicular 
or broadly ovate, 10-15 mm. long, pink, glabrous or at times somewhat pubescent, 
the inner slightly shorter, obovate, white; stamens free, numerous, filaments longer 
than the obovoid anthers; pistillate bracteoles wanting or quickly deciduous; 
pistillate tepals 3, the outer two suborbicular, 8-14 mm. long, the inner one much 
smaller, narrowly obovate or oblong; ovary 2-3-celled, placentae bifid, ovuliferous 
on all sides, styles 3, short-connate, bilobed, the stigmatic surface linear, spiral, 
continuous; capsule erect, broadly elliptic, 10-15 mm. long, hirsute to glabrous, 
wings 3 or sometimes up to 6, subequal, narrow, angled at middle or somewhat 
above, giving the fruit a rhombic or obovate outline, entire to fimbriate-serrulate, 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 



177 



seeds ellipsoid, broadly obtuse, short-stalked. A very variable species even in 
individual specimens, and given to freaks like the abortion of one ovary-cell or the 
production of extra wings on the capsule. 

Begonia oaxacana A. DC. var. stenoptera (C. DC.) Smith & 
Schubert, comb. nov. Begonia stenoptera C. DC. Smithson. Misc. 
Coll. 69, no. 12: 2. 1919. 




FlG. 25. Begonia peltata. A, Branch; X 1. 
C, Stamen; X 5. D, Style; X 5. E, Fruit; X 1. 



B, Staminate flower; X 1. 



Terrestrial, on swampy ground, 2,400-2,500 meters; El Progreso, 
Jalapa. Costa Rica; Panama. 

Petioles 5-20 cm. long; pedicels 6-22 mm. long; staminate tepals 5 mm. long; 
capsule-wings equal, marginiform. 

Begonia peltata Otto & Dietr. Allg. Gartenz. 9: 58. Feb. 1841. 
B. incana Lindl. Bot. Reg. 27: misc. 39, no. 73. May 1841. Rachia 
peltata (Otto & Dietr.) Klotzsch, Monatsber. Akad. Berlin 124. 1854; 
Begon. 69, t. 6, f. B. 1855. R. incana (Lindl.) Klotzsch, I.e. B. in- 
cana, jS. auriformis A. DC. Prodr. 15, pt. 1: 327. 1864. Begonia 
Kellermanii C. DC. Smithson. Misc. Coll. 69, no. 12: 1. 1919. 

Terrestrial or saxicolous, wet woods or thickets, 100-1,800 meters; 
Alta Verapaz; El Progreso; Zacapa; Escuintla; Sacatepe'quez; Chi- 
maltenango; Suchitepe"quez; Huehuetenango; Quezaltenango. South- 
ern Mexico. Figure 25. 

Stem erect, simple, straight, 1-9 dm. high, 1 cm. thick, woody, internodes 
short; stipules tardily deciduous, broadly lanceolate, entire, subulate at apex, thin, 
brown, villous, 15 mm. long; petioles 3-15 cm. long; leaf-blades peltate, 7-8- 



178 FIELDIANA: BOTANY, VOLUME 24 

nerved, broadly ovate, rounded at base, abruptly acuminate at apex, 6-15 cm. 
long, 4-10 cm. wide, shallowly sinuate-dentate to entire, densely and closely pale- 
lanate on both sides when young, becoming more or less glabrous above with age; 
peduncles terminal and axillary, 5-39 cm. long; inflorescence cymose, few-flowered; 
bracts very early deciduous, narrowly obovate, villous; staminate pedicels 4- 
14 mm. long; staminate tepals 4, white, entire, the outer orbicular, 5-10 mm. long, 
the inner spatulate, smaller; stamens free, anthers longer or shorter than the fila- 
ments, narrowly obovate, obtuse; pistillate flowers ebracteate; pistillate tepals 3, 
the inner one much smaller; ovary 3-celled, placentae 2-parted, ovuliferous on all 
sides; styles persistent, connate at base, bifid, the stigmatic surface forming a con- 
tinuous spiral; capsule ellipsoid, its wings subequal, lunate to subtriangular. 
There is considerable variation even in a single plant. 

Begonia pinetorum A. DC. Ann. Sci. Nat. ser. 4. 11: 131. 1859. 
B. glandulosa A. DC. ex Hook. Bot. Mag. 87: t. 5256. 1861, as to 
plant illustrated, not as to plant originally labelled by A. DC., nomen 
confusum. B. Tuerckheimii C. DC. Bot. Gaz. 20: 542. 1895 (type 
from Sesisp, Alta Verapaz, von Tuerckheim 885) . 

Wet forests and thickets, 1,200-1,350 meters; Alta Verapaz. 
Mexico. 

Rhizome creeping, its internodes very short; stipules persistent, ovate, acute, 
entire, 2 cm. long; petioles to 32 cm. long, fulvous-tomentose; leaf-blades reniform 
or obliquely elliptic and broadly acute at one side, asymmetric, cordate at base, 
to 11 cm. long and 20 cm. wide, palmately 9-nerved, undulate, fulvous-tomentose 
to glabrous beneath, glabrescent above; peduncles to 45 cm. long, tomentose; 
cymes strongly unilateral, many-flowered; bracts deciduous, oblong, entire, small; 
pedicels 4-20 mm. long; staminate tepals 2, elliptic, obtuse, entire, 4-7 mm. long, 
pilose; stamens few, anthers obovoid; pistillate bracteoles lacking; pistillate tepals 
4, subequal, 2-7 mm. long; styles 3, persistent, lunate-bilobed, placentae bilamel- 
late; capsule ellipsoid, 3- winged, the wings rounded, in the same inflorescence 
sometimes one wing much larger than the other two, sometimes the two larger 
ones subequal. 

Begonia plebeja Liebm. Vid. Medd. Kjoebenhavn 1852: 8. 
1853; Smith & Schubert, Ann. Mo. Bot. Gard. 45: 46, /. 12. 1958. 
B. glandulosa sensu J. D. Smith, Enum. PL Guat. etc. 4: 182. 1895; 
5: 34. 1899. B. uvana C. DC. Smithson. Misc. Coll. 69, no. 12: 4. 
1919. Juego, Fuego, Pie de paloma (fide Standley), Agria (fide 
Steyermark) . 

Damp woods and thickets, moist rocky slopes, 200-1,650 meters; 
Chiquimula; Jalapa; Jutiapa; Santa Rosa; Guatemala; Sacatepe"- 
quez; Suchitepe"quez. Southern Mexico; Central America. 

Herbaceous, fuscous-villous when young; rhizome usually repent, to 8 cm. long 
at least, 8-10 mm. thick, internodes very short; stipules persistent, imbricate, 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 



179 



lanceolate, acuminate-setiferous, 1-2 cm. long, entire, heavily and closely reticu- 
late-nerved; petioles erect, 6-28 cm. long, usually soon glabrous; leaf-blades 
oblique, broadly ovate or elliptic, acute, palmately 7-8-nerved, denticulate, an- 
gulate-dentate or almost evenly rounded, cordate, 7-20 cm. long, thin, soon gla- 
brous; peduncles exceeding the leaves, up to 22 cm. long; cymes bisexual, strongly 
unilateral and usually much longer than wide, few- to many-flowered; bracts de- 




B 



FIG. 26. Begonia Popenoei. A, Habit; X K- B, Staminate flower and bracts; 
X 1. C, Stamen; X 5. D, Pistillate flower and bracts; X 1. E, Fruit; X 1. 



ciduous, obovate, ciliate; staminate tepals 2, broadly ovate, 5-9 mm. long, white 
or pale pink; stamens about 15, filaments short, anthers oblong; pistillate flowers 
bracteolate; pistillate tepals 2, like the staminate; ovary 3-celled, placentae bifid, 
ovuliferous on all sides, styles short-connate, stigmas lunate-capitate; capsule sub- 
erect, oblong-ellipsoid, 12 mm. long, wings unequal, the largest subtriangular and 
as high as wide. 

Begonia Popenoei Standl. Field Mus. Bot. 8: 142. 1930. 
Rocky slopes, alt. 1-100 meters. Izabal. Honduras. Figure 26. 

Herbaceous; rhizome creeping, rooting at the nodes, 5 mm. thick, soon gla- 
brous, red, internodes 1-7 cm. long; stipules lance-oblong, setose-mucronate, entire, 
20-25 mm. long, thin, brown, sparsely hirsute; petioles erect, slender, 15-30 cm. 
long, red, very densely hirsute with spreading to reflexed slender pinkish trichomes 
up to 2 mm. or more long; leaf -blades oblique, broadly ovate, acuminate, cordate 



180 



FIELDIANA: BOTANY, VOLUME 24 



at base with a deep open sinus, palmately 7-8-nerved, 10-23 cm. long, 6.5-17 cm. 
wide, not lobed but the teeth at the ends of the nerves sometimes slightly enlarged, 
ciliate-serrulate, green and glabrous above, paler beneath with hirsute nerves; 
peduncles to 36 cm. long, 5-6 mm. thick, red, soon glabrous; cymes few-flowered, 
about 4-12 cm. wide, with sparsely hirsute branches; bracts deciduous, suborbic- 
ular, 3 mm. long, entire; pedicels slender, 1-2 cm. long; staminate flowers develop- 
ing before the pistillate; staminate tepals 4, the outer suborbicular, 6-10 mm. long, 
rose-pink, the inner smaller, paler, narrowly obovate; stamens numerous, filaments 




FIG. 27. Begonia pustulata. A, Habit; X M- B, Upper leaf-surface; greatly 
enlarged. C, Staminate flower and bracts; X 1. D, Stamen; X 5. E, Pistillate 
flower and bracts; XI. F, Style; X 5. G, Fruit; X 1. 



short, anthers oblong with the connective enlarged at apex into a broad, rounded 
appendage; pistillate tepals 2, suborbicular, fleshy, pilose; styles deciduous; cap- 
sule reflexed, obovate, the largest wing mainly basal, suboblong, obtuse, 10-14 mm. 
wide, the others 3-4 mm. wide; seeds ellipsoid, blunt. 

Begonia pustulata Liebm. Vid. Medd. Kjoebenhavn 1852: 6. 
1853. 

On limestone rocks, 350-500 meters; Alta Verapaz. Southern 
Mexico. Figure 27. 

Herbaceous; rhizome creeping, rooting at the nodes, 3-5 mm. thick, pilose or 
becoming glabrous, internodes varying from very short to 12 cm. long (probably 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 181 

in response to habitat); stipules ovate, entire, setose at apex, 12-16 mm. long, 
thin, brown, pilose; petioles erect, slender, 8-20 cm. long, densely hirsute; leaf- 
blades oblique, broadly ovate, acuminate, cordate at base, palmately 8-9-nerved, 
not lobed, 7-15 cm. long, 5-11 cm. wide, ciliate-serrulate, bullate, hispid, green or 
with gray streaks along the major veins above; peduncles 7-26 cm. long, slender, 
pilose or becoming glabrous; cymes few-flowered, bracts deciduous, elliptic, setose- 
apiculate, entire, 5-12 mm. long; pedicels pubescent; staminate tepals 4, entire, 
the outer suborbicular, pubescent or glabrous, roseate, the inner narrowly obovate; 
stamens numerous, anthers oblong, about equaling the filaments, connective 
slightly produced, obtuse; pistillate bracteoles absent; pistillate tepals 3, the outer 
like the staminate, the inner one smaller; ovary 2-3-celled, placentae bilamellate, 
styles deciduous, often irregular, barely lobed at apex; capsule reflexed, ellipsoid, 
the largest wing mainly basal, ovate or suboblong, obtuse, 9-12 mm. wide, the 
others small, lunate, seeds ellipsoid, blunt. 

Begonia Sartorii Liebm. Vid. Medd. Kjoebenhavn 1852: 14. 
1853. B. lobulata A. DC. in DC. Prodr. 15, pt. 1: 339. 1864. B. co- 
bana C. DC. Bull. Herb. Boiss. II. 8: 322. 1908. 

Damp forest, 1,150-1,650 meters; Alta Verapaz; Baja Verapaz; 
Zacapa; Chiquimula; Santa Rosa; Huehuetenango. Southern Mexico. 
Figure 28. 

Herbaceous, 1-2.5 m. tall; stem erect, simple, over 1 cm. thick, green, rufous- 
pilose at least when young, internodes 5-6 cm. long; stipules ovate-oblong, entire, 
brown, membranaceous, 2-3 cm. long; petioles 8-16 cm. long; leaf-blades reniform- 
ovate, cordate at base, short-acuminate at apex, palmate-nerved, 9-14 cm. long, 
12-20 cm. wide, coarsely and shallowly few-lobed, the margin undulate and ciliate, 
membranaceous, glabrous above, densely rufous-pilose beneath when young; cymes 
axillary on stout peduncles 14-21 cm. long, bisexual, many-flowered, hemispherical 
or subumbellate, 11-12 cm. in diameter, lax; bracts like the stipules but small, 
early deciduous; staminate tepals 2, suborbicular, 8 mm. long, entire, white or 
pale pink; stamens free, about 18, half as long as the tepals, anthers linear-obovoid, 
more than twice as long as the filaments, connective produced into a broad, 
rounded projection; pistillate tepals 2, like the staminate but smaller; styles 3, 
short-connate, truncate, not lobed; capsule ebracteate, elliptic, 10 mm. long, 
largest wing 11 mm. wide, transversely ovate, obtuse, placentae bilamellate, ovu- 
liferous throughout. 

Begonia sciadiophora Smith & Schubert, Contr. Gray Herb. 
161:28, t. 3. 1946. 

Dense wet limestone forest, ca. 900 meters; Alta Verapaz (type 
and cotype from near Chirriacte", on the Pete"n highway, Standley 
91967 and 91953). 

Herbaceous; rhizome slender, branching, densely rufous-hirsute, internodes 
evident, up to 3 cm. long; stipules lanceolate, acuminate, with a long soft bristle 
at apex, to 12 mm. long, membranaceous, ferrugineous, subglabrous; petioles very 
slender, erect, 4-14 cm. long, densely rufous-hirsute; leaf-blades peltate, 7-nerved, 



182 



FIELDIANA: BOTANY, VOLUME 24 



broadly ovate, abruptly acuminate, broadly rounded at base, 5-6.5 cm. long, 
4-5 cm. wide, entire and unlobed, glabrous above, hirsute beneath especially on 
the nerves; peduncles about equaling the leaves, sparsely hirsute; inflorescence 
laxly cymose, few-flowered, sparsely hirsute; bracts somewhat persistent, lanceo- 
late, entire, membranaceous, brown, the lowest 10 mm. long; pedicels 6-8 mm. 
long; staminate tepals 4, entire, obtuse, the outer narrowly elliptic, 5.5 mm. long, 
one red, the other white, the inner 3 mm. long, white; stamens few, anthers oblong, 
about equaling the filaments; pistillate tepals 5, unequal, elliptic, obtuse, entire; 
ovary 3-celled, placentae bilamellate; capsule decurved, subellipsoid, the largest 
wing ovate, obtuse, 14 mm. long, 9 mm. wide, the others narrowly lunate. 




FIG. 28. Begonia Sartorii, A, Branch; X % B, Staminate flower; X 1. 
C, Stamen; X 5. D, Pistillate flower; X 1. E, Style; X 5. 



Begonia stigmosa Lindl. Bot. Reg. 31: misc. 32. 1845; Smith & 
Schubert, Ann. Mo. Bot. Gard. 45: 50, /. 15. 1958. B. squarrosa 
sensu Seemann, Bot. Herald 128. 1853, nomen, not Liebmann. 

Wet forests and thickets, 650-2,000 meters; Chimaltenango; 
Suchitepe'quez; Quezaltenango; San Marcos. Southern Mexico to 
Colombia. 

Herbaceous; rhizome repent, to 12 cm. long, 1 cm. thick, coarsely lepidote, 
internodes very short; stipules tardily deciduous, lanceolate, pilose, membrana- 
ceous, 15-20 mm. long; petioles erect, to 38 cm. long, 4-10 mm. thick, covered 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 183 

with spreading to reflexed flat narrowly triangular lacerate pink scales (with some 
trichome-like divisions) 3-5 mm. long; leaf-blades oblique, very broadly ovate or 
suborbicular, abruptly acuminate and usually with a second cusp more nearly 
opposite the petiole, cordate at base, 15-30 cm. long, scarcely if at all lobed, ciliate- 
denticulate, thin, glabrous above or with a few small white trichomes, the nerves 
beneath bearing pink trichomes; peduncle 2-5 dm. long, soon glabrous, 7 mm. 
thick; cymes bisexual, nearly regular, few- to many-flowered, 7-22 cm. long, lax, 
glabrous; bracts quickly deciduous, oblong and acute (A. DC.!); pedicels slender, 
7-23 mm. long; flowers white or pink; staminate tepals 2, suborbicular, cordate at 
base, 10-15 mm. long; stamens free, very numerous, filaments much longer than 
the broadly oblong anthers, connective apiculate-produced; pistillate flowers 
ebracteolate; tepals 2, like the staminate or rarely a smaller one inside the others; 
ovary 3-celled, placentae bifid, ovuliferous on all sides, styles much connate at 
base, bifid, the stigmatic tissue linear, spiral, continuous; capsule erect, broadly 
ovoid, 6-8 mm. long, wings very unequal, two forming marginal bands, the third 
ovate, oblong or subdolabriform, 10-15 mm. wide. 

Begonia strigillosa A. Dietr. Allg. Gartenz. 19: 330. 1851; 
Smith & Schubert, Ann. Mo. Bot. Card. 45: 49, /. 15. 1958. Gireou- 
dia strigillosa Klotzsch, Monatsber. Akad. Berlin 125. 1854. Begonia 
daedalia Lem. 111. Hortic. 7: misc. 54, 8, t. 269. 1861. B. barbana 
C. DC. Bull. Soc. Bot. Belg. 35, pt. 1: 261. 1896. Flor Colorado (fide 
Steyermark). 

Epiphytic to terrestrial, damp forests, 1,260-2,700 meters; Es- 
cuintla; Guatemala; Chimaltenango; Huehuetenango; Quezalte- 
nango; San Marcos. Costa Rica. 

Herbaceous; rhizome varying from short and compact with very short inter- 
nodes and imbricate stipules to slender and branching with distinct internodes; 
stipules lanceolate with setiferous apex and often fimbriate keel, persistent; petiole 
to 18 cm. long, often spotted, bearing numerous reflexed fimbriate scales; leaf- 
blades oblique, broadly ovate, acuminate, 7-17 cm. long, entire or slightly serrate, 
rounded to angulate-dentate, glabrous above, sparsely long-pilose below especially 
on the nerves, often spotted or mottled; peduncle usually exceeding the leaves, 
sparsely pilose, often purple-spotted; cyme 2-4-branched, mostly glabrous, sub- 
symmetrical, diffuse; bracts deciduous, elliptic, entire, mucronulate; staminate 
tepals 2, suborbicular to obovate, 6-9 mm. long; stamens few, the oblong obtuse 
anthers longer than the filaments; pistillate flowers ebracteolate; tepals 2, like 
the staminate, ovary 3-celled, placentae bifid, ovuliferous on all sides, styles per- 
sistent, connate at base, somewhat bifid, stigmas linear; capsule suberect, ovoid or 
elliptic-ovoid, locules to 10 mm. long, the two larger wings subtriangular or sub- 
obtuse to acuminate. 

Begonia tinctoria Smith & Schubert, Contr. Gray Herb. 127: 
29, t. 2, f. 22-21*. 1939. (Type and only known collection from 
Suchitepe"quez, Skutch 1556.) 

About 3 dm. high; rhizome 5 mm. thick; stipules ovate, about 1 cm. long, 
attenuate into a setose tip, 5 mm. long; petioles 4-14 cm. long, bearing trichomes 



184 



FIELDIANA: BOTANY, VOLUME 24 



like those on the leaf-margins; leaf-blades oblique or transverse, elliptic, acuminate, 
5-nerved, serrate-ciliate with trichomes 3 mm. long, 5.5-8.5 cm. long, 3-4 cm. 
wide, glabrous above, pustulate beneath; peduncle exceeding the leaves, sparsely 
pilose; inflorescence cymose, bisexual, bracts quickly deciduous, unknown; flowers 
bright red; staminate tepals 2, obovate, 9-10 mm. long; anthers oblong, longer 





V 



B 



FIG. 29. Begonia trichosepala. A, Habit; X 14. B, Staminate flower; X 1. 
C, Stamen; X 5. D, Pistillate flower; X 1. E, Style; X 5. 



than the filaments; pistillate tepals 2, 7-9 mm. long; pedicels 10-35 mm. long; 
ovary glabrous, styles 3, bifid; capsule 14 mm. long including the 3 subequal 
rounded wings. 

Begonia trichosepala C. DC. Bot. Gaz. 20: 542. 1895 (type 
from Alta Verapaz, von Tuerckheim 225). B. erythrotricha C. DC. 
Bull. Herb. Boiss. II. 8: 321. 1908 (type from Alta Verapaz, von 
Tuerckheim 1 1 1336). 

Rich, wet forests, 300-1,400 meters; Alta Verapaz; Huehuete- 
nango. Endemic. Figure 29. 

Herbaceous; rhizome creeping, rooting at the nodes, 2-5 mm. thick, usually 
soon glabrous, internodes 1-8 cm. long; stipules lanceolate, acuminate, 10-18 mm. 
long, lacerate-denticulate; petioles erect, 4-17 cm. long, sparsely pilose, becoming 
glabrous; leaf -blades straight, symmetric, ovate, acuminate, shallowly cordate at 
base, 7-13 cm. long, 4-6 cm. wide, penninerved, large teeth at the ends of the prin- 



STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 185 

cipal nerves, margin denticulate between, soon glabrous above, beneath sparsely 
pilose at least on the nerves; peduncles axillary, mostly shorter than the leaves, 
soon glabrous; cymes few-flowered, bisexual; bracts deciduous, elliptic; pedicels 
pilose; staminate tepals 4, entire, white, the outer broadly obovate, to 13 mm. 
long and 11 mm. wide, more or less pilose, the inner narrowly spatulate; stamens 
on a torus or short column, numerous, anthers oblong, mostly exceeding the fila- 
ments, connective produced, obtuse; pistillate bracteoles lacking; pistillate tepals 3, 
the 2 outer large, reniform, the inner much smaller; styles deciduous, short-lobed, 
ovary 3-celled, placentae bilamellate and simple; capsule reflexed, the largest wing 
mainly basal, suboblong, obtuse, the others lunate, much smaller. 



Publication 936 



UNIVERSITY OF ILLINOIS-URBANA